Temperature Response Pattern during Afterripening of Achenes of the Winter Annual Krigia oppositifolia (Asteraceae)

Abstract Freshly-matured achenes of Krigia oppositifolia Raf. were buried in soil at near-natural temperatures for 0–35 months and then exhumed and tested in light and darkness at (12/12 hr) daily thermoperiods of 15/6, 20/10, 25/15, 30/15 and 35/20°C. Achenes required light for germination and exhibited an annual dormancy/nondormancy cycle, being dormant in spring and nondormant in autumn. High summer temperatures (30/15, 35/20°C) fully promoted afterripening, whereas low temperatures (5, 15/6°C) prevented it. As buried seeds came out of dormancy in summer, they first germinated at medium temperatures (20/10, 25/15°C), but with additional afterripening the maximum and minimum temperatures for germination increased and decreased, respectively. Thus, during afterripening, achenes exhibit type 3 temperature responses, which otherwise are known only in two perennial Asteraceae and one perennial Liliaceae. The physiological responses of achenes of K. oppositifolia are unlike those of most winter annuals, which have type 1 responses—i.e., the maximum temperature for germination increases during afterripening. Also, they are unlike the majority of Asteraceae, which have type 2 responses—i.e., the minimum temperature for germination decreases during afterripening. Type 1 responses, typical of most winter annuals, have yet to be reported in the Asteraceae.     

Effects of light, temperature and water stress on germination of Artemisia sphaerocephala

Artemisia sphaerocephala is widely used for vegetation rehabilitation, but its germination is very low after air seeding of achenes. We explored effects of light, temperature and water stress on germination. Results show that both final percent germination and germination rate were increased by temperature increment, with the highest values occurring at 15: 25°C (night: day) in dark and 20: 30°C under light. Light inhibited germination, especially at lower alternating temperatures (5: 15°C and 10: 20°C). The alternating temperature window for germination was slightly narrower under light than in dark, and germination was slower under light than in dark across the temperature range. Achenes incubated in the dark and at constant temperatures had over 80% germination at 10 to 25°C, with an optimum at 20°C. Under dark and 25μmol m^‐2 s^‐1 light flux density at 10: 20°C, final percent germination was over 94%, but if the light flux density was increased to 100 and 400 μmol m^‐2 s^‐1, final percent germination was significantly lower (64% and 38% respectively). However, achenes could keep their germination capacity for a long time (over 50 days) and germinate well after going back to the dark. Germination was also lower under water stress and few achenes germinated at ‐1.4 MPa. This was more pronounced at high and low temperatures. Given these findings and the prevailing climatic characteristics, the most suitable time for air seeding of achenes may be mid‐May.     

Morphophysiological dormancy in seeds of the ANA grade angiosperm Schisandra arisanensis (Schisandraceae)

We determined the kind of seed dormancy in Schisandra arisanensis, an ANA grade ([A]mborellales [N]ymphaeales [A]ustrobaileyales) angiosperm with medicinal value. Seeds have small underdeveloped embryos, and following seed maturity their length increased approximately 360% before radicle emergence. Germination was delayed 6–8 weeks, and the percentage and rate were much higher at 15/6, 20/10 and 25/15°C than at 30/20°C. For seeds incubated at 5/5°C (8 weeks) → 15/6°C (4 weeks) → 20/10°C (8 weeks) → 25/15°C (12 weeks) → 20/10°C (5 weeks), embryos grew at 15/6°C → 20/10°C, and almost all seeds that germinated (89%) did so at 20/10°C → 25/15°C. When seeds were incubated in a complementary temperature sequence, 25/15°C (12 weeks) → 20/10°C (8 weeks) → 15/6°C (4 weeks) → 5/5°C (9 weeks) → 15/6°C (4 weeks), embryos grew at 25/15°C → 20/10°C. Nearly all seeds that germinated (93%) did so at 25/15°C → 20/10°C and at 15/6°C following 9 weeks at 5/5°C. Based on the temperature requirements for embryo growth and seed germination, seeds of this species have non‐deep simple morphophysiological dormancy (C_1bB).     

Seed germination traits of Ailanthus altissima,Phytolacca americana and Robinia pseudoacacia in response to different thermal and light requirements

Invasion of alien plant species (IAS) represents a serious environmental problem, particularly in Europe, where it mainly pertains to urban areas. Seed germination traits contribute to clarification of invasion dynamics. The objective of this research was to analyze how different light conditions (i.e., 12-hr light/12-hr darkness and continuous darkness) and temperature regimes (i.e., 15/6°C, 20/10°C and 30/20°C) trigger seed germination of Ailanthus altissima (AA), Phytolacca americana (PA) and Robinia pseudoacacia (RP). The relationship between seed germination and seed morphometric traits was also analyzed. Our findings highlight that temperature rather than light was the main environmental factor affecting germination. RP germinated at all tested temperatures, whereas at 15/6°C seeds of AA and PA showed physiological dormancy. RP had a higher germination capacity at a lower temperature, unlike AA and PA, which performed better at the highest temperatures. Light had a minor role in seed germination of the three species. Light promoted germination only for seeds of PA, and final germination percentage was 1.5-fold higher in light than in continuous darkness. Seed morphometric traits (thickness [T], area [A] and volume [V]) had a significant role in explaining germination trait variations. The results highlight the importance of increasing our knowledge on seed germination requirements to predict future invasiveness trends. The increase in global temperature could further advantage AA and PA in terms of germinated seeds, as well as RP by enhancing the germination velocity, therefore compensating for a lower germination percentage of this species at the highest temperatures.     

Dispersion,germination and growth of seedlings ofSagittaria lancifolia L.

Sagittaria lancifolia L. a perennial helophyte growing in shallow waters has asexual reproduction by rhizomes and sexual reproduction by achenes. We studied the morphological characteristics of achenes related to its dispersion, some factors affecting its germination and the development stages and growth of seedlings. This species produces many achenes, and disperses through epizoochory and hydrochory. Achenes germinate on light only. 100% germination was obtained at 25°C. Under conditions of inundation the germination period decreases, and the percentage of germination in creases when compared to achenes with or without imbibition. Achenes in anaerobic conditions have reduced germination. The growth of seedlings ofS. lancifolia was affected by the temperature: each organ of the seedlings grows differently in dependence upon this factor. For example, roots grow faster during the early stages of the seedlings.     

Germination of Tagetes minuta L. I. Temperature Effects

Initial studies have indicated that Tagetes minuta achenes haveboth a temperature and a light requirement for germination.Temperatures tested were 10, 20, 25, 30 and 35 °C. Germinationwas optimal at 25 °C under white light conditions. Underthese conditions 100 per cent of achenes germinated within 7days of imbibition. There was no germination at 10 or 35 °Ceither in the light or in the dark. Achenes imbibed and incubatedat 35 °C for 4 days showed no visible signs of germinationbut on transfer to 25 °C, 100 per cent of these achenesgerminated within 24 h. Furthermore, achenes given this hightemperature (35 °C) treatment could be dried at 25 °C,re-imbibed at 25 °C and again 100 per cent of achenes germinatedwithin 24 h of re-imbibition. This rapid germination responsefollowing removal from the high temperature regime could alsobe induced by transfer to temperatures of 20 °C or 20 °C(16 h) alternating with 10 °C (8 h). Tagetes minuta L., weed seeds, germination, temperature, light     

Physiological dormancy in seeds of tropical montane woody species in Hawai`i

Worldwide, there is relatively little information on seed dormancy and germination of tropical montane species. Our aim was to help fill this knowledge gap by conducting seed dormancy/germination studies on woody species from this vegetation zone in Hawai`i. All species had water-permeable seeds with a fully developed embryo. Seeds of 29 species (23 genera) were incubated in light/dark at 15/6, 20/10 and 25/15°C and germination monitored at 2-week intervals for 16–128 weeks. Seeds of Chenopodium oahuense, Dubautia menziesii and Silene lanceolata were non-dormant (ND) and those of 26 other species had physiological dormancy (PD); 10 of the 26 species had conditional PD. The optimum germination temperature regime(s) was (were) 25/15°C, 17 species; 25/10 and 20/10°C, 2; 20/10°C, 6; 20/10 and 15/6°C, 2; and 15/6°C, 2. Worldwide, PD in the woody genera included in our study is more common than ND. In addition to its contribution to the world biogeography of seed dormancy/germination, this study will be useful to conservation biologists who need to germinate seeds of tropical montane species.     

Physiological dormancy in seeds of two endemic species of Begonia from Yunnan Province,China: diagnosis and classification

With the purpose of assessing the status of dormancy in seeds of two Begonia species (Begonia lithophila and Begonia guishanensis), freshly matured seeds were given gibberellic acid and moist chilling and allowed to dry after ripening. The seeds were then germinated on media with or without KNO₃ at 15, 20, 25, 30 and 18/25°C. All three treatments significantly increased germination percentages. Examination by X‐ray revealed that seeds of both species have a fully developed embryo and thus have no morphological component of dormancy; seeds readily imbibed water and KNO₃ solution. Therefore, we conclude that seeds of the two Begonia species have non‐deep physiological dormancy. Although KNO₃ significantly increased germination in both species, alternating temperatures did not, suggesting that the most favorable microhabitat for germination is small‐scale disturbances under the forest canopy.     

Biological characteristics of an invasive south African species

Senecio inaequidens, a south African species that has recently invaded parts of Europe, especially human disturbed habitats, was examined under laboratory and glasshouse conditions to assess the germination of achenes, soil seed banks, growth of seedlings and its breeding system. These observations allowed identification of ecological characteristics that may contribute to the invasiveness of the species. Achenes derived from three populations of S. inaequidens were germinated over a wide temperature range (from 14/6 °C day/night temperatures to a constant temperature of 30 °C). They exhibited a rapid achene germination especially when temperatures reached 20 °C. Germination of seeds from soil samples also exhibited a prolonged germination over time suggesting therefore different types of dormancy. Growth response was studied along a nitrogen gradient from 0.33 to 16.0 mmol l^–1 and data were interpreted using growth analysis. Seedlings of S. inaequidens exhibited a slow growth in low nutrient availability and a rapid growth rate when exposed to high nutrient availability, which contributed to producing a large leaf area and biomass. Four types of pollination were studied in S. inaequidens. Hand-pollinations showed that S. inaequidens was mostly self-incompatible. Self-pollinated capitula only showed 7.6% viable achenes, significantly lower than insect-pollinated and cross-pollinated capitula. Some individuals were more self-fertile than others ensuring the production of some offspring in a situation of colonisation. Natural pollination occurred with generalist insects. Most of these results confirm the invasive ability of S. inaequidens.     

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.     

Effect of maturation conditions on light and temperature requirements during seed germination of Citrullus colocynthis from the Arabian Desert

• Seed germination of Citrullus colocynthis, as in many other species of Cucurbitaceae, is inhibited by light, particularly at low temperatures. Germination response to light and temperature has been attributed to day length and temperature during seed maturation. This study assessed the effects of these factors on the germination response of C. colocynthis to temperature and light quality.
• Ripe fruits were collected from natural habitats during December and February and germinated at three temperatures (15/25, 20/30 and 25/35 °C) in five light treatments (dark, white light and Red:Far Red (R:FR) ratios of 0.30, 0.87 and 1.19). Additionally, unripe fruits were also collected from natural habitats and completed their maturation in growth chambers under different day lengths (6, 16 and 24 h of darkness) at 10/20 °C, and in darkness at both 10/20 °C and 25/35 °C. Mature seeds of the different treatments were germinated in the same five light treatments at 15/25 °C.
• Germination was significantly higher in the dark than that in any light treatment. Seeds matured at higher temperatures (i.e. seeds from the December collection and those matured at 25/35 °C) had significantly higher germination than those matured at lower temperatures (i.e. seeds from the February collection and those matured at 10/20 °C). Dark germination was significantly higher for the December collection than for the February collection. Seeds of the two collections germinated in the dark only at 15/25 °C. However, seeds matured in a growth chamber at 10/20 °C in darkness germinated at 15/25 °C in all light treatments, except for the R:FR ratio 0.30. Seeds of the different treatments failed to germinate in FR‐rich light.
• This study demonstrates that both temperature and day length during seed maturation play significant roles in the germination response of C. colocynthis. Additionally, the dark requirement for germination is likely beneficial for species with the larger seeds, such as C. colocynthis, which produce bigger seedlings that are able to emerge from deep soils and are competitively superior under dense vegetation and resource‐limited conditions.

     

Seed germination and dormancy in the medicinal woodland herbs Collinsonia canadensis L. (Lamiaceae) and Dioscorea villosa L. (Dioscoreaceae)

We used a double germination phenology or “move-along” experiment (sensu Baskin and Baskin, 2003) to characterize seed dormancy in two medicinal woodland herbs, Collinsonia canadensis L. (Lamiaceae) and Dioscorea villosa L. (Dioscoreaceae). Imbibed seeds of both species were moved through the following two sequences of simulated thermoperiods: (a) 30/15 °C→20/10 °C→15/6 °C→5 °C→15/6 °C→20/10 °C→30/15 °C, and (b) 5 °C→15/6 °C→20/10 °C→30/15 °C→20/10 °C→15/6 °C→5 °C. In each sequence, seeds of both species germinated to high rates (>85%) at cool temperatures (15/6 and 20/10 °C) only if seeds were previously exposed to cold temperatures (5 °C). Seeds kept at four control thermoperiods (5, 15/6, 20/10, 30/15 °C) for 30 d showed little or no germination. Seeds of both species, therefore, have physiological dormancy that is broken by 12 weeks of cold (5 °C) stratification. Morphological studies indicated that embryos of C. canadensis have “investing” embryos at maturity (morphological dormancy absent), whereas embryos of D. villosa are undeveloped at maturity (morphological dormancy present). Because warm temperatures are required for embryo growth and cold stratification breaks physiological dormancy, D. villosa seeds have non-deep simple morphophysiological dormancy (MPD). Neither species afterripened in a 6-month dry storage treatment. Cold stratification treatments of 4 and 8 weeks alleviated dormancy in both species but C. canadensis seeds germinated at slower speeds and lower rates compared to seeds given 12 weeks of cold stratification. In their natural habitat, both species disperse seeds in mid- to late autumn and germinate in the spring after cold winter temperatures alleviate endogenous dormancy.     

Seed germination ecology of the annual grass Leptochloa panicea ssp. mucronata and a comparison with L. panicoides and L. fusca

Leptochloa panicea ssp. mucronata is an annual grass that grows in relatively dry habitats. Requirements for dormancy loss and germination were determined for seeds of this species and compared to those of two species from wet habitats. Seeds of L. panicea were dormant at maturity in autumn, but when exposed to actual or simulated autumn temperatures (e.g. 20/10, 15/6 °C), they entered conditional dormancy and thus germinated to high percentages in light at 35/20 °C. Seeds buried in non-flooded soil exposed to natural seasonal temperature changes in Kentucky (USA) were non-dormant by the following summer and germinated to 80–100 % in light at 25/15, 30/15 and 35/20 °C. Seeds buried in non-flooded soil exhibited an annual conditional dormancy/non-dormancy cycle, with seeds mostly germinating to 80–100 % in light at 30/15 and 35/20 °C throughout the year but to 80–100 % in light at 25/15 °C only in summer. Results for L. panicea were compared to published data for L. panicoides and L. fusca. Whereas seeds of L. panicea buried in flooded soil failed to come out of dormancy, those of L. panicoides, an annual of moist habitats such as mudflats, exhibited an annual conditional dormancy/non-dormancy cycle, and those of L. fusca, a semi-aquatic, required flooding for both dormancy loss and germination. Differences in dormancy breaking and germination responses of seeds of Leptochloa species may help to explain why this genus occupies a wide range of habitats with regard to soil moisture conditions.     

Effect of salinity,temperature and hypersaline conditions on the seed germination in Limonium mansanetianum an endemic and threatened Mediterranean species

Abstract

Limonium mansanetianum is catalogued as critically threatened (CR) species and it is included in Valencian Catalogue of Threatened Plant Species. Limonium mansanetianum is a gypsicolous species, which only lives in a restricted area to south-centre of Valencia province (Spain). The species is a low-branched woody shrub with summer flowering. The influence of incubation temperature (10°, 15°, 20° and 25°/20?°C) and salinity (0%–3.0% NaCl) on seed germination of L. mansanetianum was studied. Best seed germination was obtained in distilled water controls. Seed germination decreased with an increase in salinity and few seeds germinated at 2.5% and 3.0% NaCl. Optimal temperature regime for germination was 15?°C where germination in 0.5% and 1.0% NaCl was not affected. Recovery and hypersaline conditions experiments showed that L. mansanetianum seeds displayed a greater tolerance to high salinity and temperature stress before germination.     

Germination requirements and dispersal timing in two heterocarpic weedy asteraceae

In SW Spain the winter annuals Anacyclus radiatus and Chrysanthemum coronarium (Asteraceae) are found as weeds in diverse crops. Both plant species are heterocarpic, i.e. the peripheral and central achenes of the capitulum are morphologically distinctive. In heterocarpic and heterospermic species the different fruit or seed morphs usually have differential ecological behaviour. In this work we have studied the morphometry, germination and dispersal timing of t he different achene morphs in A. radiatus and C. coronarium. Laboratory germination tests were carried out to evaluate the influence of incubation temperature (light/dark, 27/27, 24/18, 20/10, 15/5, 10/4 degrees C), chilling period (0, 1, 7, 21 and 60 days at 2 degrees C), GA3, nitrate and the dark on the germination percentage and rate (t50). The peripheral achenes of A. radiatus have a significantly broader wing than the central achenes. In this species, germination was inhibited in the dark but viable achenes of both types germinated completely under light exposure irrespective of test conditions. Nevertheless, the peripheral achenes germinated significantly faster compared to the central achenes (t5o 1.04 vs. 1.55 days at 24/18 degrees C). In C. coronarium peripheral achenes have three-winged ribs and are significantly longer, wider and thicker than the central achenes, which have only one- or two-winged ribs. In this species the peripheral achenes showed a much lower germination compared to the central achenes under all conditions tested (0.3-3.5% vs. 41.5-58.0%). Embryos isolated from the pericarp of peripheral achenes germinated quickly in a great extent, and when placed in close contact with a pericarp fragment they did not show any inhibition of germination, suggesting physical dormancy. To determine differences in dispersal timing between achene morphs, we monitored dispersal in a stand of both plant species. In A. radiatus the peripheral achenes were shed first, at the onset of the rainy season in late summer, whereas the central achenes dispersed from late summer to early winter. In contrast, in C. coronarium the central achenes were shed first, from early summer to early autumn, whereas the peripheral achenes dispersed mainly in early autumn. Thus, in both weed species the more dispersible and readier-to-germinate achene morph is dispersed first. The differences in the germination and dispersal behaviour between achene morphs represent an opportunistic, mixed strategy which is favourable under environmental uncertainty, and should be accounted for in order to understand the population dynamics of both weed species.     

IMPORTANCE OF RESTING STAGES IN DIATOM SEASONAL SUCCESSION1

The role of environmental factors in seasonal succession of six clones of marine diatoms was examined by testing effects of various combinations of temperature (5°, 10°, 15°, 20°C) and photoperiod (10, 12, 14, 16 h) on time to germination of resting stages and subsequent vegetative cell division rates. Resting stages were induced in batch cultures using a low nutrient medium and a cold, dark period. A three-factor ANOVA showed that species, temperature, daylength, and their interactions signficantly (P =0.0001) injuenced germination time and division rate. Resting stages of Chaetoceros similis Cleve and Odontella aurita (Lyngbye) Agardh were most successful in producing populations of cells on warm, long days (20°C/16 h). Chaetoceros didymus Ehrenberg and C. cinctus Gran were most successful under more moderate conditions (15°C/14 h). Resting stages of Ditylum bright-wellii (West) Grunow germinated and cells divided rapidly under all conditions. No consistent pattern was evident with Thalassiosira rotula Meunier. We compared results of the laboratory study with environmental conditions prevailing during the presence of these species in a British Columbia fjord. Favorable growth conditions determined for C. didymus, D. brightwellii, and T. rotula were similar to conditions present when these species were commonly found in the fjord.     

Ecology of achene dimorphism in Leontodon saxatilis

BACKGROUND [corrected] AND AIMS: Leontodon saxatilis produces two morphologically distinct achenes (morphs) in a single capitulum: one row of dark brown achenes without a pappus lies at the edge ('peripheral achenes'; 0.74 +/- 0.18 mg) while the inner ones are light brown with a pappus ('central achenes'; 0.38 +/- 0.07 mg). The hypothesis that achene heteromorphism in L. saxatilis widens its ecological amplitude was tested. KEY RESULTS: Achenes of both morphs germinated over the same range of temperatures (6-33 degrees C) but the central achenes showed significantly higher germination percentages, and the two also differed significantly in their annual dormancy cycle, with the peripheral achenes showing greater dormancy for part of the year. Seedlings from the two morphs did not differ significantly in total biomass after 2 and 4 weeks of growth, neither did they differ significantly in root and shoot weight and root:shoot ratio. Plants from both morphs growing at different regimes of soil moisture, nutrients and competition did not differ significantly in their number of achenes per capitulum. While the number of central achenes varied, that of peripheral achenes remained constant at approx. 13. Drier soil led to an increase in the number of central achenes in plants from both morphs. CONCLUSIONS: The peripheral achenes can replace the mother plant in the following growing season, whereas the central achenes are well adapted for wind dispersal and thus for colonization of new sites. However, the extent of variation in germination characteristics was similar to that found in seed populations of homomorphic plants, which suggests that germination percentage and other growth characteristics do not contribute to widening the ecological amplitude. The habitat of most heteromorphic species, the morphs of which differ greatly in germination and/or growth characteristic, are deserts or highly disturbed areas where such differences are highly advantageous, but in the moderate habitat of L. saxatilis the differences may prove a disadvantage.     

Enhancement of ABA responsiveness in wheat embryos by high temperature*†

Abstract. Mature wheat (Triticum aestivum L.) grain often possesses high-temperature dormancy which restricts the grain from germinating at warm temperatures (25–30°C). Isolated embryos from such grain exhibited little high-temperature dormancy when germinated in water. Dormancy was restored by the application of abscisic acid (ABA) to the embryos. The ability of ABA to block germination in isolated embryos was enhanced significantly by elevating the germination temperature. ABA was 100 times more effective in reducing embryonic germination at 30°C than at 15°C. These temperature effects on embryonic response to ABA are a useful system for studying the mechanism of ABA action in seed dormancy.     

Achene germination of the spring ephemeroid species Carex physodes in the Gurbantunggut Desert

Much of the seed germination research on Carex has focused on wetland species, and little is known about the species of arid habitats. Here, we investigated seed dormancy and germination of Carex physodes, which is an important component of the plant community of the Gurbantunggut Desert of the Junggar Basin in Xinjiang, China. Our studies included the effects of mechanical and chemical scarification, dry storage, treatment with GA₃, wet‐cold stratification, and burial in the field. No freshly matured achenes germinated over a range of temperature regimes after treatment with GA₃, 6 months of dry storage or removal of part of the endosperm. The mechanical scarification resulted in < 5% achene germination, however, higher percentage of achene germination occurred after removal of the pericarp (60%), H₂SO₄ scarification (30%) or scarification in 10% NaOH (85%). Six and nine months of wet‐cold stratification promoted < 40% achene germination. The optimal germination temperatures ranged from 25/10°C to 35/20°C. Maximum germination after 9 months of burial at a depth of 3 cm in the field was 36%. These results indicate that the seeds have non‐deep physiological dormancy (PD) and that the pericarp contains germination inhibitors and has strong mechanical resistance to germination.