Relationship between seed bank expression, adult longevity and aridity in species of Chaetanthera (Asteraceae) in central Chile

BACKGROUND AND AIMS: Broad surveys have detected inverse relationships between seed and adult longevity and between seed size and adult longevity. However, low and unpredictable precipitation is also associated with seed bank (SB) expression in semi-arid and arid areas. The relationship between adult longevity, SB formation, seed mass and aridity is examined in annual and perennial herbs of Chaetanthera (Asteraceae) from the Chilean Mediterranean-type climate and winter-rainfall desert areas over a precipitation range of one order of magnitude. METHODS: Seeds of 18 species and subtaxa (32 populations) were buried in field locations, and exhumed after two successive germination periods. Seeds not germinating in the field were tested in a growth chamber, and remnant intact seed tested for viability. Seed banks were classed as transient or persistent. The effect of life form, species, population and burial time on persistent SB size was assessed with factorial ANOVA. Persistent seed bank size was compared with the Martonne aridity index (shown to be a surrogate for inter-annual variation in precipitation) and seed size using linear regression. ANCOVA assessed the effect of life-form on SB size with aridity as covariate. KEY RESULTS: Three species had a transient SB and 15 a persistent SB. ANOVA revealed a significant effect of life-form on SB size with annuals having larger SB size and greater capacity to form a persistent SB than perennials. Significant inter-population variation in SB size was found in 64% of cases. Seed mass was negatively correlated with persistent SB size. Persistent seed bank size was significantly correlated with the Martonne aridity index in the perennial and annual species, with species from more arid areas having larger persistent SBs. However, when aridity was considered as a covariate, ANCOVA revealed no significant differences between the annual and perennial herbs. CONCLUSIONS: Persistent seed bank size in Chaetanthera appears to reflect environmental selection rather than any trade-off with adult longevity.     

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.     

Strong genetic differentiation on a fragmentation gradient among populations of the heterocarpic annual Catananche lutea L. (Asteraceae)

In landscapes which are predominately characterised by agriculture, natural ecosystems are often reduced to a mosaic of scattered patches of natural vegetation. Species with formerly connected distribution ranges now have restricted gene flow among populations. This has isolating effects upon population structure, because species are often confined by their limited dispersal capabilities. In this study, we test the effects of habitat fragmentation, precipitation, and isolation of populations on the genetic structure (AFLP) and fitness of the Asteraceae Catananche lutea. Our study area is an agro-dominated ecosystem in the desert–Mediterranean transition zone of the Southern Judea Lowlands in Israel. Our analysis revealed an intermediate level of intra-population genetic diversity across the study site with reduced genetic diversity on smaller scale. Although the size of the whole study area was relatively small (20?×?45?km), we found isolation by distance to be effective. We detected a high level of genetic differentiation among populations but genetic structure did not reflect spatial patterns. Population genetic diversity was correlated neither with position along the precipitation gradient nor with different seed types or other plant fitness variables in C. lutea.     

Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient

I applied a comparative approach to reveal correlated patterns of variation in phenology and seed production in four populations of two annual grasses Hordeum spontaneum and Avena sterilis, sampled in the same environments distributed along an aridity gradient in Israel. The steep aridity gradient in Israel represents two parallel clines of environmental productivity (annual rainfall) and predictability (variation in amount and timing of annual rainfall) that is likely to induce similar responses in natural plant populations distributed along the gradient, if (1) selection is strong, (2) species share the same ecological niche, and (3) there is genetic variation for ecologically important traits. I found in plants of both species (1) ultimate advance in onset of flowering, and (2) more but smaller seeds, with increasing aridity. The broad sense heritabilities of onset of flowering, seed size and seed yield in both species were very high, moderate and low, respectively. It appears that the observed adaptive complex of traits have evolved in both species in response to this specific array of environments.     

Ploidy, geographical distribution and morphological differentiation of Parasenecio auriculata (Senecioneae; Asteraceae) in Japan

Parasenecio auriculata is a woodland perennial herb widely distributed in Northeastern Asia, constituted by a poorly understood polyploidy with a diploid (2n=2x=60) and a tetraploid (2n=4x=120). In this study, for a better understanding of the polyploidal evolution, cytogeography and morphological variation were analyzed in Japanese P. auriculata, including two varieties; var. bulbifera endemic to central Hokkaido and var. kamtschatica widely distributed in northern Honshu and Hokkaido. The occurrence of two polyploidal levels was reconfirmed. While var. bulbifera is predominantly tetraploid, var. kamtschatica is comprised of diploid and tetraploid. Morphological variation among 22 quantitative characteristics is continuous and not distinctive among cytotypes or varieties, but plant size tended to be larger in the order, diploid of var. kamtschatica, var. bulbifera, and tetraploid of var. kamtschatica. The cytotype distribution showed a conspicuous geographical pattern. Besides var. bulbifera endemic to the central Hokkaido, the diploid of var. kamtschatica is mainly found in Southern Hokkaido, and the tetraploid has a disjunct distribution in eastern and northern Hokkaido and northern Honshu. Such a geographical pattern is possibly attributable to the differentiation of climatic preference among cytotypes and varieties, and may have been established in association with the climatic cline along the Japanese archipelago.     

Summer annual plants exhibit germination responses to an aridity gradient consistent with both delayed germination and phenotypic plasticity

Plant Ecology - Two adaptive strategies to face variable environments have been proposed for annual plants in deserts: delayed and plastic (predictive) germination. All theories and models have...     

Genetic diversity and morphological differentiation in Liatris helleri (Asteraceae), a threatened plant species

Liatris helleri (Asteraceae) is an insect-pollinated herbaceous perennial endemic to several high-elevation sites in the Blue Ridge Mountains of North Carolina. Allozymes were used to describe the genetic diversity and population structure in nine populations of this rare, federally listed species. Differences in leaf morphology were also examined for greenhouse-grown plants representing several populations. The proportion of the total genetic diversity found among populations, as indicated by the allozyme data, was 16%. Higher levels of population differentiation were found for differences in leaf shape; population of origin accounted for 37% of the variation in maximum leaf width, while families within populations accounted for 7%. In contrast to many endemic species, L. helleri maintains fairly high levels of genetic diversity. For the species, the percent polymorphic loci was 87.5, the average number of alleles at variable loci was 3.00 and the gene diversity was 0.276. Mean population values were percent polymorphic loci =58.4, mean number of alleles per polymorphic locus =2.59 and gene diversity =0.219. The estimated gene flow was low (Nm=0.85–1.32) and a relatively high correlation (r=0.55; p<0.005) was found between linear geographic and genetic distance. This suggests that the populations are partially isolated by distance, despite the limited range (<60 km) of the species. We recommend that population distinetiveness be maintained in restoration efforts.     

Adaptation of metabolism and evaporative water loss along an aridity gradient

Broad-scale comparisons of birds indicate the possibility of adaptive modification of basal metabolic rate (BMR) and total evaporative water loss (TEWL) in species from desert environments, but these might be confounded by phylogeny or phenotypic plasticity. This study relates variation in avian BMR and TEWL to a continuously varying measure of environment, aridity. We test the hypotheses that BMR and TEWL are reduced along an aridity gradient within the lark family (Alaudidae), and investigate the role of phylogenetic inertia. For 12 species of lark, BMR and TEWL decreased along a gradient of increasing aridity, a finding consistent with our proposals. We constructed a phylogeny for 22 species of lark based on sequences of two mitochondrial genes, and investigated whether phylogenetic affinity played a part in the correlation of phenotype and environment. A test for serial independence of the data for mass-corrected TEWL and aridity showed no influence of phylogeny on our findings. However, we did discover a significant phylogenetic effect in mass-corrected data for BMR, a result attributable to common phylogenetic history or to common ecological factors. A test of the relationship between BMR and aridity using phylogenetic independent constrasts was consistent with our previous analysis: BMR decreased with increasing aridity.     

Phylogeny of Chaetanthera (Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages

Penalized likelihood analysis of previously published chloroplast DNA (cpDNA) ndhF sequences suggests that the central-southern Andean genus Chaetanthera diverged ca. 16.5 million years (my) ago, well before the uplift of the Andes to their present heights. Penalized likelihood analysis based on new nuclear ribosomal DNA (rDNA) internal transcribed spacer (ITS) sequences indicates that the most relictual lineages occupy high elevation Andean habitats that did not exist until some 10my later. This result is contrary to the expectation that younger habitats should be occupied by phylogenetically younger lineages. The results are interpreted with respect to the development of aridity in lowland habitats during the Miocene and Pliocene, which presumably extinguished the lowland relatives of the high elevation taxa or, in effect, forced them upwards in search of adequate moisture. As the more northerly lineages were being displaced upward, others diversified in the mediterranean-type climate area of central Chile, giving rise to additional high elevation taxa again, at an early date, as well as lowland taxa. Some species of Chaetanthera from lowland central Chile appear as the phylogenetically youngest taxa, suggesting secondary adaptation to lowland aridity. At the same time, at least two high elevation species, Chaetanthera peruviana and Chaetanthera perpusilla, appear to have been derived recently from a lower elevation ancestor, while some middle to low elevation taxa seem to have evolved recently out of a high elevation complex. The results suggest that the younger high elevation habitats have served as both "cradle" and "museum" for Chaetanthera lineages.     

Annual plant–shrub interactions along an aridity gradient

Annual plants in semi-arid and arid areas are often closely associated with shrubs. The degree of association largely depends on the balance of negative and positive effects between these contrasting plant life-forms, ranging from interference to facilitation. Since positive interactions are predicted to become less important with increasing rainfall, the interaction balance is expected to shift along aridity gradients. However, this prediction has not been tested on a community level and for different life-history stages across large geographical gradients. Here, we show such changes for annual plant populations and communities in four contrasting sites along a steep climatic gradient, ranging from the arid desert to mesic Mediterranean regions in Israel. Above-ground productivity, richness, seedling density, and seed bank density of the annual plant community, as well as fecundity of annual plant populations, were generally higher under shrubs than in areas between shrubs at the arid end of the gradient, but significantly lower at the humid end. Net effects of shrubs on annuals expressed as relative interaction intensity indicated a steady and consistent shift from net positive or neutral effects in the desert to net negative effects in the mesic part of the gradient. These findings emphasize the usefulness of studies along large-scale gradients encompassing a wide range of environmental conditions for understanding community level interactions among coexisting species.     

High outcrossing in the annual colonizing species Ambrosia artemisiifolia (Asteraceae)

Background and Aims

Variation in mating patterns may be particularly evident in colonizing species because they commonly experience wide variation in plant density. Here, the role of density for the mating system of Ambrosia artemisiifolia (common ragweed), a wind-pollinated annual colonizing species previously reported as self-compatible, is explored.

Methods

The effect of population density on the proportion of self- and cross-fertilized seeds was examined using allozyme markers and experimental arrays conducted over two seasons in the field. Also the reproductive success of isolated plants located in diverse habitats was measured. The potential occurrence of a physiological mechanism preventing self-fertilization, i.e. self-incompatibility, following controlled self- and cross-pollinations in the glasshouse was examined.

Key Results

Outcrossing rates estimated using allozyme markers were uniformly high, regardless of the spacing between plants. However, when single plants were isolated from congeners they set few seeds. Observations of pollen-tube growth and seed set following controlled pollinations demonstrated that plants of A. artemisiifolia possess a strong self-incompatibility mechanism, contrary to earlier reports and assumptions.

Conclusions

The maintenance of high outcrossing rates in colonizing populations of A. artemisiifolia is likely to be facilitated by the prodigious production of wind-borne pollen, high seed production and extended seed dormancy.Key words: Self-incompatibility, outcrossing rate, density dependence, colonization, wind-pollination, Ambrosia artemisiifolia (ragweed), Asteraceae     

Estimates of nitrogen fixation by trees on an aridity gradient in Namibia

Summary Nitrogen (N₂) fixation was estimated along an aridity gradient in Namibia from the natural abundance of ¹⁵N (¹⁵N value) in 11 woody species of the Mimosacease which were compared with the ¹⁵N values in 11 woody non-Mimosaceae. Averaging all species and habitats the calculated contribution of N₂ fixation (N _f ) to leaf nitrogen (N) concentration of Mimosaceae averaged about 30%, with large variation between and within species. While in Acacia albida N _f was only 2%, it was 49% in Acacia hereroensis and Dichrostachys cinerea, and reached 71% in Acacia melifera. In the majority of species N _f was 10–30%. There was a marked variation in background ¹⁵N values along the aridity gradient, with the highest ¹⁵N values in the lowland savanna. The difference between ¹⁵N values of Mimosaceae and non-Mimosaceae, which is assumed to result mainly from N₂ fixation, was also largest in the lowland savanna. Variations in ¹⁵N of Mimosaceae did not affect N concentrations, but higher ¹⁵N-values of Mimosaeae are associated with lower carbon isotope ratios (¹³C value). N₂ fixation was associated with reduced intrinsic water use efficiency. The opposite trends were found in non-Mimosaceae, in which N-concentration increased with ¹⁵N, but ¹³C was unaffected. The large variation among species and sites is discussed.This paper is prepared in memory of J. Visser, who took part in the collection of species, but died in 1990     

Genetic,cytogenetic and morphological diversity in Helicrysum leucocephalum (Asteraceae) populations

Helichrysum (family Asteraceae) is a large, heterogeneous and polyphyletic genus. Most of the Helichrysum species have ornamental and medicinal values. Helicrysum leucocephalum Boiss., has wide geographical distribution in Iran and forms several local populations. The present investigation was performed to study inter populations, genetic, cytogenetic and morphological diversity of this medicinal species. The AMOVA test showed significant genetic difference among the studied populations but the Mantel test did not show correlation between the genetic distance and geographical distance of these populations. STRUCTURE and reticulation analyses showed some degree of genetic admixture and gene exchange among the studied populations. These populations had different chromosome numbers and also differed significantly in the size of their chromosomes.     

Quantitative morphological variation inTetramolopium (Asteraceae) in Hawaii and the Cook Islands

Multivariate analyses of quantitative characters were undertaken to examine the pattern of variation resulting from adaptive radiation ofTetramolopium in Hawaii and to investigate the systematics of an undescribed species in the Cook Islands and a group of populations of uncertain affinity on Kalaupapa Peninsula in Hawaii. Taxa differentiated on quantitative characters in the absence of qualitative characters. Differentiation was observed in multiple characters which could be organized into a few principal components. The analyses supported the species rank of the Cook Islands plants. The relationship of the Kalaupapa Peninsula populations to the rest of the taxa needs further study.     

Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert

The Namib Desert is considered the oldest desert in the world and hyperarid for the last 5 million years. However, the environmental buffering provided by quartz and other translucent rocks supports extensive hypolithic microbial communities. In this study, open soil and hypolithic microbial communities have been investigated along an East–West transect characterized by an inverse fog-rainfall gradient. Multivariate analysis showed that structurally different microbial communities occur in soil and in hypolithic zones. Using variation partitioning, we found that hypolithic communities exhibited a fog-related distribution as indicated by the significant East–West clustering. Sodium content was also an important environmental factor affecting the composition of both soil and hypolithic microbial communities. Finally, although null models for patterns in microbial communities were not supported by experimental data, the amount of unexplained variation (68–97 %) suggests that stochastic processes also play a role in the assembly of such communities in the Namib Desert.     

Phenological responses to temperature of an annual and a perennial Lesquerella species

BACKGROUND AND AIMS: The annual Lesquerella fendleri, native to the south-western desert of United States and Mexico, and the perennial L. mendocina, native to Argentina, may have potential as new crops for cold-arid environments. The introduction of a new crop requires an understanding of environmental influences on growth and development, particularly temperature, which has been recognized as the main factor affecting the rate of development in crops. The objective of this study was to examine differences in the phenology of L. fendleri and L. mendocina and in the response to temperature in both vegetative and reproductive phases. METHODS: Plants of each species were grown at a range of constant temperatures under controlled conditions and developmental responses were analysed and quantified. KEY RESULTS: The rate of development of L. fendleri increased linearly with temperature in the phase from emergence (EM) to floral bud appearance (FBA) over the range 9-20 degrees C, and for the phase from FBA to first flower open (FL) over the range 9-24 degrees C. In contrast, the rate of development of L. mendocina was insensitive to temperature during the phase EM to FBA. In the phase FBA to FL, L. mendocina had a lower sensitivity to temperature than L. fendleri. In addition, L. fendleri exhibited a quantitative response to supra-optimal temperatures (reducing rate of development with further increases in temperature) whereas L. mendocina showed a qualitative response, with development ceasing to progress at temperatures above the optimum. CONCLUSIONS: This differential behaviour at high temperatures could explain the biennial habit found for L. mendocina sown during late spring under field conditions, whereas it behaves as an annual when sown in autumn-winter. The possibility is discussed of using this information for establishing the coincidence of critical stages with environmental conditions that can limit potential and actual yield through agronomic practices.     

Biomass-density relationship varies with water use efficiency across an aridity gradient

Accumulating evidence has shown that the scaling exponent (α) of the aboveground biomass-density relationship is not a constant value. Debate continues over what determines the variation in α. By measuring foliar stable carbon isotope, plant morphological traits and α along an aridity gradient from eastern to western China, we confirmed that the variation in α was accounted for by changes in plant morphological traits which are adaptive strategies for enhancing water use efficiency during drought stress. This information can be crucial for understanding and predicting community and ecosystem processes.     

Accumulated effect of elevated ultraviolet-B radiation over multiple generations of the arid-environment annual Dimorphotheca sinuata DC. (Asteraceae)

Two D. sinuata populations, established from the same seed batch, were cultured concurrently under ambient (2.5–8.9 kj m^?2 d^?1) and enhanced (4.7–11.4 kj m^?2 d^?1) ultraviolet-B radiation for two consecutive generations. The two populations, designated low- and high-UV-B cultivars, were each grown for an additional generation under both UV-B fluences at different alignments. Differences in plant performance were examined between UV-B treatments (immediate UV-B effects), cultivars (accumulated UV-B effects), alignments and their interactions. Accumulated UV-B had a greater effect on plant performance than immediate UV-B. The former induced earlier reproductive effort, substantial (up to 35%) reductions in dry mass (stems, leaves and reproductive organs), decreased stem and inflorescence production, and diminished steady-state fluorescence yields, chlorophyll a concentrations, pollen tube growth and germination of seeds set. The latter caused only diminished non-photochemical quenching, reduced chlorophyll a, soluble sugar and starch concentrations, decreased pollen germination, and increased carotenoid contents. Interactions between UV-B fluence level and alignment occurred for all measured dry mass parameters, leaf area, photosynthetic pigment and soluble sugar concentrations, and germination success of seeds set. These results indicate altered DNA integrity rather than an indirect UV-B effect of photomorphogenic origin.     

Cytological and morphological differentiation in Caralluma burchardii (Asclepiadaceae)

Meve, U. 1995. Cytological and morphological differentiation in Carallurna burchardii (Asclepiadaceae). — Nord. J. Bot. 15: 459–467. Copenhagen. ISSN 0107–055X.
Caralluma burchardii is revised based on detailed investigations of vegetative and floral morphology, as well as chromosome numbers. The species is now divided into two taxa, the dodecaploid ssp. burchardii , an endemic of the Canary Islands, and ssp. rnaura , a hexaploid from the west coast of Morocco. The high ploidy levels involved, the highest known in the Asclepiadaceae, are discussed in conjunction with biogeographical aspects.