A role for fruit structure in seed survival and germination of Swartzia langsdorffii Raddi beyond dispersal

• Diaspore structure has been hypothesised to play a role in seed viability and/or germination of recalcitrant seeds, especially for Swartzia langsdorffii. Thus, this work aims to (i) investigate the in situ contribution of pericarp and aril on seed viability and germination, and (ii) identify morphoanatomical traits of S. langsdorffii diaspores that allow its desiccation‐sensitive seeds to remain viable.
• The role of the pericarp and aril in seed survival and germination was investigated by placing the whole fruit, whole seeds (arillate seed) and bare seeds (without aril) in soil in the forest understorey, assessing germination, emergence, dead, firm and predated seeds, and water content of pericarps, arils and seeds. Correlation analysis was performed between environmental variables and physiological parameters. Histochemical features of diaspores were also investigated.
• Pericarp water content fell after several months, while the aril maintained its water content. Seeds did not lose water even without the presence of the pericarp and aril. However, presence of the pericarp promoted seed water content, viability and germination long after dispersal. The embryo had a thickened outer periclinal cell wall.
• Pericarp and aril are not essential to prevent water loss in seeds, but do help to retain seed moisture, favouring viability maintenance and promoting germination during the rainy season. Morphoanatomical features of seeds are suggested as main factors that reduce water loss. Survival of these desiccation‐sensitive seeds upon dispersal during the dry season appears to be facilitated by multiple diaspore features that prevent viability loss.

     

Seasonal rhythms of seed rain and seedling emergence in two tropical rain forests in southern Brazil

Seasonal tropical forests show rhythms in reproductive activities due to water stress during dry seasons. If both seed dispersal and seed germination occur in the best environmental conditions, mortality will be minimised and forest regeneration will occur. To evaluate whether non-seasonal forests also show rhythms, for 2 years we studied the seed rain and seedling emergence in two sandy coastal forests (flooded and unflooded) in southern Brazil. In each forest, one 100 x 30-m grid was marked and inside it 30 stations comprising two seed traps (0.5 x 0.5 m each) and one plot (2 x 2 m) were established for monthly monitoring of seed rain and a seedling emergence study, respectively. Despite differences in soil moisture and incident light on the understorey, flooded and unflooded forests had similar dispersal and germination patterns. Seed rain was seasonal and bimodal (peaks at the end of the wetter season and in the less wet season) and seedling emergence was seasonal and unimodal (peaking in the wetter season). Approximately 57% of the total species number had seedling emergence 4 or more months after dispersal. Therefore, both seed dormancy and the timing of seed dispersal drive the rhythm of seedling emergence in these forests. The peak in germination occurs in the wetter season, when soil fertility is higher and other phenological events also occur. The strong seasonality in these plant communities, even in this weakly seasonal climate, suggests that factors such as daylength, plant sensitivity to small changes in the environment (e.g. water and nutrient availability) or phylogenetic constraints cause seasonal rhythms in the plants.     

Sensitivity of seed germination to water stress in high-altitude populations of a threatened palm species

• Divergence in seed germination patterns among populations of the same species is important for understanding plant responses to environmental gradients and potential plant sensitivity to climate change. In order to test responses to flooding and decreasing water potentials, over 3 years we germinated and grew seeds from three habitats of Euterpe edulis Mart. occurring along an altitudinal gradient.
• Seed germination and root growth were evaluated under different water availability treatments: control, flood, −0.4 MPa, −0.8 MPa, in the years 2012, 2013 and 2014, and in the final year of the experiment (2014) at −1.0 MPa and −1.5 MPa.
• Seeds from the montane habitat did not germinate in the flooding treatment. Seed germination of all three habitats decreased in the −1.5 MPa treatment and the montane habitat had lowest germination in this treatment. Time required for half of the seeds to germinate increased up to −0.8 MPa. Seeds from montane habitats germinated more slowly in all treatments. The only difference in seed germination synchrony was an increase in the submontane population under the flooding treatment. However, synchrony decreased at the lowest water potentials. Roots of the montane population were more vigorous in most treatments, except at −0.8 MPa.
• The unusual ability of these seeds to germinate at low water potentials might be related to early seed germination at the onset of the rainy season, which potentially decreases seed predation pressure. Seeds of the montane population were more sensitive to both types of water stress. A predicted increase in the frequency and intensity of extreme high rainfall or drought events may predispose early stages of this population to adverse factors that might negatively affect population viability with elevational in future climate change scenarios.

     

热带雨林林内小生境对木奶果和染木种子萌发与幼苗存活的影响

森林生态系统中小生境的差异可能影响种子萌发与幼苗的建立,并形成物种不同的分布格局。为探讨生境异质性对植物分布的影响,该研究在西双版纳热带季节性雨林中选取5个具有不同海拔和地形的小生境,人工散布木奶果与染木种子,定期监测种子萌发与幼苗存活,同步记录小生境的近地面（距地面3 cm）温度、近地面(3 cm)空气湿度和表层土壤(5 cm)含水量变化,并在实验室开展种子脱水和不同水势条件下的种子萌发实验。结果表明：（1）实验所选的5个小生境中,沟底地带的土壤水分含量在一年中始终都高于其他小生境;5个小生境的空气相对湿度在雨季与雾凉季都接近饱和,只有在干热季出现明显的差别,即随海拔的升高,空气湿度下降。同时,在旱季,近地面温度随海拔的升高而升高, 出现1~2个月的山地逆温现象。（2）木奶果与染木具有不同的种子传播与萌发规律,而且染木种子在不同生境下,其种子萌发率和幼苗存活率有显著的差异。其中：木奶果种子在雨季成熟散布并快速萌发,以幼苗的形式度过干热季,其种子可以在相对较低的水势下萌发;染木种子在雨季末期成熟并进入休眠期,以种子的形式度过干热季,其种子相对较抗脱水。研究认为,地形是决定西双版纳热带季节性雨林生境异质性的重要因子,小生境在温度和水分方面的差异在干热季时尤为明显,并对木奶果和染木种子的萌发或幼苗存活产生了重要影响, 而木奶果与染木种子自身的特性使其对于西双版纳季节性的干旱具有不同的适应对策。     

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Combined effect of temperature and water stress on seed germination of four Leptocereus spp. (Cactaceae) from Cuban dry forests

Cactus seeds have developed adaptations to survive with high temperatures and low soil moisture in their habitats. We studied the effect of the combination of four water potentials (0, −0.2, −0.4 and −0.6 MPa) and two temperatures (25°C and 35°C) on germination and seedling mass of four endemic Leptocereus species from Cuba. There were two semi-arid coast species (L. arboreus and L. santamarinae) and two species inhabiting dry inland karstic hills (L. ekmanii and L. scopulophilus). We hypothesized that: (a) a decrease in water potential and an increase in temperature would result in low and slow germination, as well as in low seedling mass, and (b) ungerminated cactus seeds after exposure to combined water and temperature stress would have a high recovery capacity. The minimum time required for the seeds to start germination (Tmin), mean germination time (MGT) and germinability were evaluated. In addition, seed mass, the recovery after treatments of non-germinated seeds, and the seedling fresh mass obtained under different treatments were compared between species. In general, germination was only obtained at 25°C and germinability and seedling mass were drastically affected by the reduction from 0 MPa to −0.2 MPa. Seeds showed thermoinhibition at 35°C at all water potentials. There was a tendency to increase the MGT with decreasing water potential in three species. Low seed recovery occurred at all combined treatments for three species. If the predictions of increased temperature and decreased rainfall for the Caribbean region occur, a reduction in the germination of the Leptocereus species studied is expected.     

Dormancy cycles in buried seeds of three perennial Xyris (Xyridaceae) species from the Brazilian campo rupestre

• Dormancy cycles are an important mechanism for avoiding seed germination under unfavourable periods for seedling establishment. This mechanism has been scarcely studied in tropical species. Here, we studied three tropical and perennial species of Xyris, X. asperula, X. subsetigera and X. trachyphylla, to investigate in situ longevity and the existence of seasonal seed dormancy cycles.
• Seeds of three species of Xyris were buried in their natural habitat, with samples exhumed bimonthly for 18 months. Germination of exhumed seeds was assessed under a 12‐h photoperiod over a broad range of temperatures. Seeds of X. trachyphylla were also subjected to treatments to overcome secondary dormancy.
• Seeds of all species are able to form a persistent seed bank and exhibit seasonal changes in germinability. Secondary dormancy was acquired during the rainy summer and was overcome during the subsequent dry season (autumn/winter). Desiccation partially overcomes secondary dormancy in X. trachyphylla seeds.
• Soil seed bank persistence and synchronisation of seed germination under favourable conditions for seedling establishment contribute to the persistence and regeneration of X. asperula, X. subsetigera and X. trachyphylla in their natural environment.

     

Patterns of Seed Size,Germination and Seed Viability of Tropical Tree Species in Southern India1

Seed weight, days to germination and seed viability were observed for 99 species growing in the Western Ghats of Karnataka, India. Seed size was strongly correlated with days to germination; smaller seeds germinated faster than larger seeds. Species which flowered during the rainy season had lighter seeds than species which flowered during the dry season. It was also found that seed size and viability of seeds were related to the season of fruiting. Species which fruit during the rainy season had heavier seeds and shorter viability than species which fruit during the dry season. These flowering and fruiting patterns and varying seed sizes are argued to be adaptations to the time of dispersal, time of moisture availability in the habitat and seedling survival.     

Responses of Primula vulgaris to light quality in the maternal and germination environments

• In the model species Arabidopsis thaliana phytochromes mediate dormancy and germination responses to seasonal cues experienced during seed maturation on the maternal plants. However, the effect of the maternal light environment on seed germination in native wild species has not been well studied. This is particularly important given its practical application in the context of environmental restoration, when there can be marked changes in the canopy.
• Plants of Primula vulgaris were grown in the field over two vegetative seasons under four shading treatments from low to high ratio of red to far‐red light (R:FR). Leaf and seed traits were assessed in response to the light treatments. The germination of seeds from these four maternal environments (pre‐dispersal) was investigated at seven light and five temperature treatments (post‐dispersal).
• Thinner leaves, larger leaf area and greater chlorophyll content were found in plants growing in reduced R:FR. Shading in the maternal environment led to increased seed size and yield, although the conditions experienced by the maternal plants had no effect on seed germination. Seeds responded strongly to the cues experienced in their immediate germination environment. Germination was always enhanced under higher R:FR conditions.
• The observed phenotypic trait variation plays a major role in the ability of P. vulgaris to grow in a wide range of light conditions. However, the increased germination capacity in response to a higher R:FR for all maternal environments suggests potential for seedling establishment under vegetative shade only in the presence of canopy gaps.

     

Synchrony of Seed Dispersal, Hydrology and Local Climate in a Semi-arid River Reach in California

The temporal availability of propagules is a critical factor in sustaining pioneer riparian tree populations along snowmelt-driven rivers because seedling establishment is strongly linked to seasonal hydrology. River regulation in semi-arid regions threatens to decouple seed development and dispersal from the discharge regime to which they evolved. Using the lower Tuolumne River as a model system, we quantified and modeled propagule availability for Populus fremontii (POFR), Salix gooddingii (SAGO), and Salix exigua (SAEX), the tree and shrub species that dominate near-channel riparian stands in the San Joaquin Basin, CA. A degree-day model was fit to field data of seasonal seed density and local temperature from three sites in 2002–2004 to predict the onset of the peak dispersal period. To evaluate historical synchrony of seed dispersal and seasonal river hydrology, we compared peak spring runoff timing to modeled peak seed release periods for the last 75 years. The peak seed release period began on May 15 for POFR (range April 23–June 10), May 30 for SAGO (range May 19–June 11) and May 31 for SAEX (range May 8–June 30). Degree-day models for the onset of seed release reduced prediction error by 40–67% over day-of-year means; the models predicted best the interannual, versus site-to-site, variation in timing. The historical analysis suggests that POFR seed release coincided with peak runoff in almost all years, whereas SAGO and SAEX dispersal occurred during the spring flood recession. The degree-day modeling approach reduce uncertainty in dispersal timing and shows potential for guiding flow releases on regulated rivers to increase riparian tree recruitment at the lowest water cost.     

Reduced germination success of temperate grassland seeds sown in dung: consequences for post‐dispersal seed fate

• Endozoochory is one of the main drivers shaping temperate grassland communities by maintaining plant populations of its constituents and enabling plants to colonize new habitats. Successful endozoochorous dispersal implies that seeds not only get consumed and survive the digestive tract but are also able to develop into viable seedlings in a dung environment.
• We experimentally assessed the germination probability and timing of 15 annual and perennial temperate European grassland species in cattle and horse dung and in different climatic conditions (greenhouse and outdoor conditions).
• Interspecific variation in germinability and germination timing are found, while life strategy had only an effect on germination timing. We found adverse effects of both cattle and horse dung on the germination characteristics of all tested grassland species, but the effects of cattle dung were more pronounced. In comparison with the control treatment, fewer seeds emerged in dung and more time was needed to germinate. Also, germination metrics clearly differed between the artificial greenhouse and outdoor conditions, with generally a lower germinability in outdoor conditions.
• According to our results, a large cost seems to be associated with endozoochorous dispersal in this stage of the life cycle, as seed dispersal effectiveness strongly depends on the quality of the deposition site with a lowered survival and germination probability when seeds are deposited in dung.

     

Effects of season and population size on pollination and reproductive output in a Mediterranean shrub

• Pollinator guilds may change throughout extended flowering periods, affecting plant reproductive output, especially in seasonal climates. We hypothesised a seasonal shift in pollinator guild and an autumn reduction in pollinator abundance, especially in small and sparse populations.
• We recorded pollinator identity, abundance and behaviour in relation to flower density from plant to population throughout the extended flowering of Ononis tridentata. We evaluated female reproductive output by recording pollination success and pre‐dispersal seed predation in eight populations of contrasting size and density. Offspring quality was also characterised through seed weight and germination.
• A diverse guild of insects visited O. tridentata in spring, while only Apis mellifera was observed in autumn. Visitation frequency did not vary seasonally, but the number of flowers per foraging bout was lower, and seeds were heavier and had a higher germination rate in autumn. Plant and neighbourhood flowering display were not related to pollinator visitation frequency or behaviour. However, the rate of fertilised ovules, seed set and autumn flowering display size were positively related to population density.
• The maintenance of pollination in autumn enhances the reproductive performance of O. tridentata due to higher quality of autumn seed, and to a large reduction in seed predator pressure. We also suggest that observed changes in pollinator behaviour could be one of the processes behind seasonal variation in seed performance, since geitonogamous crosses were less likely to occur in autumn.

     

Rainfall seasonality predicts the germination behavior of a tropical dry‐forest vine

Seed dormancy is considered to be an adaptive strategy in seasonal and/or unpredictable environments because it prevents germination during climatically favorable periods that are too short for seedling establishment. Tropical dry forests are seasonal environments where seed dormancy may play an important role in plant resilience and resistance to changing precipitation patterns. We studied the germination behavior of seeds from six populations of the Neotropical vine Dalechampia scandens (Euphorbiaceae) originating from environments of contrasting rainfall seasonality. Seeds produced by second greenhouse‐generation plants were measured and exposed to a favorable wet environment at different time intervals after capsule dehiscence and seed dispersal. We recorded the success and the timing of germination. All populations produced at least some dormant seeds, but seeds of populations originating from more seasonal environments required longer periods of after‐ripening before germinating. Within populations, larger seeds tended to require longer after‐ripening periods than did smaller seeds. These results indicate among‐population genetic differences in germination behavior and suggest that these populations are adapted to local environmental conditions. They also suggest that seed size may influence germination timing within populations. Ongoing changes in seasonality patterns in tropical dry forests may impose strong selection on these traits.     

Hydrodynamics and seed dispersal in the lower Amazon

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Seed traits and bird species influence the dispersal parameters of wetland plants

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Factors Limiting Tropical Rain Forest Regeneration in Abandoned Pasture: Seed Rain,Seed Germination,Microclimate, and Soil1

Vegetation, seed rain, seed germination, microclimate, and soil physical and chemical parameters were measured in a recently abandoned pasture and adjacent primary rain forest in southern Costa Rica. The goal of this study was to assess the importance of these factors in limiting forest regeneration in abandoned pastures. Seed rain of animal dispersed species decreased dramatically in the pasture >5 m from the forest/pasture edge; fewer wind dispersed seeds fell in the pasture than in the forest, but the difference was much less than for animal dispersed seeds. Percent seed germination of most species studied was similar in the forest and in pasture with grasses; seed germination was lower during the dry season in areas of pasture cleared of grasses. Air temperature, vapor pressure deficit (VPD), and photon flux density (PFD) were much higher in the pasture than in the forest at 1 m above the ground. VPD and PFD at ground level and soil temperature were similar in the pasture and the forest, indicating that pasture grasses strongly modify microclimatic conditions near the soil surface. The lowest gravimetric water content recorded in the pasture during the dry season was 0.5 and leaf relative water contents of the two species measured in the forest and pasture were identical, suggesting that plants in the pasture were not water stressed. Levels of most soil nutrients were lower in the pasture as compared to the forest; however, aboveground and root biomass for seedlings grown in pasture and forest soils did not differ significantly. Although a number of factors impede forest recovery in abandoned pastures, these results suggest that the most imporrant limitation is lack of seed dispersal.     

Sensitivity cycling in physically dormant seeds of the Neotropical tree Senna multijuga (Fabaceae)

• Cycling of sensitivity to physical dormancy (PY) break has been documented in herbaceous species. However, it has not been reported in tree seeds, nor has the effect of seed size on sensitivity to PY‐breaking been evaluated in any species. Thus, the aims of this study were to investigate how PY is broken in seeds of the tropical legume tree Senna multijuga, if seeds exhibit sensitivity cycling and if seed size affects induction into sensitivity.
• Dormancy and germination were evaluated in intact and scarified seeds from two collections of S. multijuga. The effects of temperature, moisture and seed size on induction of sensitivity to dormancy‐breaking were assessed, and seasonal changes in germination and persistence of buried seeds were determined. Reversal of sensitivity was also investigated.
• Fresh seeds were insensitive to dormancy break at wet–high temperatures, and an increase in sensitivity occurred in buried seeds after they experienced low temperatures during winter (dry season). Temperatures ≤20 °C increased sensitivity, whereas temperatures ≥30 °C decreased it regardless of moisture conditions. Dormancy was broken in sensitive seeds by incubating them at 35 °C. Sensitivity could be reversed, and large seeds were more sensitive than small seeds to sensitivity induction.
• Seeds of S. multijuga exhibit sensitivity cycling to PY‐breaking. Seeds become sensitive during winter and can germinate with the onset of the spring–summer rainy season in Brazil. Small seeds are slower to become sensitive than large ones, and this may be a mechanism by which germination is spread over time. Sensitive seeds that fail to germinate become insensitive during exposure to drought during summer. This is the first report of sensitivity cycling in a tree species.

     

Diversity of epicotyl dormancy among tropical montane forest species in Sri Lanka

• Fruiting season of many Sri Lankan tropical montane species is not synchronised and may not occur when conditions are favourable for seedling establishment. We hypothesised that species with different fruiting seasons have different seed dormancy mechanisms to synchronise timing of germination with a favourable season for establishment. Using six species with different fruiting seasons, we tested this hypothesis.
• Germination and imbibition of intact and manually scarified seeds were studied. Effect of GA₃ on germination was examined. Embryo length:seed length (E:S) ratio of freshly matured seeds and of those with a split seed coat was determined. Time taken for radicle and plumule emergence and morphological changes of the embryos were recorded.
• The radicle emerged from Ardisia missionis, Bheza nitidissima and Gaetnera walkeri seeds within 30 days, whereas it took >30 days in other species. Embryos grew in seeds of B. nitidissima and G. walkeri prior to radicle emergence but not in Microtropis wallichiana, Nothapodytes nimmoniana and Symplocos cochinchinensis. A considerable delay was observed between radicle and plumule emergence in all six species. Warm stratification and/or GA₃ promoted germination of all species.
• All the tested species have epicotyl dormancy. Seeds of B. nitidissima and G. walkeri have non‐deep simple morphophysiological epicotyl dormancy, and the other four species have non‐deep physiological epicotyl dormancy. Differences in radicle and epicotyl dormancy promote synchronisation of germination to a favourable time for seedling development. Therefore, information on dormancy‐breaking and germination requirements of both radicle and epicotyl are needed to determine the kind of dormancy of a particular species.

     

Trade-offs between seed dispersal and dormancy in an amphi-basicarpic cold desert annual

Background and Aims

Several studies have demonstrated trade-offs between depth of seed dormancy and dispersal ability for diaspore-dimorphic species. However, relatively little is known about trade-offs between these two life history traits for a species that produces more than two diaspore morphs. The aim of this study was to investigate the relationship between seed dormancy and dispersal in Ceratocarpus arenarius, an amphi-basicarpic cold desert annual that produces a continuum of dispersal unit morphs.

Methods

A comparison was made of dispersal and dormancy breaking/germination responses of dispersal units from ground level (a), the middle of the plant canopy (c) and the top of the plant canopy (f). Various features of the morphology and mass of dispersal units and fruits (utricles) were measured. The role of bracteoles in diaspore dispersal by wind, settlement onto the soil surface and dormancy/germination was determined by comparing responses of intact dispersal units and fruits. Movement of dispersal units by wind and animals, seed after-ripening, germination phenology and the presence of water-soluble germination inhibitors in bracteoles were tested using standard procedures.

Key Results

Dispersal units a, c and f differed in morphology and mass; in the majority of cases, extremes were exhibited by a and f, with c being intermediate. Overall, relative dispersal ability was f > c > a, whereas relative intensity of dormancy was a > c > f. Bracteoles increased dispersal distance by wind, enhanced settlement of diaspores onto the soil surface and mechanically inhibited germination.

Conclusions

The results provide evidence for a model in which there is a continuous inverse-linear relationship between diaspore dispersal ability and depth of dormancy. Thus, dispersal unit heteromorphism of C. arenarius results in a continuum, from no dispersal ability/high dormancy (dispersal unit a) to high dispersal ability/low dormancy (unit f), which may be a bet-hedging strategy in the cold desert environment.     

Misunderstanding on germination sensu stricto leads us to a false positive germination-dormancy balance in diaspores of Paepalanthus chiquitensis Herzog (Eriocaulaceae), a threatened everlasting flowering species

Although germination sensu stricto and germination–dormancy balance are essential to each other, few reports demonstrate the effects of their relationship on patterns of seed–seedling transition. We studied this relationship by using diaspores of Paepalanthus chiquitensis, a threatened everlasting flowering species. We assessed three aspects: (a) water dynamics in germinating diaspores, (b) thermal stimulation (35, 40, 50, 60 and 70°C for 30 min) and (c) imbibition (4 hr) in growth stimulators (100, 200 or 300 mg · L⁻¹ GA₃; 0.1% KNO₃). Our findings demonstrate that there are no barriers in the diaspore for diffusion. Additionally, there is high variability in diaspore permeability, and the germination of the species has two steps divided into five stages. These stages were defined by inflections on velocity in water dynamics, which ranged from −0.05 to 0.05 g · hr⁻¹, where only the last stage (Phase V) is associated with the second step of germination sensu stricto. The imbibition phase was the shortest germination phase (1.3 hr), whereas the predominantly biochemical phase was the longest (5 days after sowing [DAS]). Germinability was high (77.5–86.5%), with high daily embryo protrusion (2.46–3.03 diaspores · day⁻¹). Protocorm emergence marked the end of the first step of germination sensu stricto (9 DAS), and first protocorm rupture (11 DAS) marked the end of the second step. The growth stimulus only concentrated the germination process, showing that the diaspores do not have primary dormancy. Knowing these diaspores’ peculiarities during germination sensu stricto avoids a false positive statement on the existence of diaspore dormancy.