Variability and Cryptic Heteromorphism of Ambrosia artemisiifolia Seeds: What Consequences for its Invasion in France?

BACKGROUND AND AIMS: Ambrosia artemisiifolia is a ruderal weed introduced from North America to Europe. It produces large amount of achenes which are highly heterogeneous in size. Due to the preponderant role of propagules in invasive plant processes, the achene mass variability related to germination, dispersal strategy and life history traits of offspring were investigated within this species. METHODS: The variability in achene mass was quantified among six populations sampled in different habitats. The effects of achene mass variation on germination were studied. The percentages of floating and non-floating achenes were evaluated in the studied populations. The consequences of floatability on the growth and traits of the offspring were studied. KEY RESULTS: Mean achene mass ranged from 1.72 to 3.60 mg, depending on the populations, and was highly variable. Variation among achenes within plants accounted for 63.9 % of the variance, whereas variances among plants within each population (22.2 %) and among populations (13.9 %) were lower. Achene masses were also positively correlated to the total germination percentage for four populations out of six. Two kinds of achenes were distinguished: floating and non-floating. The majority of floating achenes (90 %) sank 24 h after water immersion. Whatever the population, floating achenes were lighter, more dormant and germinated faster than non-floating achenes. Plants which issued from floating achenes had better growth than those from non-floating achenes. CONCLUSIONS: The capacity of A. artemisiifolia to be invasive in Europe appears to be high, possibly due to its huge plasticity in seed mass which may help it to cope with a wide range of conditions and to establish in disturbed habitats. Furthermore, the recent invasion of southern France by A. artemisiifolia could be partially explained by water dispersal of achenes through rivers and has pinpointed its colonization potential along French rivers.     

Variation in Dispersal Traits in a Narrow-endemic Plant Species, <Emphasis Type="Italic">Centaurea corymbosa</Emphasis> Pourret. (Asteraceae)

The existence of genetic variability for dispersal is a crucial issue for organisms facing increased habitat fragmentation and climate change. We study the genetic basis and evolutionary potential for diaspore traits related to dispersal in Centaurea corymbosa. Using diaspores collected in natural conditions in four of the six extant populations of this narrow-endemic plant species and diaspores produced in a common garden experiment, we study the variation for pappus and achene sizes, and diaspore mass. Using a sample of achenes from the common garden experiment, we find that the best predictor of terminal velocity is a linear combination of pappus length, achene width, and achene weight. We find significant differences among populations for all traits in both conditions, as well as significant differences among families within population. Although the differences among populations for some traits are not exactly the same in controlled conditions compared to natural conditions, the ranking of populations according to their mean trait values is consistent in both conditions. Our study is therefore one of the first to show a correlation between phenotypic differentiation for dispersal traits in natural conditions vs. controlled conditions. We also show evidence of genetic variation for traits commonly thought to be involved in dispersal ability, suggesting the potential for evolutionary changes following environmental change and management actions.Co-ordinating editor: J.F. Stuefer     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Inbreeding effects on fitness traits in the heterocarpic herb Leontodon autumnalis L. (Asteraceae)

Heterocarpic plants are characterized by the production of distinct types of fruits that usually differ in their ecological behavior. In the Asteraceae, differences are mainly found between peripheral non-dispersal and central dispersal achenes (single-seeded fruits). Inbreeding depression is considered as an evolutionary force as it may reduce several fitness traits, and in the case of heterocarpic plants, it could influence fitness traits (e.g., seed set, germination rate, growth rate) of each fruit morph, which may have important ecological and evolutionary consequences. In particular, differential effects on fitness traits and dispersal of selfed and outcrossed progeny can strongly determine the viability of extant populations and the potential to colonize new habitats. We conducted a hand-pollination experiment in greenhouse conditions to test whether inbreeding affects the fitness of achene morphs in the heterocarpic herb Leontodon autumnalis (Asteraceae). Results show that achene morphs significantly differ in their ecological behavior, peripheral achenes germinating more and faster than central achenes. The significant interaction between pollination treatment and achene morph for germination probability might indicate a link between dormancy and mating system in L. autumnalis: germination was higher for outcrossed achenes in central achenes whereas the opposite pattern was exhibited by peripheral achenes. Selfing dramatically reduced seed set, probably as a consequence of strong self-incompatibility mechanisms rather than inbreeding effects. Inbreeding depression significantly affected late life-cycle traits, such as growth rate and biomass at flowering. Overall, results suggest that inbreeding depression seems to be an important selective force maintaining outcrossing in L. autumnalis.     

QUANTITATIVE GENETICS OF SIZE,SHAPE, LIFE-HISTORY,AND FRUIT CHARACTERISTICS OF THE SEED-HETEROMORPHIC COMPOSITE HETEROSPERMA PINNATUM. I. VARIATION WITHIN AND AMONG POPULATIONS

We document phenotypic and genetic variation within and among populations of the seed heteromorphic species Heterosperma pinnatum Cav. (Compositae) in the production of seed morphs and in a variety of life-history and morphological characteristics that might be correlated with seed and head traits. Each trait is found to have significant genetic variance in most or, usually, all populations. Significant among-population genetic variation exists for all traits except number of achenes per head and seedling shape, although some traits have much less genetic variation among than within populations. Number and percentage of intermediate achenes per head, total number of achenes per head, and lengths of central and peripheral achenes had little among-population genetic variation compared to within-population variation. Most traits had slightly less genetic variation among than within populations; however, some traits (percentage of central achenes, length of awns, date that the first flowering head opened, date that the first fruiting head opened, and death date) had more among-population genetic variation. The proportions of achene morphs produced had high broad-sense heritabilities and high among-population genetic variance, except in the case of intermediate achenes. All phenological variables had high among-population genetic variation. Within-population heritabilities were high for dates of first flowering head and fruiting head but low for death date and reproductive interval. Family and population means measured in the greenhouse for traits having high broad-sense heritability or among-population genetic variance were closely correlated with field means for the corresponding families or populations. The amounts of phenotypic variation were similar for traits that were measured in both the field and the greenhouse. These lines of evidence suggest that greenhouse results provide reasonable estimates of genetic variation in the field for this species. Numerous studies have reported variation in the proportion of seed morphs for different heteromorphic-seeded species and have discussed adaptive scenarios for the evolution of seed proportions; however, our investigation is one of only a few that have documented the amount of phenotypic and genetic variation within and among populations.     

Plastic bet-hedging in an amphicarpic annual: an integrated strategy under variable conditions

Amphicarpy is a form of diversified bet-hedging expressed mostly in annual plants, where two types of offspring are produced with two distinct ecological roles: long-range aerial dispersers and highly competitive subterranean, sedentary fruit. Emex spinosa is a semi-arid, amphicarpic annual, inhabiting habitats with different levels of environmental variation. We tested the hypothesis that, in E. spinosa, bet-hedging may be “fine-tuned” by plasticity in the phenotype ratio (aerial/subterranean fruit mass) as a function of environmental conditions. We conducted a greenhouse experiment, manipulating nutrient availability and intraspecific density, to determine the pattern of ratio shifts. In order to determine whether the integrated strategy is an adaptation to variable habitats, a similar common garden experiment was conducted, comparing two natural populations differing in environmental variability. The offspring ratio shifted in response to both nutrient availability and plant density. In pots containing single plants the ratio increased steeply with nutrient availability, while in pots containing eight plants a more moderate increase occurred. These shifts were the result of plasticity in allocation to both achene types, as well as ontogenetic effects on aerial achene production. The degree of response increased with the heterogeneity of the habitat of origin. We found evidence for an adaptive integrated strategy, with bet-hedging “fine-tuned” by phenotypic plasticity. Strenuous conditions tended to shift the offspring ratio towards securing subterranean reproductive success, while favorable conditions resulted in a shift towards dispersible achenes. The authors Asaf Sadeh and Hagai Guterman contributed equally to this study.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

Effects of pollen donors on seed formation inEspeletia schultzii (Compositae) populations at different altitudes

The influence of different pollen donors on seed formation was investigated in three populations ofEspeletia schultzii that differ in environmental conditions and life history characteristics. Self pollen and pollen from different donors (< 15m apart) within each population was used in a diallel design in order to test the genetic base of seed set variation. Three measures of seed formation were used: (1) achene number; (2) proportion of filled achenes (fruits) that distinguishes between achenes with seeds and empty achenes; (3) proportion of aborted seeds that distinguishes between viable and aborted seeds. Self-pollinations resulted in empty achenes. Achene number did not vary between the different pollen donors. A bimodal pattern of filled achenes was found in two populations in two consecutive years. On the other hand, a unimodal pattern was found in crosses between more distant donors (> 30m). These patterns seems to be the results of a sporophytic incompatibility system. Seed abortion was highest at the higher elevations and seems to be correlated with elevation rather than with any genetic effect.     

Role of within-individual variation in capitulum size and achene mass in the adaptation of the annual Centaurea eriophora to varying water supply in a Mediterranean environment

To clarify the adaptive value of variation in capitulum size and achene mass, plants of Centaurea eriophora were studied in a glasshouse and in a natural population. C. eriophora plants consist of a basal leaf rosette from which an erect stem grows, with lateral branches of various orders ending in capitula of different orders. Primary, secondary and tertiary capitula are comparable in size and they produce similar numbers of achenes, which are similar in weight (large achenes). These capitula are formed during April, May and early June, and constitute the normal or primary flowering. Following ripening of tertiary capitula, leaves senesce, but, later during June and the first half of July, a secondary flowering of a variable number of smaller capitula may occur if wet conditions persist for longer than usual. Plants that have almost senesced develop small lateral branches 1-2 cm long bearing a few small leaves and ending in a capitulum about half the diameter of capitula from the primary flowering period. The number of achenes produced in these capitula (small achenes) and their weight are 70 and 30% less, respectively, than those of capitula formed during primary flowering. These reductions appear to result from restricted availability of resources. Large and small achenes have similar dispersal characteristics and possess similar germination potential. However, large achenes produce seedlings that are capable of emerging from greater burial depths, providing the resulting plants with a potential advantage. The normal flowering period coincides with the optimum time of year for flowering and fruiting in the south of Spain, and only if rainfall lasts longer than usual does secondary flowering occur. Secondary flowering extends the normal flowering and fruiting periods, thereby providing a supplementary crop of smaller, yet viable, fruits. It can be considered to be an adaptive response to the unpredictable Mediterranean climate, optimizing the use of available resources.     

Role of mucilage during achene germination and sprout growth of the endangered Tibetan medicinal herb Mirabilis himalaica (Nyctaginaceae) exposed to abiotic stresses

Aim Mirabilis himalaica (Nyctaginaceae) is an endangered medicinal plant mainly distributed in the plateau region of northern Tibet, China. The outer surface of M. himalaica achenes is covered by a pectinaceous mucilaginous layer upon hydration. However, the role of the achene mucilage is poorly understood. In this study, we investigated the effects of mucilage on achene germination and sprout growth under abiotic stress to explain how M. himalaica survive the alpine environment.Methods We investigated the effect of mucilage on achenes germination by contrast the capacity of water absorption, dehydration and respiration of intact achene and the achene with mucilage removal. We performed abiotic stresses experiments including drought stress, salt stress, cold stress and high temperature stress, and quantified the effects of mucilage removal on achene germination rate, root and shoot lengths of seedlings.Important findings Mucilage is extremely hydrophilic, and the mass of intact achenes can be 9-fold greater than that of demucilaged achenes. The removal of the mucilaginous layer did not significantly change final germination percentages under ideal conditions, but intact achenes (i.e. with mucilage) took longer to germinate. The mucilage significantly decreased seed respiration rates by acting as a physical barrier that prevented oxygen diffusion. Germination rates, shoot and root growth of intact achenes were higher than those of demucilaged ones during exposures to cold, heat, osmotic and salt stresses. Achene mucilage presumably plays an ecologically important role in the life cycle of M. himalaica by aiding the critical achene germination and early seedling growth in the stressful habitats of the plateau region of northern Tibet.     

Influence of achene heteromorphism on life-cycle traits in the annual weed gallant soldier (Galinsoga parviflora Cav.)

Galinsoga parviflora (gallant soldier), Asteraceae, produces two morphologically distinct achenes in a single capitulum: peripheral and central. The morphological, phenological and generative reproduction traits of the progeny derived from peripheral and central achenes that were cultivated in greenhouse conditions were analyzed. Differences between the progeny of various morph types were manifested at different stages of life. The plants of both morph types developed at a similar rate and they entered key phenological phases at the same time except for the flowering stage. The average height of the studied plants was similar on the same days of the experiment. The study showed that plants from peripheral achenes realize the generative reproduction in other pattern than plants from central achenes. The plants from central achenes produce more capitula per plant at the beginning of the fruiting stage, whereas plants from peripheral seeds achieve the highest number of capitula per plant at later stages. Finally, on day 130 of the experiment, the number of capitula per plant of both morphs equalized. At the early fruiting stage, the number of achenes per capitulum of plants from the two morphs did not differ significantly. The number of achenes within one capitulum decreases with plant age in both populations, but the rate of that decline is greater in the progeny of peripheral seeds. The number of peripheral achenes per capitulum in plants from both achene types decreases at a comparable rate. In contrast, the number of central achenes is reduced at a faster rate in plants germinated from peripheral achenes. After day 83, the individuals from peripheral diaspores intensified their reproductive effort; they produced more capitula but with fewer number of achenes than in individuals from central diaspores. On day 130, the number of dispores per plant of the two morphs equalized. The changes of fecundity of peripheral and central progeny with the age of the plant (at different stages of the life cycle of a single plant) contribute to an intensive seed rain throughout the fruiting period. By producing a large number of less fertile capitula with a stable number of peripheral achenes, plants derived from peripheral achenes are able to supply a higher share of peripheral achenes than if they would produce fewer but more fertile capitula. This strategy increases the pool of peripheral achenes.     

The role of pollination in the population biology of the monocarpic species Cirsium palustre and Cirsium vulgare

Summary The absence of cross-pollination in Cirsium palustre and Cirsium vulgare resulted in reduced achene production while the achenes produced were heavier than those produced after cross-pollimation. Establishment of plants from non-cross-pollinated achenes is comparatively higher, facilitating the founding of a population from isolated individuals in these wind-dispersed species.If achene weight is the result of a balance between cross-pollination and self-pollination (and/or apomixis), the first causing more and thereby lighter achenes to develop, increasing density of flowering individuals in a population may lower achene weight and consequently seedling survival. This may account for the frequently observed decline of populations of fugitive monocarpic perennials.Publication of the Meijendel-comité, Nieuwe Serie no. 58     

The role of the soil seed bank in the unpredictable habitat of Anthemis chrysantha J. Gay,a critically endangered species

The ability of Anthemis chrysantha to form a soil seed bank (SB) was investigated in order to understand better the adaptation of this rare annual species to its arid and unpredictable natural habitat. The natural seed bank population was analyzed in five consecutive plant cohorts (2006–2010) by sampling at two different times: in May, after the germination period, and in October, after the first dispersal episodes due to the beginning winter rainfall. In addition, to determine the persistence in the soil of the two achene morphs of this species’ (white and dark achenes), an artificial SB was created where achenes were buried and exhumed successively after each season, during two years. In all the cohorts studied, seedling emergence from May samples indicated the existence of a persistent seed bank (PSB). Moreover, the dark achenes were largely responsible for the permanent fraction because, after two years of burial in the artificial SB, 85.0% of them remained apparently healthy without having germinated, versus 9.9% of white achenes. Both types of buried achenes exhibited an annual conditional dormancy/non-dormancy cycle, induced by low winter temperatures. The PSB dynamics appeared to oscillate between the minimum values at the end of the germination period in spring (up to 2000 achenes per square meter) and the maximum values of the dispersal episodes in early autumn (up to 6000 achenes per square meter), with fluctuations of achene density due to the variability in annual rainfall. Hence, the SB showed a decline due to the failure of fruiting in the 2008 cohort, caused by drought, although the low value of ca. 560 achenes per square meter was able to establish the following population. Our study highlights the importance of the PSB, which, in “bad” years, may be critical for the persistence of this species.     

Effect of source–sink ratio on seed set and filling in sunflower (Helianthus annuus L.)

Poor seed development in sunflower may result from insufficient assimilate supply (source limitation). To test this hypothesis, the effects of changed source–sink ratio on seed set (measured as percentage of empty achenes) and seed filling (measured as dry mass per filled achene) in individual plants were investigated. Source–sink ratio, defined as leaf area per floret (LAF), was experimentally altered using invasive (floret removal, defoliation) and non‐invasive (pulse of chilling, short days or shading during leaf or floret initiation) treatments. Shading at floret initiation proved the most effective non‐invasive method. Generally, an increase, or decrease, in LAF improved, or impaired, both seed set and filling. Increasing LAF by 2.0 cm²[95% confidence interval (1.5, 2.5)] decreased the percentage of empty achenes by 36.9%‐points (?41.9, ?30.9) and increased dry mass per filled achene by 20.1 mg (13.6, 26.7) in the capitulum centre. The effect of source–sink ratio on seed set was always strongest in the centre, whereas peripheral whorls were not affected. Achene mass was affected in all parts of the capitulum. It is concluded that source limitation is a major cause for empty achenes in sunflower plants grown under non‐stress conditions.     

POPULATION DYNAMIC CONSEQUENCES AND EVOLUTION OF SEED TRAITS OF HETEROSPERMA PINNATUM (ASTERACEAE)

We summarize research on variation in achene morphology of Heterosperma pinnatum. Each flowering head has a broad range of achene morphologies. There is no between-year seed bank. There are no ecologically relevant differences in achene size, seedling size, or seedling growth rates of the different morphs. Achenes located centrally in the heads lose dormancy earlier during the period between autumn achene production and the onset (May-July) of the rainy season. This results in relatively more early emergence of central achenes. Awned central achenes also have greater adhesive dispersibility. Emergence, survival, and fecundity were measured for achenes sown into natural populations in central Mexico. When harsh conditions result in few seeds surviving to reproduce, the advantage tends to go to the more “conservative” peripheral achenes. Greenhouse experiments show a strong genetic component to the determination of achene proportions within and among populations and a variety of genetic correlations. Populations with a high proportion of awned achenes tend to be found in vegetation types where they are likely to be ephemeral. A high proportion of central achenes tend to be found in sites with little pre-rainy season precipitation. These patterns are interpreted in terms of selection for dispersal and the within-year timing of germination.     

How much genetic variation is stored in the seed bank? A study of Atriplex tatarica (Chenopodiaceae)

We investigated to what extent the soil seed bank differed genetically and spatially in comparison to three consecutive life history stages (seedlings, mature plants, and fruiting plants) in a natural population of Atriplex tatarica. Representatives of particular life history stages from twenty subunits within a large population were randomly collected and subjected to allozyme analysis. Comparison of population polymorphism among various life history stages showed significant differences in observed heterozygosity (H(O)) and F statistics (F(IS) and F(ST)), but nonsignificant ones in the cases of number of alleles per polymorphic locus (A) and gene diversity (H(S)). These results indicate an increasing number of heterozygotes, a decreasing level of inbreeding and an increase of the partitioning genetic diversity among populations with increasing population age. Spatial autocorrelation was used to calculate f, the average co-ancestry coefficient between individuals within distance intervals of two meters along a 39 m long transect. Significant positive fine scale genetic structure was detected in mature and fruiting plants but not in soil seeds and seedlings stages. The results of the presented study on A. tatarica indicated that significant differences exist in genetic differentiation, differentiation in allele frequencies and spatial autocorrelation among early (soil seeds and seedlings) and late (mature and fruiting plants) life history stages but not within early and late ones. This pattern suggests that, rather than storing genetic variability in the soil or germination and establishment success, self-thinning might be the major microselective force in populations of A. tatarica.     

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.     

Domestication and the distribution of genetic variation in wild and cultivated populations of the Mesoamerican fruit tree Spondias purpurea L. (Anacardiaceae)

Domestication occurs as humans select and cultivate wild plants in agricultural habitats. The amount and structure of variation in contemporary cultivated populations has been shaped, in part, by how genetic material was transferred from one cultivated generation to the next. In some cultivated tree species, domestication involved a shift from sexually reproducing wild populations to vegetatively propagated cultivated populations; however, little is known about how domestication has impacted variation in these species. We employed AFLP data to explore the amount, structure, and distribution of variation in clonally propagated domesticated populations and sexually reproducing wild populations of the Neotropical fruit tree, Spondias purpurea (Anacardiaceae). Cultivated populations from three different agricultural habitats were included: living fences, backyards, and orchards. AFLP data were analysed using measures of genetic diversity (% polymorphic loci, Shannon's diversity index, Nei's gene diversity, panmictic heterozygosity), population structure (F(ST) analogues), and principal components analyses. Levels of genetic variation in cultivated S. purpurea populations are significantly less than variation found in wild populations, although the amount of diversity varies in different agricultural habitats. Cultivated populations have a greater proportion of their genetic variability distributed among populations than wild populations. The genetic structure of backyard populations resembles that of wild populations, but living fence and orchard populations have 1/3 more variability distributed among populations, most likely a reflection of relative levels of vegetative reproduction. Finally, these results suggest that S. purpurea was domesticated in two distinct regions within Mesoamerica.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.