The flora of Mediterranean ecosystems contains families with species having fully and under-developed embryos in their seeds. After-ripening for physiological dormancy release and smoke influence germination in many species. We investigated how after-ripening and embryo growth interact with smoke to influence the temporal dynamics of seedling emergence among fire ephemerals. Seeds were placed in the field and under standardized (50% relative humidity, 30°C) laboratory conditions to test the effects of summer conditions on physiological dormancy loss. Germination was tested with water or smoke compounds (smoke water, KAR1) at a simulated autumn/winter temperature (18/7°C). The timing and amount of seedling emergence with smoke was observed for seeds exposed to near-natural conditions. During summer, physiological dormancy was broken in all species, enabling germination at autumn/winter but not summer temperatures; no embryo growth occurred in seeds with under-developed embryos. At the start of the wet season, seedling emergence from seeds with fully developed embryos occurred earlier than from seeds with under-developed embryos. In a non-consistent manner among our study species, smoke and smoke compounds influenced the rate of embryo growth and amount of germination. Effects of smoke were noticeable in terms of number of emergents in the first emergence season. Among ecologically similar species, we have shown (1) that both thermal and embryo traits exclude germination in the summer, (2) how embryo size influences the timing of seedling emergence in autumn–winter, and (3) a reduced requirement for smoke in the second emergence season after a fire with a shift to reliance on seasonal cues for emergence.

Morphophysiological dormancy (MPD) is predominantly found in seeds of temperate regions and is uncommon in arid biomes. MPD has been reported in a number of Hibbertia (Dilleniaceae) species of temperate Australia, and in a single species of the arid zone, H. glaberrima. This study aimed to examine the dormancy and germination ecology of seeds of H. glaberrima. Seeds were subjected to temperature stratification treatments designed to mimic summer and autumn conditions in the Pilbara region of Western Australia. Seed germination and embryo growth were measured. We also tested the interaction between temperature stratification and cycles of drying and wetting designed to mimic sporadic rainfall events. All temperature and moisture treatments were tested in combination (+/–) with the smoke-derived chemical karrikinolide (KAR1). Exposing dormant seeds to temperatures suitable for warm stratification (35°C) for ≥ 8 weeks, followed by incubation at 25°C, resulted in significantly higher germination compared with non-stratified seeds. Exposing seeds to dry/wet cycling in conjunction with temperature stratification did not significantly increase germination. Exposure to KAR1 increased germination under most conditions. Once seeds are shed during October to December, they are exposed to hot and sporadically wet conditions over summer, allowing MPD to be overcome in a proportion of the seed population. Seeds may germinate in autumn (March to April), in conjunction with cooler temperatures. More deeply dormant individuals may require more than one summer to overcome dormancy. Similar to other species occurring in fire-prone ecosystems, fire also plays a crucial role in the germination ecology of H. glaberrima.

Convallariamajalis has double dormancy and hypogeal germination, but no information is available on embryo growth or on the effects of light and gibberellic acid (GA3) on germination in this genus. Therefore, we investigated embryo growth and other germination features in seeds of C. keiskei and compared the data with those of Trillium camschatcense in another study. Until now, in seeds with double dormancy, embryo growth and germination (epigeal) have been studied in detail only for seeds of T. camschatcense. Phenology of embryo growth and emergence of cotyledonary petiole/root (hereafter root) and shoot in seeds of C. keiskei were monitored outdoors. Effects of temperature, light and GA3 on embryo growth and root and shoot emergence were tested under laboratory conditions. Roots emerged the first spring following seed dispersal in autumn. The embryo grew soon after root emergence, and germination was hypogeal. Seeds with an emerged root formed buds from which a shoot (leaf) emerged above ground during the second spring. Alternating temperatures and light had negative effects on root emergence, and GA3 did not substitute for cold stratification in root emergence. Seeds of C. keiskei have double dormancy, but it differs from that in T. camschatcense. Based on differences in embryo growth before (T. camschatcense) versus after (C. keiskei) root emergence, and on epigeal (T. camschatcense) versus hypogeal (C. keiskei) germination, we suggest that two types of deep simple double morphophysiological dormancy (MPD) be recognized. Since embryo growth in C. keiskei does not fit the standard definition of MPD, we propose to expand this definition.

Seeds with epicotyl dormancy reside in soil up to 15 months (or longer), being exposed to a sequence of temperatures, before seedlings completely emerge (i.e. with both roots and shoots). Heretofore, few studies have examined precise temperatures, especially in sequences, for promotion of radicle and cotyledon emergence and how they relate to environmental cues in nature. Viburnum is the best known genus to exhibit epicotyl dormancy and, as such, we investigated the Japanese V.furcatum, hypothesizing a similar kind and level of dormancy. The underdeveloped embryos in mature seeds in October were spatulate shaped, unlike those in other Viburnum species, and they elongated from late June to August of the following year. Radicles emerged after embryo growth until mid-October, followed by cotyledons from mid-April to mid-May. Temperatures required for embryo growth, radicle and cotyledon emergence in the laboratory approximated closely those in the field. Embryo elongation and radicle emergence occurred at warm temperature regimes, and gibberellic acid (GA3) did not substitute for this warm temperature requirements. Following a 120-d cold stratification of seeds with an emerged radicle, shoots emerged from seeds at 10, 15, 15/5, 20/10 and 25/15°C. We identified that seeds of V. furcatum have deep simple epicotyl morphophysiological dormancy like the majority of other Viburnum species. For propagation of the species from seeds, the nearly 2-year period for seedling emergence could be shortened to 8 months: start fresh seeds at 25/15°C (60 d) and then move them through a sequence of 15/5°C (30 d) →  0°C (120 d) →  20/10°C (30 d).

Many temperate plant genera, like Sambucus, have species with range disjunctions among North America, Europe and/or Asia. Cold stratification (sometimes in combination with warm stratification) is the primary mechanism to break seed dormancy in these species. For some of these genera showing Northern Hemispheric disjunctions, members also occur in subtropical or tropical regions, mostly confined to higher elevations where climate and vegetation differ from those in northern latitudes. We made two hypotheses concerning germination for the subtropical Taiwanese Sambucus chinensis: (1) seeds from populations exposed to warm temperatures would require warm stratification, and (2) seeds from populations exposed to cold temperatures need cold stratification. We investigated the germination (including embryo growth) of non-stratified seeds over a range of temperatures and tested the effects of cold stratification and of gibberellins GA3 and GA4 on germination. The amount and timing of germination among populations varied substantially in response to temperature treatments. Seeds from all populations of this species required warm temperatures for dormancy break and germination, regardless of environmental conditions. As such, the majority of seeds had non-deep simple morphophysiological dormancy, which, until now, has not been reported in any members of Sambucus. The seed characteristics of the subtropical S. chinensis are different from those of temperate members of the genus in which cold stratification is the predominate treatment to overcome dormancy.

The purpose of the present study was to investigate the dormancy break and germination requirements of seeds from the rhamnaceous vine Berchemia scandens. The fleshy fruit contains a two-locular stone with an endocarp described as ‘bony, thickish’. Scarified and non-scarified stones increased by about 30–50% in mass during imbibition over a 24-h period. The endocarp of the stone does not completely enclose the seeds and a soft tissue region is present. This region is the primary area of water entrance to the seed, as shown by dye-tracking and by sealing it. Freshly matured and overwintered seeds of B. scandens germinated to low percentages at all temperatures during 2 weeks of incubation in light, and they germinated from moderate to high percentages during 12–14 weeks of incubation in light. While cold stratification had a relatively modest effect on the promotion of total germination across most temperatures assessed (if seeds were left for long enough), it had a somewhat stronger effect on germination rate. Cold-stratified seeds germinated equally well in light and darkness. The class of dormancy found in seeds of B. scandens would be physiological. The anatomy of the stones readily allows water imbibition, showing that seeds of B. scandens lack physical dormancy, an uncommon trait in Rhamnaceae.

Although the underdeveloped embryo, and thus morphological (MD) or morphophysiological (MPD) seed dormancy, is basal in angiosperms, it also occurs in advanced groups. A synthesis of the literature, combining phylogeny and the kind of seed dormancy in the highly evolutionarily advanced order Dipsacales, shows that MPD (or MD) occurs throughout all clades except the most advanced one, Valerina. Seeds of taxa in the Valerina clade have fully developed embryos and physiological dormancy (PD) or are non-dormant (ND); thus, PD and ND are derived conditions in Dipsacales. Assuming that types of seed dormancy have not changed since the Early Tertiary, the fossil record suggests that MPD (or MD) was present in extant genera of Dipsacales by the Palaeocene, but PD (or ND) not until the Miocene. Molecular dating indicates that the ages of dipsacalean lineages with MPD and PD are older than those indicated by the fossil evidence.

Requirements for dormancy break and embryo growth were determined for seeds of the western North American species, Osmorhiza depauperata. Seeds were collected in August 2001 from Sandia Crest (3200 m elevation) and Las Huertas (2300 m), New Mexico (USA). Embryos in fresh seeds were c. 0.6 mm long, and they had to grow to c. 9–10 mm before the radicle emerged from the mericarp. Embryo growth occurred at low temperatures (1 and 5°C), and seeds germinated to high percentages at 1°C during 32 weeks of incubation in the light. No seeds germinated at 5, 15/6, 20/10, 25/15 or 30/15°C during 32 weeks of incubation. Although a 4–18 week warm-temperature (25/15°C) pretreatment increased germination rates at 1°C, it was unnecessary for a high percentage of seeds to germinate. Gibberellic acid (GA3, 10–1000 mg l–1) did not substitute for cold stratification. Seeds from the low-elevation population contained larger embryos and required less time to germinate than those from the high-elevation population. O. depauperata seeds have deep complex morphophysiological dormancy (MPD), which is similar to two other western North American congeners and an Asian congener, but different from two eastern North American congeners. Results from this study suggest that: (1) phylogenetic niche conservatism has played a role in the persistence of deep complex MPD in the three western North American species of Osmorhiza; and (2) the stimulatory effect from a warm pretreatment in species needing only cold stratification for dormancy break is a preadaptation that initiated the development of an absolute warm requirement in species needing both warm and cold stratification.

The most often used time-line for distinguishing a transient seed bank from a persistent seed bank is one calendar year. Thus, species whose seeds live in or on the soil for <1 year have a transient seed bank, whereas those whose seeds live for ≥1 year have a persistent seed bank. However, dormancy cycling of seeds buried in soil has not been given due consideration in these models. When dormancy cycling is considered, it is shown that seeds of both autumn-germinators and spring-germinators are in the dormant state when they are 1 year old. Thus, unless the seeds live until at least the second germination season (i.e. usually 16–18 months following dispersal), they are, in effect, part of a transient seed bank, having lived through only one germination season. We propose that for seeds of such species to be considered part of a short-term persistent seed bank, they should remain viable and germinable until at least the second germination season, and to be part of a long-term persistent seed bank, until at least the sixth germination season. Our definitions are applicable to seeds with physiological, physical or morphophysiological dormancy, which often require >1 year after maturity to come out of dormancy in nature. We discuss modifications of the seedling emergence method for detection of a soil seed bank, so that they correspond to our definitions of seed-bank strategies.

Dormancy-breaking requirements and types of dormancy were determined for seeds of Lonicera fragrantissima Lindl. & Paxt., L. japonica Thunb., L. maackii (Rupr.) Maxim. and L. morrowii A. Gray. Seeds of all four species have underdeveloped spatulate embryos that are about 20–40%fully developed (elongated) when dispersed. Embryos in freshly matured, intact seeds grew better at 25/15°C than at 5°C. Gibberellic acid (GA3) (tested only in the light) was more effective in breaking dormancy in L. maackii and L. morrowii than in L. fragrantissima and L. japonica. Warm- followed by cold stratification was required to break dormancy in seeds of L. fragrantissima, whereas seeds of L. japonica required cold stratification only. Thus, seeds of L. fragrantissima have deep simple morphophysiological dormancy (MPD) and those of L. japonica nondeep simple MPD. About 50%of the seeds of L. maackii required warm- or cold stratification only to come out of dormancy and 50% of those of L. morrowii required warm stratification only, whereas the other 50% did not require stratification to germinate. Thus, about half of the seeds of the two species has nondeep simple MPD, and the other half has morphological dormancy (MD). In these laboratory tests, seeds of L. japonica, L. maackii, and L. morrowii generally germinated to significantly higher percentages in light than in darkness; seeds of L. fragrantissima were not tested in darkness. Peaks of germination for seeds of L. fragrantissima, L. japonica, L. maackii and L. morrowii sown on a soil surface and covered with Quercus leaves under near-natural temperature conditions shortly after seed maturity and dispersal in late June 1997, late November 1997, early November 1996 and late June 1998, respectively, occurred in early March 1998, late February 1998, late March 1997 and early October 1998, respectively. The germination phenologies of seeds of the same species and seed lots buried in soil were similar to those of seeds under leaf litter. High percentages of seeds of all four species germinated both under litter (78–96%) and beneath the soil surface (78–97%). These germination patterns correspond closely with the requirements for embryo growth and dormancy break in the four Lonicera species.