Effects of storage on germination ofDioscorea composita (Dioscoreaceae) seeds

Effects of storage were tested on germination ofDioscorea composita (Dioscoreaceae) seeds. Freshly collected seeds and seeds stored at 25°C in paper bags from 1 to 11 mo or for 4 and 5 yr were used in most of the experiments. Seeds were tested for germination at 20, 25, 30, 35, 25–20, and 25–35°C in white light and in darkness. Initiation of germination was delayed in freshly harvested seeds, and dormancy was reduced in seeds stored for about 9 mo. Viability of the seeds decreased after 4 and 5 yr of storage.     

Effects of seed maturation time and dry storage on light and temperature requirements during germination in invasive Prosopis juliflora

The effects of time of seed maturation and dry seed storage and of light and temperature requirements during seed incubation on final germination percentage and germination rate were assessed for the invasive shrub Prosopis juliflora (Sw.) D.C., grown under desert environmental conditions of the United Arab Emirates (UAE). Seeds were collected from Fujira on the northern coast of the UAE at different times during the growing seasons (autumn, winter and spring) and were germinated immediately and after 8 months of dry storage under room temperature (20±3 °C). Seeds were germinated at three temperatures (15, 25 and 40 °C) in both continuous light and darkness. The results showed significant effects for time of seed collection, seed storage, light and temperature of seed incubation and many of their interactions on both germination percentage and rate. Fresh seeds matured during autumn and winter germinated significantly greater at 40 °C and in light than at lower temperatures and in dark. Storage significantly increased germination percentage and rate; the increase was greater for seeds matured during winter than for seeds matured during spring. This indicates that dormancy breakage was greater in seeds of winter than seeds of spring. The need for high temperature to achieve greater germination was significantly reduced after seed storage, especially for seeds matured in autumn and winter.     

Dormancy studies in black gram (Phaseolus mnngo L.)

Mechanical scarification of seeds ofPhaseolus mungo L. resulted in 100 per cent gormination in comparison to 90 per cent dormant control. Seeds treated with concentrated sulphuric acid for 8 min led to 85 per cent germination. Heat treatment at 70 °C for 24 h exhibited 75 per cent gormination, while boiled water treatment for 7 min caused 65 per cent germination.     

After-Ripening in Festuca idahoensis Seeds: Adaptive Dormancy and Implications for Restoration

Festuca idahoensis (Idaho fescue) was a common native perennial bunchgrass in the sagebrush steppe of the western United States until the introductions of domestic livestock and alien plants. Restoration of Idaho fescue to degraded sites will likely involve reseeding, and one of the factors affecting reseeding success is germinability of the seeds employed. We investigated effects of after-ripening and storage temperature on germinability of Idaho fescue seeds collected from a central Oregon site. Six months of after-ripening were required before maximum germination was obtained. Storage of dry seeds at either room temperature (20°C) or at cooler, alternating temperatures (5/15°C) did not alter the rate at which dormancy was lost. Storage at the warmer temperature promoted rapid germination in seeds that had broken dormancy. Seed longevity varied greatly from year to year. Seeds produced in a very dry year had poorer germination and shorter longevity than seeds produced during a year with near normal precipitation. Because seed dispersal occurs in late July and early August for Idaho fescue in central Oregon, a six-month after-ripening requirement ensures that the greatest potential germination coincides with the spring period most likely to provide sufficient moisture for seedling establishment.     

Germination of stored and scarified seeds of Passiflora caerulea L. (Passifloraceae)

Abstract

This work evaluates the influence of storage and scarification in the germination of Passiflora caerulea L., in order to acquire the necessary knowledge to cultivate this economically important species. Seeds stored one month under different conditions (no storage, fermentation, desiccation) were submitted to various pre‐treatments (aril removal, mechanical and chemical scarification or combinations of these). Emergence was recorded periodically for 11 months in a greenhouse. Germination time, germination percentage and germination speed were calculated. Histochemical tests were applied to seeds maintained under the three storage conditions, for observing possible changes in the integument and storage tissue. Viability was maintained at least for the storage period tested, as germination occurred after that time. Because the seeds stored under desiccation germinated, the species is orthodox. Chemical scarification prevented germination in most cases. Although germination levels were low, they were higher in stored seeds (fermented and desiccated) than in fresh ones. Fermented seeds (which exhibited no storage tissue and less lignin in the integument) devoid of the aril germinated earlier, faster and in greater quantity. The type of dormancy of this species is discussed.     

Strict requirement of fluctuating temperatures as a reliable gap signal in <Emphasis Type="Italic">Picris hieracioides</Emphasis> var. <Emphasis Type="Italic">japonica</Emphasis> seed germination

Picris hieracioides var. japonica (Asteraceae), which grows in occasionally disturbed habitats such as riverbanks, is rarely observed under dense vegetation. We examined the effect of the experience and timing of receiving leaf-transmitted light in gap-detecting seed germination in this plant. Seeds under unfiltered light, which simulated the light conditions of seeds on the soil surface in a canopy gap, germinated at a constant temperature of 20°C. However, most seeds in darkness, which simulated the light conditions of seeds buried in the soil without receiving leaf-transmitted light, germinated under temperature fluctuations of over 4°C. Seeds in darkness after receiving leaf-transmitted light for 1 week, which simulated the light conditions of seeds buried in the soil after receiving leaf-transmitted light, germinated under temperature fluctuations of over 8°C. Finally, seeds under continuous leaf-transmitted light, which simulated the light conditions of seeds on the soil surface below preexisting vegetation, germinated under temperature fluctuations of over 12°C. Seeds that experience unfiltered light, which suggests that they are in a gap, should not delay germination. In contrast, seeds that have received leaf-transmitted light should delay germination until the vegetation above is removed. Seeds exposed to leaf-transmitted light required larger temperature fluctuations in darkness than did untreated seeds, and seeds under continuous leaf-transmitted light required the largest temperature fluctuations. The various germination reactions to each gap signal in P. hieracioides var. japonica seeds allow the more reliable detection of gaps for subsequent seedling establishment. The requirement for gap signals that created high precision of timing in the germination process results in the germination of this species only in gaps. Therefore, P. hieracioides var. japonica is rarely found under dense vegetation.     

Effect of in situ storage,light, and moisture on the germination of two wetland tropical trees

Seed germination was evaluated for Annona glabra L. and Pachira aquatica Aubl. in the wetland conditions of La Mancha, Veracruz, Mexico. These species have recalcitrant seeds and hydrochoral dispersal. Germination experiments were carried out under varying moisture (high, middle, and low) and light (below canopy and open sky) levels as well as after being stored in contrasting natural conditions. Seeds were stored both floating in water and buried in wetland soil for 15, 30, 60, and 90 days. P. aquatica seeds germinated faster in low and medium moisture, regardless of light intensity. After 45 days, for example, they exhibited 87–73% germination in medium-moisture/canopy and low-moisture/canopy treatments, respectively. In high moisture, seeds reached similar percentages after 60 days (80%). Storage by burial caused the death of seeds regardless the time they spent underground, while those stored in water germinated at a rate of over 90%. A. glabra seeds germinated better (98%) in low-moisture/sunny conditions. As to storage, they responded favorably to both burial and water techniques but germinated more readily in treatments that involved a long storage period. Evaluation of the germination behavior of A. glabra and P. aquatica seeds subject to varying in situ storage and germination conditions illustrates the response capacity of each species during early phases of development.     

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。     

Seed dormancy and germination of the rare,high elevation Balkan endemic <Emphasis Type="Italic">Cerastium dinaricum</Emphasis> (Caryophyllaceae)

Short or long-term ex situ conservation is becoming increasingly important in conservation of plants in today’s changing environments. One of the important steps in ex situ conservation is the collection and storage of seeds and the consequent establishment of seed germination protocols. Cerastium dinaricum (Caryophyllaceae) is an endemic, high elevation and rare species of European conservation concern. Because of its severely fragmented distribution along the Dinaric Alps, the populations are likely to undergo further shrinkage in the future, which addresses the need of a long-term effective conservation management. From the potential ex situ population management perspective, we focused our study on germination ecology of C. dinaricum. The study revealed that temperature considerably affected the germination of seeds, which germinate better at 20 °C rather than 10 °C. A period of cold-wet stratification also significantly improved the final germination percentage with more pronounced increase at 20 °C, while addition of GA3 increased the final germination percentage by breaking the dormancy of non-stratified seeds. Mechanical scarification did not improve germination; on the contrary, it resulted in the lowest germination success. Seeds grown in complete darkness germinated significantly better compared to control when they were exposed to cold-wet stratification. Contrary to previous studies on some alpine species, which germinate better when exposed to light, dark treatment resulted in the highest germination percentages with 70 and 90% germination success after 4 and 8 weeks of stratification, respectively.     

Seed Dormancy,Seedling Establishment and Dynamics of the Soil Seed Bank of Stipa bungeana (Poaceae) on the Loess Plateau of Northwestern China

Studying seed dormancy and its consequent effect can provide important information for vegetation restoration and management. The present study investigated seed dormancy, seedling emergence and seed survival in the soil seed bank of Stipa bungeana, a grass species used in restoration of degraded land on the Loess Plateau in northwest China. Dormancy of fresh seeds was determined by incubation of seeds over a range of temperatures in both light and dark. Seed germination was evaluated after mechanical removal of palea and lemma (hulls), chemical scarification and dry storage. Fresh and one-year-stored seeds were sown in the field, and seedling emergence was monitored weekly for 8 weeks. Furthermore, seeds were buried at different soil depths, and then retrieved every 1 or 2 months to determine seed dormancy and seed viability in the laboratory. Fresh seeds (caryopses enclosed by palea and lemma) had non-deep physiological dormancy. Removal of palea and lemma, chemical scarification, dry storage (afterripening), gibberellin (GA₃) and potassium nitrate (KNO₃) significantly improved germination. Dormancy was completely released by removal of the hulls, but seeds on which hulls were put back to their original position germinated to only 46%. Pretreatment of seeds with a 30% NaOH solution for 60 min increased germination from 25% to 82%. Speed of seedling emergence from fresh seeds was significantly lower than that of seeds stored for 1 year. However, final percentage of seedling emergence did not differ significantly for seeds sown at depths of 0 and 1 cm. Most fresh seeds of S. bungeana buried in the field in early July either had germinated or lost viability by September. All seeds buried at a depth of 5 cm had lost viability after 5 months, whereas 12% and 4% seeds of those sown on the soil surface were viable after 5 and 12 months, respectively.     

Dormancy,viability and germination of Magnolia lanuginosa (Wall.) Figlar & Noot. seeds: A threatened tree species of Northeast India

The population of Magnolia lanuginosa a rare tree species of northeastern India has declined drastically owing to habitat destruction, low natural regeneration and over harvesting for its multipurpose uses. The present study was carried out to understand the type of dormancy and analyse the effect of storage on viability and germination behaviour of M. lanuginosa under various physical and chemical treatments. Seeds subjected to physical treatments such as water (cold, hot, and boiling), acid (H₂SO₄) and manual scarification failed in breaking dormancy. Seeds treated with growth regulators (GA₃) had a significant effect on germination. It reduced the germination time and the shortest T₅₀ was observed in seeds treated with 2000 mg/l of GA₃ (non-scarified seeds) and 1000 mg/l of GA₃ (scarified seeds). The use of KNO₃ did not have any significant effect in breaking dormancy. However, the use of KNO₃ along with GA₃, increased the germination percentage. Seeds cold stratified (CS) for 60 days at 5 °C was effective in breaking dormancy and resulted in 84.26% germination. This indicates the prevalence of Type-1 Non deep physiological dormancy in M. lanuginosa seeds that requires a crucial CS period for proper embryo growth and development. The seeds stored in moist sand at 5 °C remained viable even after 120 days with 48.88% viability. The study would be helpful in devising seed germination protocols for mass production and reintroduction of the species into the wild.     

Drought tolerance and germination response to light and temperature for seeds of saline and non-saline habitats of the habitat-indifferent desert halophyte <Emphasis Type="Italic">Suaeda vermiculata</Emphasis>

Few plants are habitat-indifferent halophytes (i.e., grow well in both saline and non-saline soils). These plants offer a good opportunity to study drought and salinity tolerances during germination for seeds developed and matured in soils differ in salinity. Here, we assessed drought tolerance during germination, as simulated with PEG, and response of germination to light and temperature for Suaeda vermiculata, a habitat-indifferent shrub. Seeds matured in saline and non-saline soils were germinated in six PEG concentrations (0 to ? 1.0 MPa) and put in three incubators adjusted at different temperatures in both light and dark regimes. Drought tolerance was greater for seeds of the non-saline than those of saline soils, especially at higher temperatures. Seeds of the saline soils germinated in the lowest osmotic potentials (? 0.8 and ??1.0 MPa) only at lower temperatures, but seeds of the non-saline soils germinated to higher levels at all temperatures. Tolerances to drought and high temperatures were greater in light for seeds of saline soils, but in darkness for seeds of non-saline soils. Germination rate index did not differ significantly between seeds of the two soil types in higher osmotic potentials, but was significantly greater in seeds of non-saline at lower osmotic potentials. Most seeds that failed to germinate in the PEG concentrations recovered their germination when transferred to distilled water. Germination recovery levels and speeds increased with the decrease in osmotic potentials. Seeds of the saline soil postpone their germination until arrival of suitable temperatures and effective rainfalls that ensure seedling survival in salty habitats of the arid unpredictable deserts.     

Seed dormancy-breaking and germination requirements ofDrosera anglica,an insectivorous species of the Northern Hemisphere

Seeds of Drosera anglica collected in Sweden were dormant at maturity in late summer, and dormancy break occurred during cold stratification. Stratified seeds required light for germination, but light had to be given after temperatures were high enough to be favorable for germination. Seeds stratified in darkness at 5/1 °C and incubated in light at 12/12 h daily temperature regimes of 15/6, 20/10 and 25/15 °C germinated slower and to a significantly lower percentage at each temperature regime than those stratified in light and incubated in light. Length of the stratification period required before seeds would germinate to high percentages depended on (1) whether seeds were in light or in darkness during stratification and during the subsequent incubation period, and (2) the temperature regime during incubation. Seeds collected in 1999 germinated to 4, 24 and 92 % in light at 15/6, 20/10 and 25/15 °C, respectively, after 2 weeks of stratification in light. Seeds stratified in light for 18 weeks and incubated in light at 15/6, 20/10 and 25/15 °C germinated to 87, 95 and 100 %, respectively, while those stratified in darkness for 18 weeks and incubated in light germinated to 6, 82 and 91 %, respectively. Seeds collected from the same site in 1998 and 1999, stratified in light at 5/1 °C and incubated in light at 15/6 °C germinated to 22 and 87 %, respectively, indicating year-to-year variation in degree of dormancy. As dormancy break occurred, the minimum temperature for germination decreased. Thus, seed dormancy is broken in nature by cold stratification during winter, and by spring, seeds are capable of germinating at low habitat temperatures, if they are exposed to light.     

Variation in seed dormancy and germination among populations of Silybum marianum (Asteraceae)

Milk thistle (Silybum marianum) is a medicinal plant; however, lack of consistency in past dormancy studies has hindered propagation of this species from seeds. We tested the germination responses of freshly harvested and after-ripened (stored for 2 and 7 months; 25°C at 50% relative humidity) seeds from three populations (P1, P2 and P3) in Iran at varying constant or alternating temperatures, with or without GA₃ and in light and continuous darkness. No germination occurred in freshly harvested seeds incubated at any condition without GA₃ application, indicating that all the seeds were dormant. Seeds from P1 and P2, which developed under relatively dry, warm conditions, germinated over a wider range of temperatures after 2 months of dry storage, indicating type 6 of non-deep physiological dormancy (PD). Seeds from P3, which developed under relatively wet, cool conditions, incubated at constant temperatures (especially on GA₃), exhibited an increase in maximum temperature for germination, indicating type 1 of non-deep PD. Light improved germination of after-ripened seeds, and GA₃ application substituted for the light requirement for germination. This is the first report that environmental conditions during seed development may be correlated with differences in the type of non-deep PD. We conclude that milk thistle seeds are positively photoblastic and photodormant and the germination responses of after-ripened seeds from different populations are different under darkness. Therefore, the impacts of genetic differences and maternal effects on the induction of dormancy during seed development should be considered in attempts to domesticate this medicinal plant.     

Differential germination behaviour in seeds of the Indian arid zone

Seed germination studies of a number of Indian arid zone species were conducted and the seeds were classified into groups on the basis of germination behaviour. Results of seed germination were also compared with those of old seed collections. Most of the seeds studied showed some form of dormancy, the hard seed coatedness being the most common. InBorreria articularis, germination inhibitors were found. Seeds of different years required different duration of acid scarification for maximum germination and these differences have been attributed to rainfall and other such environmental conditions in those years.     

Asymbiotic seed germination and in vitro seedling development of the threatened orchid Hoffmannseggella cinnabarina

Hoffmannseggella cinnabarina has not been found in the wild for the last 70?yr in the State of S?o Paulo and, therefore, wild populations of this native orchid are thought to be extinct. This investigation studied seed storage at a low temperature, in vitro germination, and seedling development of H. cinnabarina in order to establish an optimized protocol for propagation, and thus assure species conservation. Seeds of different ages were incubated on Knudson C (KC), Murashige and Skoog, and Vacin and Went media with or without 1???M of N⁶-benzyladenine (BA) and exposed to either 12 or 16?h of light (30???mol?m^?2?s^?1 at 26?±?2°C). Seed surface sterilization was deleterious to 3-mo-old seeds and severely reduced the viability of the 4-mo-old seeds. More mature seeds were not affected by the sterilization procedure. In general, the germination of 4-, 5-, 6-, and 7-mo-old seeds increased when BA was added to the culture medium especially under 16?h of light. Germination rates were highest with 8- and 9-mo-old seeds, and application of BA failed to enhance germination rates further. Developmental studies revealed that this cytokinin reduced seed and protocorm mortality rates; however, protocorm development was negatively affected in its presence. Seedling development was more pronounced when KC medium with 16?h of light was used. Long-term seed storage at 4°C did not provide promising results. The protocol described in this study proved to be efficient and relevant to in vitro seed germination and initial development of H. cinnabarina, and thus will contribute to conservation of this orchid species.     

Effects of long-term storage on phytochrome-mediated germination in lettuce seeds

The effects of long-term seed storage on the physiological properties of phytochrome-mediated germination including water uptake, the temperature and light flunnce dependencies of germination and dark germination were studied. The fluenceresponse relationships of the brief irradiation with monochromatic red (660 nm, 7.5 W m⁻²) and far-red (750 nm, 6.6 W m⁻²) light at various times after sowing were also studied. The samples used consisted of three lots of seeds ofLactuca sativa L. cv. MSU-16, which had been harvested in 1976, 1979 and 1985 and stored dry for 9, 6 and 0 years, respectively, in darkness at 23±2 C until the experiments were carried out in July–August, 1985. Seeds with the longer storage periods showed the higher ability to germinate in both continuous darkness and continuous white fluorescent light at 20–30 C. In the seeds stored for 6 or 9 years, red light irradiation for 20 sec given at 15 min or more after sowing at 25 C induced as high a percent germination (85–95%) as those under continuous white fluorescent light. In the freshly harvested seeds, however, germination under continuous white fluorescent light (46%) was considerably lower than the germination induced by the red pulse (97%). Germination of the seeds decreased when the intervals between sowing and a far-red irradiation for 20 sec increased up to 100 min (or 30 min in the freshly harvested seeds). The far-red pulse given later than 100 min (or 6 hr in the freshly harvested seeds) after sowing resulted in an increased germination up to the dark-germination levels with increasing intervals between sowing and the pulse irradiation. Before or at 3 min after sowing, the seeds stored for 6 or 9 years were responsive to the far-red pulse although they were not or hardly responsive to the red pulse, while the freshly harvested seeds were responsive to both the far-red and the red pulses. These data indicate that normal functions of phytochrome completely survived in the dry seeds during storage at 25 C for as long as 6 or 9 years and that these functions are restored into full operation by means of imbibition. The differences in the dependence of germination on the time and fluence of a single pulse of red or far-red light seems to be related to the smaller water content throughout the imbibition in the seeds with the longer storage periods. The greater ability to germinate in the dark indicates the greater amounts of P_FR or the greater responsivity to P_FR, in the seeds with the longer storage periods.     

Breaking Setaria parviflora seed dormancy by nitrates and light is part of a mechanism that detects a drawdown period after flooding

We explored the hypothesis that, in flood-prone habitats, nitrates can signal to seeds that a drawdown period has begun. To investigate this issue, Setaria parviflora (Poir.) Kerguélen seeds were buried in a never-flooded upland and a nearby, flood-prone lowland grassland. Seeds were exhumed during the flooding period. Additionally, grassland mesocosms with buried S. parviflora seeds were flooded during 20 d (controls drained). After both field and mesocosm pretreatments germination was assayed in laboratory at 25 °C in a medium with or without nitrates, under red light pulses or in darkness. Seeds exhumed from the never-flooded upland showed no specific requirements to germinate. In contrast, seeds exhumed from the flooded lowland germinated ca. 65% when nitrates were combined with red light pulses, significantly higher than in the rest of the treatments. Seeds exhumed from drained mesocosms germinated equally in all treatments. However, in the seeds exhumed from the flooded mesocosms, nitrates increased germination by more than 20% compared with seeds imbibed in water. Seeds germinated ca. 85% when nitrates were combined with red light pulses, significantly higher than in the other treatments. We can conclude that after flooding, S. parviflora seeds require nitrate and light to germinate. Therefore, a large fraction of seeds do not germinate unless nitrates are combined with light, indicating a drawdown period after floods and vegetation gaps.     

Germination response and viability of an endangered tropical conifer Widdringtonia whytei seeds to temperature and light

The tropical conifer Widdringtonia whytei Rendle is an endangered species endemic to Mulanje Mountain in Malawi. A study was conducted for the first time under controlled conditions in order to assess the effects of temperature and light on germination and viability of W. whytei seeds. Seeds incubated at a constant temperature of 20 °C attained the highest cumulative germination percentage (100%) followed by 87% germination under fluctuating temperatures of 15 °C night/25 °C day. No seed germination occurred at temperatures below 15 °C. Seeds that failed to germinate at temperatures below 15 °C showed the highest (> 90%) viability compared to the seeds incubated at 25 °C (60%). Across temperature regimes, germination was significantly higher under light (44.7%) than dark (35.6%) conditions. It is concluded that temperature is one of the critical factors for germination of W. whytei seed. The ability of W. whytei seeds to germinate both in light and darkness implies that the species would unlikely form a persistent soil seed bank, an attribute which is common in species that survive in habitats frequently disturbed by fires.     

The ecology of germination and reproduction of less frequent and vanishing species of the Czechoslovak flora I.Alyssum saxatile L.

Lhotská M. (1988): The ecology of germination and reproduction of less frequent and vanishing species of the Czechoslovak flora I.Alyssum saxatile L.—Folia Geobot. Phytotax., Praha, 23: 321–324.—The conditions of germination of both fresh and stratified seeds ofAlyssum saxatile were investigated in the laboratory at 13 various temperature values. The temperature of 15/25°C was found to be optimum. In stratified seeds germination was accelerated, and the range of range of temperatures suitable for germination became wider. Seeds sown out after ripening sprouted in the first half of June, and by autumn they developed rather robust rosettes.