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.     

Effects of environmental stress and nutlet morph on proportion and within-flower number-combination of morphs produced by the fruit-dimorphic species Lappula duplicicarpa (Boraginaceae)

Phenotypic plasticity in species with heteromorphic diaspores is considered to be a plastic bet-hedging strategy in environmentally fluctuating habitats. The primary aim of this study was to determine the influence of level of abiotic (soil moisture and nutrient availability) and biotic (pure and mixed density) factors on the long-glochid (LN; high dispersal, low dormancy)/short-glochid (SN; low dispersal, high dormancy) nutlet ratio and on nutlet combination of the fruit-dimorphic cold desert winter annual/spring ephemeral Lappula duplicicarpa derived from different nutlet morphs. We hypothesized that stress would decrease the LN: SN ratio. All levels of the four treatments resulted in significant variation in the LN: SN ratio and in the proportion of each nutlet combination. Reproductive biomass varied less than plant mass in plants from both morphs. In favorable environments, the LN: SN ratio and number of SN–3LN and 4LN combinations were relatively high, while in unfavorable environments they were relatively low, thus supporting our hypothesis. Relative allocation to reproductive biomass was significantly negatively correlated with allocation to vegetative biomass. The downward shift that occurred in the LN: SN diaspore ratio in response to stressful growth conditions in Lappula duplicicarpa is assumed to be a bet-hedging strategy.     

Phenotypic plasticity and bet‐hedging in a heterocarpic winter annual/spring ephemeral cold desert species of Brassicaceae

Phenotypic plasticity can play an important role in colonization and survival of plants in an environmentally fluctuating habitat. The primary aim of this study was to determine the influence of level of abiotic (soil moisture and nutrient availability) and biotic (density and herbivory) factors on phenotypic plasticity in the number and proportional mass allocation to the heteromorphic dehiscent upper (high dispersal, low dormancy) and indehiscent lower (low dispersal, high dormancy) siliques of individuals of the cold desert winter annual/spring ephemeral Diptychocarpus strictus derived from different seed morphs. Plants produced from seeds sown in an experimental garden were subjected to different levels of soil moisture, nutrient supply, density and simulated herbivory. Mass allocated to vegetative and reproductive components was measured and number of upper and lower siliques counted. Except for number ratio of upper: lower siliques under nutrient supply, levels of the four treatments resulted in significant variation in total plant mass, reproductive mass, number of siliques (upper and lower), number and mass of each silique morph, individual seed mass, upper/lower silique ratio and mass allocation to each organ in an individual. In favorable environments, the upper/lower silique ratio was relatively high, while in unfavorable environments it was relatively low. The relative allocation to upper and lower siliques was significantly negatively correlated, suggesting that allocation to upper siliques in good growth conditions occurred at the expense of allocation to lower siliques. This appears to be the first report on the effect of herbivory on diaspore morph ratio in heterocarpic plants and one of only a very few on the effect of density on morph ratio in this group. In D. strictus, stressful vs nonstressful growth conditions caused a shift in the ratio of heteromorphic diaspores, which themselves are assumed to be bet‐hedging.     

异子蓬二型种子的萌发与休眠特性及其生态适应

异子蓬为中亚荒漠的特有种,其果实为胞果,花被片在结果时宿存包被果实形成浆果状的散布单位.对异子蓬的散布单位、果实形态进行比较观察,并在5/15℃、5/25℃、15/25 ℃(暗/光=12h/12h)3个温变周期下进行种子萌发实验,结果表明:(1)异子蓬的散布单位绿色、浆果状,有大小两种形态;大小两种散布单位内包被的果实在形状、大小、颜色上存在着明显差异.(2)大的散布单位内包被的果实圆形,扁平,褐色,无光泽,直径(2.64±0.03) mm,重量(3.68±0.04) mg;小的散布单位内包被的果实双凸镜形,黑色,有光泽,直径(2.40±0.04) mm,重量(2.86±0.06) mg.(3)褐色种子在3个温变周期下的萌发率都在81%以上,萌发速度快,而黑色种子萌发率低,萌发慢,具有休眠现象.(4)划破种皮及低温层积处理可显著提高黑色种子在3个温变周期下的萌发率,延长储存时间也可有效地促进黑色种子的萌发,表明黑色种子处于非深度生理休眠状态.异子蓬产生的两种不同类型的种子及其在时间和空间上的萌发差异对荒漠异质环境具有重要的适应意义.     

Comparison of germination and seed bank dynamics of dimorphic seeds of the cold desert halophyte Suaeda corniculata subsp. mongolica

Background and Aims

Differences in dormancy and germination requirements have been documented in heteromorphic seeds of many species, but it is unknown how this difference contributes to maintenance and regeneration of populations. The primary aim of this study was to compare the seed bank dynamics, including dormancy cycling, of the two seed morphs (black and brown) of the cold desert halophyte Suaeda corniculata and, if differences were found, to determine their influence on regeneration of the species.

Method

Seeds of the two seed morphs were buried, exhumed and tested monthly for 24 months over a range of temperatures and salinities, and germination recovery and viability were determined after exposure to salinity and water stress. Seedling emergence and dynamics of the soil seed bank were also investigated for the two morphs.

Key Results

Black seeds had an annual dormancy/non-dormancy cycle, while brown seeds, which were non-dormant at maturity, remained non-dormant. Black seeds also exhibited an annual cycle in sensitivity of germination to salinity. Seedlings derived from black seeds emerged in July and August and those from brown seeds in May. Seedlings were recruited from 2·6 % of the black seeds and from 2·8 % of the brown seeds in the soil, and only 0·5 % and 0·4 % of the total number of black and brown seeds in the soil, respectively, gave rise to seedlings that survived to produce seeds. Salinity and water stress induced dormancy in black seeds and decreased viability of brown seeds. Brown seeds formed only a transient soil seed bank and black seeds a persistent seed bank.

Conclusions

The presence of a dormancy cycle in black but not in brown seeds of S. corniculata and differences in germination requirements of the two morphs cause them to differ in their germination dynamics. The study contributes to our limited knowledge of dormancy cycling and seed bank formation in species producing heteromorphic seeds.     

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.

     

盐生植物角果碱蓬种子二型性对环境的适应策略

角果碱蓬(Suaeda corniculata)是藜科一年生盐生植物, 在我国分布于北方盐碱滩涂和盐碱荒漠地区。角果碱蓬具有棕色和黑色两种异型体种子(简称棕色和黑色种子)。对采自内蒙古鄂托克前旗盐渍化生境的角果碱蓬二型种子的形态、休眠和萌发特性开展对比研究, 测定了二型种子休眠和萌发行为对温度、光照和盐分(NaCl)的响应, 以揭示盐生植物异型种子对温带盐漠生境的适应对策。结果表明: (1)二型性种子在大小、种皮特性和结实比例方面有显著差异。与黑色种子相比, 棕色种子个体较大, 种皮透水性强。黑色种子与棕色种子的结实比例约为5.6 : 1。(2)新成熟的棕色种子的萌发对各温度梯度和光照条件不敏感, 萌发率较高(84%-100%); 而新成熟的黑色种子萌发率较低(8%-78%), 萌发对光照敏感。(3)黑色种子具有浅度生理休眠, 种皮划破、赤霉素处理和低温层积均可有效地提高种子的萌发率。(4)二型种子萌发对土壤盐分的胁迫具有不同的响应。与黑色种子相比, 棕色种子对盐分胁迫不敏感, 在较高的盐分浓度下仍有较高的萌发率, 低温层积处理能够降低黑色种子对盐胁迫的敏感性, 有效地提高种子的初始萌发率、萌发恢复率和最终萌发率。角果碱蓬二型种子不同的形态、休眠和萌发特性, 提高了该物种在高度异质性生境中的适合度, 对种群成功地适应温带盐漠环境具有重要的意义。     

Germination of dimorphic seeds of the desert annual halophyte Suaeda aralocaspica (Chenopodiaceae), a C4 plant without Kranz anatomy

Background and Aims

Suaeda aralocaspica is a C₄ summer annual halophyte without Kranz anatomy that is restricted to the deserts of central Asia. It produces two distinct types of seeds that differ in colour, shape and size. The primary aims of the present study were to compare the dormancy and germination characteristics of dimorphic seeds of S. aralocaspica and to develop a conceptual model of their dynamics.

Methods

Temperatures simulating those in the natural habitat of S. aralocaspica were used to test for primary dormancy and germination behaviour of fresh brown and black seeds. The effects of cold stratification, gibberellic acid, seed coat scarification, seed coat removal and dry storage on dormancy breaking were tested in black seeds. Germination percentage and recovery responses of brown seeds, non-treated black seeds and 8-week cold-stratified black seeds to salt stress were tested.

Key Results

Brown seeds were non-dormant, whereas black seeds had non-deep Type 2 physiological dormancy (PD). Germination percentage and rate of germination of brown seeds and of variously pretreated black seeds were significantly higher than those of non-pretreated black seeds. Exposure of seeds to various salinities had significant effects on germination, germination recovery and induction into secondary dormancy. A conceptual model is presented that ties these results together and puts them into an ecological context.

Conclusions

The two seed morphs of S. aralocaspica exhibit distinct differences in dormancy and germination characteristics. Suaeda aralocaspica is the first cold desert halophyte for which non-deep Type 2 PD has been documented.Key words: Borszczowia, cold desert halophyte, physiological seed dormancy, seed germination, Suaeda     

Variation of seed heteromorphism in Chenopodium album and the effect of salinity stress on the descendants

Background and Aims

Chenopodium album is well-known as a serious weed and is a salt-tolerant species inhabiting semi-arid and light-saline environments in Xinjiang, China. It produces large amounts of heteromorphic (black and brown) seeds. The primary aims of the present study were to compare the germination characteristics of heteromorphic seeds, the diversity of plant growth and seed proliferation pattern of the resulting plants, and the correlation between NaCl stress and variation of seed heteromorphism.

Methods

The phenotypic characters of heteromorphic seeds, e.g. seed morphology, seed mass and total seed protein were determined. The effects of dry storage at room temperature on dormancy behaviour, the germination response of seeds to salinity stress, and the effect of salinity on growth and seed proliferation with plants derived from different seed types were investigated.

Key Results

Black and brown seeds differed in seed morphology, mass, total seed protein, dormancy behaviour and salinity tolerance. Brown seeds were large, non-dormant and more salt tolerant, and could germinate rapidly to a high percentage in a wider range of environments; black seeds were salt-sensitive, and a large proportion of seeds were dormant. These characteristics varied between two populations. There was little difference in growth characteristics and seed output of plants produced from the two seed morphs except when plants were subjected to high salinity stress. Plants that suffered higher salinity stress produced more brown (salt-tolerant) seeds.

Conclusions

The two seed morphs of C. album exhibited distinct diversity in germination characteristics. There was a significant difference in plant development and seed proliferation pattern from the two types of seeds only when the parent plants were treated with high salinity. In addition, seed heteromorphism of C. album varied between the two populations, and such variation may be attributed, at least in part, to the salinity.     

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.     

新疆干旱区植物藜的种子异型性及其萌发机理

新疆干旱区分布的植物藜(Chenopodium album)的种子有黑色和褐色两种类型。对藜的异型性种子从形态结构、不同环境因素及激素或化学物质对萌发的影响以及同工酶谱等方面进行了研究,并对其萌发及适应异质环境的机理进行了讨论。结果表明:(1)藜的异型性种子在形态结构、萌发休眠特性等方面都存在明显差异:黑色种子种皮厚且硬,休眠,萌发慢,萌发率低;褐色种子种皮薄而软,不休眠,萌发快且萌发率高;(2)黑色种子的休眠可通过切除胚根外缘种皮得以完全解除;(3)赤霉素、乙烯利对黑色种子的萌发无明显促进作用;KNO3可较显著促进黑色种子的萌发;协同使用乙烯利和KNO3时,可显著提高黑种子萌发率,完全打破休眠;(4)黑色种子和褐色种子的酯酶、过氧化物酶及过氧化氢酶同工酶谱带存在差异;(5)黑色种子的萌发需要光照,而褐色种子则对光不敏感;低温贮藏对二者的萌发均无显著影响,尽管黑色种子的萌发率有波动。研究结果初步显示黑色种子的休眠是内源(胚)和外源(种皮)因素共同所致。藜的种子异型性及其萌发机理的形成是其对新疆干旱区异质化环境的高度适应。     

异子蓬二型性种子的种皮结构及离子含量差异

异子蓬(Borszczowia aralocaspica)是中亚荒漠区系特有的一年生盐生植物,其种子具有二型性。通过检测异子蓬二型性种子中的生理生化物质以及观察种皮的结构,比较异子蓬二型性种子中营养物质、养分全量以及种皮结构的差异,结果表明:(1)异子蓬的棕色种子只有一层薄种皮,而黑色种子不但具有一层薄的内种皮还有一层坚硬致密的外种皮;(2)棕色种子的质量大、质量频数分布较为集中,并且吸水迅速,在吸水率较低时就能够大量萌发;(3)棕色种子的可溶性糖、可溶性蛋白质、全氮、全磷以及K+、Na+、K+/Na+和Mg2+/Na+都显著高于黑色种子。说明在繁殖子代、产生二型性种子时,异子蓬对棕色种子和黑色种子之间的繁殖投入存在着巨大的差异,这可能是导致异子蓬二型性种子产生差异性萌发机制的重要原因。     

喀什霸王的结实和种子萌发特性

喀什霸王(Zygophyllum kaschgaricum)是生长于中国新疆南部荒漠环境的稀有种及二级保护植物。当前, 该物种在自然种群中呈分散式及片段化分布, 且种群密度低, 种群老龄化较严重。因此, 为了了解该物种种子萌发特性及其对荒漠环境的响应, 该文采用室内控制实验方法, 对该物种的自然坐果率、结籽率、种子吸水特性、种子休眠和萌发特性及对干旱胁迫的响应进行了比较研究。结果表明: 喀什霸王在自然种群中的坐果率及结籽率较低, 且种子的败育率较高。不同干藏时间种子的吸水速率间存在显著差异; 随着干藏时间的延长, 种子的吸水率逐步增强。刚成熟的种子在不同温度及光周期下均可萌发; 其中高温(10/20 ℃, 20/30 ℃)及黑暗条件下的萌发率比低温(10/5 ℃, 5/2 ℃)及光照条件下的萌发率高。不同干藏时间的种子在不同浓度赤霉素(GA₃)下的萌发率均较高; 但低温储藏时间对该物种种子的打破休眠及萌发率没有促进作用。以上结果说明该物种存在非深度生理休眠; 而干藏时间、高温且黑暗及高浓度(50 mmoloL ^-1) GA₃是打破休眠及促进种子萌发的最合适条件。高温条件下的干旱胁迫对喀什霸王种子萌发具有抑制作用; 春季和秋季降水量决定种子的萌发率。总之, 喀什霸王种子在物候上表现出的春秋季萌动及非深度生理休眠以提高幼苗存活力及保障种群稳定性, 是一种对新疆南部干旱及高温胁迫荒漠环境的适应策略。     

Reproductive patterns in annual legume species on an aridity gradient

Reproductive patterns are analysed in annual legumes of west Asia, and their relationships to increasing aridity determined by multivariate analysis. Dormancy, seed size, dispersal and fecundity are shown to be partially substitutable in terms of their effect on survival and population growth.The range of patterns show greatest diversity under mesic conditions in coastal, mediterranean areas with high winter rainfall, low incidence of frost and long growing season. Increasing aridity leads to increasing reproductive homogeneity, in which a subset of patterns-those composed of high levels of seed dormancy, high seed to pod ratios, restricted dispersal capabilities and early flowering-become predominant. These findings corroborate earlier theoretical and empirical evidence concerning desert annuals.The majority of widespread species are shown to possess arid-type patterns. This exerts considerable influence on both the type and degree of ecotypic differentiation within species. The only feature that responds consistently to climatic change is flowering time, particularly among species characterized by high seed dormancy. In the few widespread species that do not display arid-type characteristics, ecotypic differentiation appears more frequently in a larger suite of traits.The unequal environmental demands made on species showing different levels of dormancy (e.g. variation in length of growing season from year to year and place to place) leads to strong asymmetries governing the relationships between reproduction and ecological amplitude. In this respect arid-type strategists with high seed dormancy appear to have greater chances of expansion than others. This has implications when choosing pasture legumes to improve mediterranean grasslands.     

Delayed germination and dispersal in desert annuals: Escape in space and time

Summary A model is developed to consider the interplay between dispersibility and delayed germination in desert annuals. The model explores the effect of low levels of dispersal, considered realistic for annual plants, on optimal germination fraction. The model also demonstrates the effect of the amount and accuracy of predictive (responsive to the environment) dormancy on the optimal innate germination fraction (not responsive to environmental conditions).Optimal germination fraction is found to be very sensitive to changes in despersibility especially at the limited dispersibilities that are realistic for annual plants. As dispersibility increases, optimal germination fraction increases. If plants make two kinds of seeds with differing despersibility, reproduction is maximized if the low dispersal seeds have delayed germination and the high dispersal seeds have quick germination. If dormancy mechanisms permit seeds to germinate when environmental conditions allow successful maturation, and remain dormant when environmental conditions do not permit successful maturation, what fraction of seeds should remain dormant under predicted good conditions as a hedge against inaccurate prediction of the environment? If environmental cues that break dormancy are uncorrelated with environmental conditions that permit successful maturation, predictive dormancy has little or no effect on the optimal innate germination fraction. When predictive dormancy lowers the probability of germinating when environmental conditions preclude successful maturation, the optimal innate germination fraction increases with increasing germination control by predictive dormancy. With a moderate degree of germination control by predictive dormancy, the optimal innate dormancy is still sensitive to changes in dispersal in the low dispersal ranges characteristic of annual plants.Evidence is presented from plant species that have both dispersal and germination dimorphisms to support the predicted correlation of high germination fractions with high dispersal.     

种子异型性及其生态意义的研究进展

种子异型性是指同一植株产生不同形状或行为种子的现象。根据异型种子在植株上的生长位置, 种子异型性可划分为地上下结实性和地上种子异型性两类。此现象已在26科129属292种被子植物中报道。异型性种子植物主要分布于干旱半干旱区、荒漠和盐渍土地区等干扰强烈的环境, 在菊科和藜科中最为常见, 主要出现在一年生植物中。种子异型性在避免密集负效应、减弱同胞子代间的竞争、采取两头下注策略以适应时空异质性环境等方面具有重要的进化生态意义。该文系统总结了国内外种子异型性的研究工作, 主要内容包括: 1)种子异型性的概念、类型和种类, 2)具有异型种子植物的生境和生活型, 3)异型种子的生态学特性, 4)种子异型性的理论模型, 5)种子异型性的生态意义。在综述文献的基础上, 对今后的研究进行了展望。针对国内外的研究现状, 提出两点建议: 1)系统调查具有种子异型性现象的植物种类, 摸清其生物学特性; 2)确定研究种子异型性现象的模式植物, 从生态学、生理学和分子生物学等学科角度来研究种子异型性的个体发育机制及分子调控机理。     

Dimorphism of the seed-dispersing organ in a pantropical coastal plant,Scaevola taccada: heterogeneous population structures across islands

Scaevola taccada is characterized by dimorphic fruits, with one type having cork and pulp (C-morph) and the other type having only pulp (NC-morph). Although within-individual dimorphism has not been observed, both morphs can occur at the same sites. The cork floats on seawater, and the pulp is eaten by birds. Thus, the morphs may have different seed dispersal abilities, via sea currents and birds, respectively. This study aimed to determine the functional characteristics of the dimorphic fruits. First, the potential seed dispersal ability of sea currents and birds was compared between the two fruits morphs by conducting a floating test and by measuring the proportions of different parts of the fruits, respectively. Next, the frequencies of the two morphs across different substrates (beaches, rocks, and cliffs) in the southern islands around Japan were analyzed. Most C-morph fruits remained floating for more than 180 days in a seawater tank, whereas all NC-morph fruits sank after approximately 1 week. The NC-morph fruits had a more pulp volume and less indigestible material than the C-morph fruits. Although both types of plants were found on many islands and all substrates, the C-morph was dominant on beaches, whereas the NC-morph was most frequent on cliffs. The frequencies of the morphs on different substrates might be influenced by the differences in their seed dispersal abilities. The two morphs may differ in fitness on different substrates. These findings improve our understanding of plant adaptations for dispersal.     

Diaspore ecology of Mertensia maritima: effects of physical treatments and their relative timing on dispersal and germination

This paper reports an experimental study of water dispersal potential and germination of the shingle beach plant Mertensia maritima in which we consider the effects of physical factors (cold treatment, mechanical wear of the pericarp and salt-water exposure) on the diaspores. Our approach also includes testing effects of different orders of the treatments, in contrast to most earlier studies of diaspore ecology. A cold period was necessary to break seed dormancy, and prolonged cold treatment (stratification at 2°C) enhanced germination. Mechanical wear of the pericarp before cold treatment did not affect germination, whereas mechanical wear after cold treatment increased germination significantly. Seeds exposed to 6 weeks of cold treatment before floating in salt water for 6 weeks did not germinate. In contrast, for seeds given the same cold treatment after floating, the germination was more than 50%. Most undamaged and slightly damaged nutlets stayed afloat throughout the dispersal experiment (9 weeks) in 3% salt water, whereas seeds that fell out of damaged nutlets sunk immediately. Thus, the results suggest that the potential for long-distance dispersal is high unless the diaspores (nutlets) are severely damaged, but the order of cold treatment and water dispersal seems to be of great importance for germination: seeds dispersed in autumn (before cold periods) have a much higher probability of germinating than seeds dispersed in winter or early spring (after a cold period). Similar effects of the relative timing of physical processes have hitherto only been reported for two other water-dispersed beach plants. Future studies in diaspore ecology should consider such timing effects as they may be important determinants of the distribution and abundance of plants.     

Factors affecting seed germination and seedling establishment of a long-lived desert shrub (<Emphasis Type="Italic">Coleogyne ramosissima</Emphasis>: Rosaceae)

Long-lived desert shrubs exhibit infrequent, episodic recruitment from seed. In spite of this long time scale, selection on life history attributes that affect seedling recruitment should be strong. We studied factors affecting germination phenology and seedling establishment for Coleogyne ramosissima, a dominant shrub species in the ecotone between warm and cold deserts in western North America. We also examined ecotypic differentiation in establishment strategy in response to selection regimes in two contrasting habitats. We followed patterns of dormancy loss, germination, emergence, and survival in reciprocal field experiments at warm winter Mojave Desert and cold winter Colorado Plateau study sites. Seed germination took place in late winter, under winter rain conditions at the warm desert site and under snow at the cold desert site. Distinctive germination phenologies for the two seed populations at contrasting field sites followed predictions based on laboratory germination experiments. There was no seed bank carryover across years. Seedling survival at the end of three growing seasons was remarkably high (mean survival 54%). Most seedling mortality was due to sprout predation by rodents early the first spring in unprotected caches. Emergence and establishment at each site were significantly higher for seeds from the local population, supporting the idea of ecotypic differentiation in establishment strategy. Establishment success was an order of magnitude greater overall at the Colorado Plateau site, which represents the leading edge of an upward elevational shift in distribution for this species under the current climatic regime. The Mojave Desert site is on the trailing edge of this shift, and recruitment there is apparently a much less frequent occurrence.     

Performance and responses to competition in two congeneric annual species: does seed heteromorphism matter?

Variations in seed characteristics observed in heteromorphic species may affect various stages of their life cycles, e.g. seed dormancy, germination characteristics or even adult plant performance. Highly specialised seed morphs – described as colonisers and maintainers – exhibit a trade‐off between colonisation capacity and competitive traits. The performance of distinct seed morph progeny under competitive conditions, and especially in multi‐species arrangements, had previously not been given much attention. In this study, we compared performance and response to competition among distinct seed morph progenies in two congeneric, co‐occurring species: the invasive Bidens frondosa and the non‐invasive Bidens tripartita. We hypothesised that maintainer seed morphs of both species would perform better under increased plant densities and within inter‐morphic mixtures, while coloniser morphs would show stronger responses to increased densities and perform relatively poorly in inter‐morphic mixtures. We conducted a growth trial and a greenhouse experiment, which revealed that seed morph progeny differed significantly in plant height when grown without competition, while under competitive conditions such differences became less apparent. The observed pattern was more strongly pronounced in B. frondosa, which showed a general predominance in stature and biomass over its non‐invasive congener. Although seed morphs performed equally well under competitive conditions, increased plant height and more rapid germination can favour the maintainer seed morph on sites where vegetation is already present.