The Asteraceae with up to 30,000 species occurs on all continents except Antarctica and in all major vegetation zones on earth. Our primary aim was to consider cypselae dormancy-break and germination of Asteraceae in relation to ecology, vegetation zones and evolution. Cypselae are desiccation-tolerant and in various tribes, genera, species and life forms of Asteraceae are either non-dormant (ND) or have non-deep physiological dormancy (PD) at maturity. All six types of non-deep PD are found among the Asteraceae, and dormancy is broken by cold or warm stratification or by afterripening. Soil cypselae banks may be formed but mostly are short-lived. Much within-species variation in dormancy-break and germination has been found. Using data compiled for 1192 species in 373 genera and 35 tribes of Asteraceae, we considered ND and PD in relation to life form, vegetation zone and tribe. Senecioneae and Astereae had the best representation across the vegetation zones on earth. In evergreen and semi-evergreen rainforests, more species have ND than PD, but in all other vegetation zones, except alpine/high-latitude tundra (where ND and PD are equal), more species have PD than ND. Tribes in the basal and central grades and those in the Heliantheae Alliance have both ND and PD. The high diversity and lability of non-deep PD may have enhanced the rate of species diversification by promoting the survival of new species and/or species in new habitats that became available following globally disruptive events since the origin of the Asteraceae in the Late Cretaceous.

Germination of seeds of some summer annuals in Kentucky (eastern USA) in late-winter lead to the hypothesis that under present climate conditions the whole length of the winter cold stratification (CS) period is not required for dormancy-break of seeds of summer annuals with physiological dormancy (PD). We evaluated our data from germination phenology studies of 45 species (69 datasets) and buried-seed studies of 33 species (44 datasets). We determined time and temperature of germination after CS and percentage of the total number of hours of CS during winter (% of winter CS) seeds received prior to start of germination. In the phenology studies, mean temperature during the week of first germination for C3 and C4 species was 11.1 and 14.4°C, respectively, and % of winter CS was 80.8 and 87.4, respectively. In the buried-seed studies, % of CS for C3 and C4 species was 40.8 and 48.1, respectively, when they germinated to 25% at 20/10°C. For 32 of 33 species in the buried-seed studies, the minimum temperature at which seeds germinated decreased with increased CS; thus, seeds had Type 2 non-deep PD. The time of germination is controlled by a number of hours of CS, a decrease in minimum temperature at which seeds can germinate and a temperature increase in early spring. Seeds can germinate at relatively high temperatures as early as December and January, but they continue to be CS until spring. Temperature increases in eastern North America due to global warming are not likely to inhibit the germination of summer annuals with PD in spring.

This review provides a revised and expanded word-formula system of whole-seed primary dormancy classification that integrates the scheme of Nikolaeva with that of Baskin and Baskin. Notable changes include the following. (1) The number of named tiers (layers) in the classification hierarchy is increased from three to seven. (2) Formulae are provided for the known kinds of dormancy. (3) Seven subclasses of class morphological dormancy are designated: ‘dust seeds’ of mycoheterotrophs, holoparasites and autotrophs; diaspores of palms; and seeds with cryptogeal germination are new to the system. (4) Level non-deep physiological dormancy (PD) has been divided into two sublevels, each containing three types, and Type 6 is new to the system. (5) Subclass epicotyl PD with two levels, each with three types, has been added to class PD. (6) Level deep (regular) PD is divided into two types. (7) The simple and complex levels of class morphophysiological dormancy (MPD) have been expanded to 12 subclasses, 24 levels and 16 types. (8) Level non-deep simple epicotyl MPD with four types is added to the system. (9) Level deep simple regular epicotyl MPD is divided into four types. (10) Level deep simple double MPD is divided into two types. (11) Seeds with a water-impermeable seed coat in which the embryo-haustorium grows after germination (Canna) has been added to the class combinational dormancy. The hierarchical division of primary seed dormancy into many distinct categories highlights its great diversity and complexity at the whole-seed level, which can be expressed most accurately by dormancy formulae.

The genus Nolana (Solanaceae) comprises numerous species endemic to the coastal Atacama Desert of Chile and Peru of high ornamental potential and conservation value. The environments in which these species have evolved and are present today correspond to particular conditions in the midst of a hyper-arid habitat, so the study of their germination requirements and characterisation of seed dormancy becomes important in terms of conservation but also for ecological and evolutionary purposes. Different treatments were performed on mericarps of 12 species of Nolana: control (intact seeds imbibed in distilled water), scarification in funicular plug and distilled water and scarification in funicular plug and addition of GA3 (500 ppm); their permeability to water was also tested. It was determined that the species did not present physical dormancy, as had been previously reported, but rather physiological dormancy (PD). Germination results after treatments were not homogeneous among all 12 species, indicating differences in their dormancy levels. Also, the important role of the endosperm in the prevention of germination for the studied Nolana species was highlighted. Regarding the relationship between the level of PD (expressed as the percentage of germination for the most successful treatment) and the latitudinal distribution of the species or their phylogenetic closeness, it was determined that, for the studied species, their proximity in terms of clades was more relevant than their latitudinal distribution.

For physiologically dormant (PD) species in fire-prone environments, dormancy can be both complex due to the interaction between fire and seasonal cues, and extremely deep due to long intervals between recruitment events. Due to this complexity, there are knowledge gaps particularly surrounding the dormancy depth and cues of long-lived perennial PD species. This can be problematic for both in situ and ex situ species management. We used germination experiments that tested seasonal temperature, smoke, dark and heat for 18 PD shrub species distributed across temperate fire-prone Australia and assessed how germination was correlated with environmental factors associated with their home environments. We found extremely high levels of dormancy, with only eight species germinating above 10% and three species producing no germination at all. Seven of these eight species had quite specific seasonal temperature requirements and/or very strong responses to smoke cues. The maximum germination for each species was positively correlated with the mean temperature of the source population but negatively correlated with rainfall seasonality and driest months. The strong dependence on a smoke cue for some of the study species, along with examples from other studies, provides evidence that an obligate smoke response could be a fire-adapted germination cue. Germination response correlated with rainfall season of the source populations is a pattern which has often been assumed but little comparative data across sites with different rainfall seasonality exists. Further investigation of a broader range of species from different rainfall season environments would help to elucidate this knowledge gap.

In Chile, two quinoa ecotypes are grown: salares, also present in the highlands of Bolivia, and coastal, in central and southern areas of the country, at sea level. Genotypes from the coastal ecotype have characteristics that differentiate them from the most popular quinoa genotypes grown in the Andean Region of South America. The objectives of this study were: (1) to determine the cardinal temperatures for seed germination in quinoa genotypes from coastal and salares ecotypes cultivated in Chile, and (2) to study the presence of physiological dormancy (PD) in these genotypes. Seed germination from nine quinoa genotypes, two from salares and seven from coastal ecotypes, was evaluated in a gradient of temperatures between 11 and 42°C. Germination was also evaluated at 20°C at 0, 7 and 15 months from harvest. Results showed that seed from the nine genotypes germinated at their maximum percentage between 11 and 35°C. However, their faster germination occurred between 25 and 35°C. There was a significant difference between optimum temperature for germination between genotypes from coastal (28°C) and salares (30°C). An increase in germination rates after 7 months of storage suggested the presence of a non-deep PD in seeds from coastal ecotype, which may be useful to improve pre-harvest sprouting resistance in quinoa breeding programmes.

Only a few studies have been performed on seed germination of perennial ephemeral species native to the cold deserts of central Asia. We hypothesized that seeds of the cold desert perennial ephemeral Leontice incerta exhibit versatility in the timing of germination, that is, having the capacity to germinate at any time in summer, autumn and next spring. At dispersal in late May, only about 30% of the seeds could germinate; thus, a high percentage of the seeds was dormant. Seeds had a fully developed embryo, and dry storage, cold stratification, warm stratification and gibberellin promoted germination; we concluded that they have non-deep physiological dormancy. Seeds buried under natural conditions during summer germinated to 57–86% in autumn (late October) when exhumed and incubated at 5/2–25/15°C. However, seeds were sown in soil exposed to natural temperature and (low) precipitation did not germinate until next spring when the soil was moist. Thus, like various cold desert annuals, seeds of the perennial L. incerta can germinate in summer, autumn and next spring, depending on the availability of soil moisture (rainfall). Rainfall in cold deserts can play an important role in shaping seed germination traits of desert plants.

Fire is a frequent disturbance in the Cerrado and is one of the major factors affecting vegetation structure and diversity. Fire events open gaps within the herbaceous layer and increase temperature fluctuation in the soil surface. In addition to being an important environmental filter for germination, fire is a germination cue for species with physiological dormancy. This study aimed to evaluate the germination of native grasses, using daily temperature fluctuations and heat shock to overcome physiological dormancy in native grasses. We also evaluated seed longevity after dispersal for some species. We conducted the daily temperature fluctuation experiments on seeds of ten native grass species, which were collected and then placed in germination chambers simulating thermal fluctuation throughout the day (19–55°C). We also subjected seeds to different heat-shock treatments: 100°C for 1 min, 100°C for 3 min and 200°C for 1 min. To determine seed longevity, we stored seeds for 6 and 12 months after collection and then set them to germinate (27°C, 12/12 h light). Non-germinated seeds from all experiments were tested for viability. Most species had low longevity and germination percentages. Those that had physiological dormancy were stimulated to germinate when exposed to temperature fluctuations. One species resisted temperatures up to 200°C. For all other species, neither treatment affected germination percentages. Our results indicate the importance of these environmental filters for seedling recruitment of these species, considering the low longevity and the presence of physiological dormancy.

In seasonal climates, germination timing is mainly controlled by temperature, especially in species with physiological seed dormancy. The germination response to temperature may, however, vary among populations across the distribution range of species. Understanding how populations along climate gradients vary in their sensitivity to temperature is important for determining their vulnerability to climate variability and change. Here, we investigated the germination response of two Erica species with physiological seed dormancy (E. australis and E. umbellata) to changes in temperature throughout the seasons (simulated autumn through to spring) and to the local climate in six localities across a latitudinal gradient in western Iberia. Effects were studied with and without exposing the seeds to a heat shock. The local climate of seed provenance emerged as a key factor in modifying the germination sensitivity to germination temperature and their variation through the seasons. Although each species showed idiosyncratic germination responses to temperature treatments and across the gradient, germination of both species was sensitive to warmer temperatures and to a heat shock. Both showed similar seasonal germination patterns: as we moved from south to north, populations tended to have a larger germination peak in spring, which was greater at colder temperatures. We conclude that rising temperatures associated with climate change will affect these species, particularly at their northern ranges, where many seeds will remain dormant during warmer winters. Arguably, models aiming at assessing climate change impacts in these species need to include such variability across latitude.

This study investigated the induction and loss of dormancy in oilseed rape (Brassica napus). Twenty genotypes were preliminary screened; from these, two genotypes, RGS003 and Hayola 308, which possess high potential for dormancy induction (HSD) and medium potential to induce secondary dormancy (MSD), were selected. The stratification of seeds at alternating temperatures of 5–30°C (in dark) significantly relieved secondary dormancy, but dormancy was not fully released. The ψb(50) values were −1.05 and −1.06 MPa for the MSD and the HSD before dormancy induction. After inducing dormancy, the ψb(50) values for the MSD and the HSD were increased to −0.59 and −0.01 on day 0 stratification at 20°C. The hydrothermal time (θHT) value was low for one-day stratification for HSD in comparison with other stratification treatments. Water stress can induce dormancy (if the seeds have the genetic potential for secondary dormancy) and warm stratification (in dark) can only reduce the intensity of dormancy. The seeds with a high potential of dormancy induction can overcome dormancy at alternating temperatures and in the presence of light. It can, therefore, be concluded that a portion of seeds can enter the cycle of dormancy ↔ non-dormancy. The secondary dormant seeds of B. napus cannot become non-dormant in darkness, but the level of dormancy may change from maximum (after water stress) to minimum (after warm stratification). It seems that the dormancy imposed by the conditions of deep burial (darkness in combination with water stress and more constant temperatures) might be more important to seed persistence than secondary dormancy induction and release. The dormancy cycle is an important pre-requisite in order to sense the depth of burial and the best time for seed germination.

The broad aim of this work was to study intraspecific variation of seed germination in Phillyrea angustifolia L. (Oleaceae), a species with a hard (water-permeable) endocarp. Germination of seeds from six different wild populations was correlated with traits related either to seed morphology or to environmental parameters. Germination of naked seeds (seeds without endocarp) at the optimum germination conditions was similar among populations and individuals, but great differences could be detected regarding the germination of seeds with endocarp both at inter- and intra-populational levels. Differences among populations could be related to climatic parameters and to morphometric variables of seeds with endocarp. A higher germination was associated with populations growing in habitats with more severe summer (higher temperature, lower precipitation and a longer drought period) and producing elongated seeds (lower Feret ratio and roundness). Moreover, seeds from eight different individuals within a population were tested independently, and great differences regarding the germination of seeds with endocarp could be detected among individuals. Our results suggest that the morphological variation found in P. angustifolia endocarp is both under strong maternal genetic control as well as influenced by environmental factors, as indicated by the high variability among individuals within one population and the significant correlation between climate variables and seed germination among populations. Finally, it is emphasized that standardization of plant propagation protocols should take into account the degree of intraspecific variation of Mediterranean species.

Common teasel is an invasive and widespread weed in Argentina. Germination experiments were carried out from 2011 to 2014 to determine the effect of various environmental factors on germination and emergence. Germination of recently dispersed seeds was 12% in darkness at constant temperature. In contrast, seed exposure to light and alternating temperatures enhanced germination to 95%. The requirement of light and alternating temperatures suggest that common teasel has physiological dormancy. Several experiments were carried out to determine whether (1) seed responses to light and alternating temperatures have a hormonal basis, (2) seed coats can suppress germination, and (3) time and thermal conditions during seed storage reduce light and alternating temperature requirements. Germination was reduced (≤ 13%) by a gibberellin synthesis inhibitor. In contrast, the presence of gibberellins and an abscisic acid synthesis inhibitor increased germination to 95 and 38%, respectively. Results suggest that a higher ratio among gibberellins and abscisic acid (GA/ABA) leads to a break in dormancy. Germination was 100% when embryos were excised, suggesting that seed coats may suppress germination by mechanical restriction. Likewise, germination was enhanced by hydrogen peroxide (70%). This compound is known to increase GA/ABA ratio in agreement with a hormonal control of dormancy proposed for common teasel. An increment of storage time reduces light and alternating temperature requirements, allowing seeds to germinate in darkness. Taking these results together confirms that common teasel has physiological dormancy. Seedling emergence was progressively reduced from 70 to 8% by increased burial depth from 0 to 3 cm. Information from these experiments may facilitate development of effective management for common teasel.

This article briefly reviews how long weed seeds can live in the soil and what happens to them during burial. Freshly matured seeds of many weeds are water-permeable, but those of others are water-impermeable. Water-permeable seeds may have morphological, physiological, or morphophysiological dormancy, with physiological dormancy (PD) being the one most commonly found in buried weed seeds. Further, nondeep PD is very common in buried weed seeds, and many of them exhibit annual dormancy cycles in response to seasonal temperature changes. The time of year when seeds are nondormant varies with the species, i.e., autumn, spring, or spring to summer. A light requirement for germination plays an important role in preventing nondormant seeds from germinating in the soil. To germinate, soil disturbance that exposes seeds to light must occur at a time of year when seeds are nondormant. Buried seeds of some species come out of dormancy and remain nondormant regardless of seasonal changes in environmental conditions; however, a light requirement for germination prevents them from germinating in the soil. Water-impermeable seeds have either physical dormancy (PY) or a combination of PY and PD, with PY being the most common, e.g., in members of the Fabaceae and Malvaceae. Seeds with PY have a water gap in the seed coat that opens in response to an environmental signal, thereby allowing water to enter. When disturbance brings seeds to the soil surface, temperatures that are higher than those in the soil can cause the water gap to open. Consequently, the water gap indirectly serves as a depth sensor. A challenge for the future is to use information about buried weed seeds to better manage weeds in crops, and modeling is an important step in that direction.

The fruit plus accessory parts, and thus not only the fruit or seed, is the diaspore in some species of flowering plants. Atriplex centralasiatica, a summer annual salt-secreting cold-desert halophyte of central Asia that produces flat and humped diaspores, is such a species. The dispersal/germination units of this diaspore-heteromorphic species are fruits enclosed in persistent ‘bracteoles’. Germination of black and brown fruits (i.e. ‘bracteoles’ removed) of this species has been studied in some detail previously, but little attention has been given to the germination biology of the intact diaspores. The aim of this study was to compare the germination biology of the intact diaspores of A. centralasiatica, and their fates in the field during 12 months on the soil surface and buried at 5-cm depth. Fresh flat diaspores germinated to 42.7–51.3% in light (12-h photoperiod) and to 16.0–59.3% in constant dark, over a temperature range of 15–20/30°C, while no freshly matured humped diaspores did so under any temperature/light regime. Neither gibberellin (GA3) nor potassium nitrate (KNO3) had an effect on germination of either diaspore. Both diaspores readily imbibed water, and removal of ‘bracteoles’ released most of the dormancy. Thus, the ‘bracteoles’ are primarily responsible for diaspore dormancy in A. centralasiatica. Humped diaspores persisted for a longer period in the soil than flat ones. The germination requirements of intact natural dispersal/germination units of A. centralasiatica differ from those previously reported for fruits of this species.

To increase our knowledge of the diversity of seed dormancy and germination in Rubiaceae, we investigated seed desiccation sensitivity and germination of three Psychotria species. Seeds of P. gardneri, P. nigra and P. zeylanica germinated to high percentages at <15% seed moisture content. Intact seeds of P. zeylanica and P. nigra imbibed water and thus do not have physical dormancy. More than 50% of the seeds of P. zeylanica, P. nigra and P. gardneri took 33, 53 and 110 d, respectively, at 25°C for the radicle to emerge, and embryo growth occurred before and after radicle emergence. Thus, seeds have morphophysiological dormancy. Shoot emergence of P. nigra and P. zeylanica seeds was delayed 50 and 80 d after radical emergence, respectively; thus, seeds have epicotyl morphophysiological dormancy (eMPD). This is the first report of eMPD in Rubiaceae. Since warm stratification promoted both radicle and shoot emergence in seeds of P. zeylanica and P. nigra, the level of eMPD is non-deep simple. Hence, dormancy of the studied Psychotria spp. can be described as C1bBb (radicle)–C1bBb (epicotyl), i.e. the embryo is underdeveloped and grows prior to radicle emergence and epicotyl emergence under warm temperatures (Bb), and both the radicle and epicotyl have non-deep simple physiological dormancy broken by warm temperatures (C1b). In two Psychotria species studied in detail, radicle emergence occurs at the beginning of the rainy season and plumule emergence at the peak rainy season when conditions are most favourable for rapid seedling development.

The dispersal unit of many species of Brassicaceae is an indehiscent fruit, but relatively few studies have tested the effect of the pericarp on seed germination in this family. Our aim was to determine the effect of the pericarp on seed dormancy in six species of Brassicaceae native to the cold desert of north-west China. Intact dispersal units and isolated seeds of Chorispora sibirica, Euclidium syriacum, Goldbachia laevigata, Spirorrhynchus sabulosus, Sterigmostemum fuhaiense and Tauscheria lasiocarpa were stored dry at ambient laboratory conditions for 0–12 months and tested for germination in light and in dark at 5/2, 15/2 and 30/15°C. The amount of water absorbed by fruits and by seeds within the fruits was determined. For four species, intact fruits, isolated seeds and isolated seeds plus the removed pericarps were used to test for the mechanical versus possible chemical role of the pericarp in seed dormancy. Fresh isolated seeds, which have a fully developed embryo, germinated to lower percentages and rates than afterripened seeds. Thus, seeds have non-deep physiological dormancy. The pericarp significantly reduced germination, but inhibition was not due to lack of water uptake by seeds or to chemical inhibitors. Afterripened seeds of the six species germinated to 0–50% inside the fruits. We conclude that the pericarp plays a dominant role in seed dormancy of the six study species, and it does so by mechanically restricting embryo growth. Thus, the pericarp has the potential to spread germination over an extended period of time.

Determining the phylogenetic and biogeographic distribution of physical dormancy remains a major challenge in germination ecology. Here, our goal was to describe a novel water-impermeable seed coat mechanism causing physical dormancy (PY) in the seeds of Chaetostoma armatum (Melastomataceae). Although seed coat permeability tests indicated a significant increase in seed weight after soaking in distilled water, anatomical and dye-tracking analyses showed that both water and dyes penetrated the seed coat but not the embryo, which remained in a dry state. The water and dye penetrated the lumen of the exotestal cells, which have a thin outer periclinal face and thickened secondary walls with U-shaped phenolic compounds. Because of this structure, water and dye do not penetrate the inner periclinal face of the exotestal cells, indicating PY. Puncturing the seeds increased germination more than tenfold compared to that of the control, but GA3 did not increase germination further. A significant fraction of the seeds did not germinate after puncturing, indicating that embryos are also physiologically dormant (PD). This paper constitutes the first report of the water-impermeable seed coat in the Myrtales and the first report of physiophysical (PD+PY) dormancy in a shrub from a tropical montane area.

Seeds of Sisymbrium officinale display physiological dormancy and require nitrate to germinate. Rupture of the testa precedes radicle protrusion through the endosperm (germination sensu stricto). While both endosperm rupture and testa rupture (TR) required nitrate, endosperm rupture was fully inhibited by abscisic acid (ABA) but TR was not inhibited. The gibberellic acid (GA)-synthesis inhibitor paclobutrazol prevented TR, which was reverted by exogenous GA4 but not by nitrate. The orientation of TR was transverse, which prompted the question whether seeds elongate prior to radicle protrusion, concurrent with an increase in water content. Between 9 h and 1 d no increase in length or water content was observed. During incubation in ABA the length of imbibed seeds without TR did not increase between 1 and 5 d, whereas nitrate added to ABA induced TR and a 94% increase in length. At the same time the water content of seeds without TR increased by 18%, while the water content of seeds with TR increased by 38%. Length and water content were correlated in a single-seed analysis for seeds with TR, but not for seeds without TR. Increased length was also observed in Arabidopsis seeds with nitrate-induced TR. These results indicate that prior to endosperm rupture dormancy release by nitrate is accompanied by TR, seed elongation and an increase in water content. A new multiphasic model is proposed for the imbibition curve, where the second phase of the classical triphasic curve is split into three sub-phases, of which phases IIB and IIC are associated with TR.

Seeds of legumes are generally considered to have physical dormancy and to be orthodox, but most seed biologists are unaware of the various kinds and combinations of dormancy and storage behaviour in seeds of this family. The aim of our study was to document the dormancy and storage behaviour of seeds of 100 native and introduced tropical Fabaceae species in Sri Lanka and classify them into germination/storage behaviour categories. Moisture content (MC) was < 16% for fresh seeds of 94 species and >29% for those of six. Seeds of these six species had low tolerance for desiccation and for low temperatures. Thus, seeds of six species are non-orthodox and 94 species orthodox. Nine of the 100 species were non-dormant, and 2, 3 and 86 had physiological, physiological epicotyl and physical dormancy, respectively. Six germination/storage behaviour categories were identified among the 100 species. However, as in extratropical regions of the world, orthodox storage behaviour and physical dormancy are characteristic of seeds of the majority of species of Fabaceae in tropical Sri Lanka.

Although seed dormancy of temperate legumes is well understood, less is known about it in species that grow in subalpine/alpine areas. This study investigated dormancy and germination of four Vicia species from the Tibetan Plateau. Fresh seeds of V. sativa were permeable to water, whereas those of V. angustifolia, V. amoena and V. unijuga had physical dormancy (PY). One year of dry storage increased the proportion of impermeable seeds in V. angustifolia, but showed no effect on seed coat permeability in V. amoena or V. unijuga. Seeds of all four species also had non-deep physiological dormancy (PD), which was especially apparent in the two annuals at a high germination temperature (20°C). After 1 year of storage, PD had been lost. The hydrotime model showed that fresh seeds obtained a significantly higher median water potential [Ψb(50)] than stored seeds, implying that PD prevents germination in winter for seeds dispersed without PY when water availability is limited. After 6 months on the soil surface in the field, a high proportion of permeable seeds remained ungerminated, further suggesting that PD plays a key role in preventing germination after dispersal. Addition of fluridone, an inhibitor of abscisic acid (ABA) biosynthesis, evened-out the differences in germination between fresh and stored seeds, which points to the key role of ABA biosynthesis in maintaining dormancy. Further, fresh seeds were more sensitive to exogenous ABA than stored seeds, indicating that storage decreased embryo sensitivity to ABA. On the other hand, the gibberellic acid GA3 increased germination rate, which implies that embryo sensitivity to GA is also involved in seed dormancy regulation. This study showed that PY, PD or their combination (PY+PD) plays a key role in timing germination after dispersal, and that different intensities of dormancy occur among these four Vicia species from the Tibetan Plateau.