Genetic enrichment of the arctic clonal plant Saxifraga cernua at its southern periphery via the alpine sexual Saxifraga sibirica

Isolation of populations at the margins of a species range may lead to decreasing genetic diversity via genetic drift and inbreeding. Hybridization between peripheral populations of two species can, however, counteract genetic impoverishment. The mainly clonal, polyploid plant Saxifraga cernua has a wide arctic distribution but also extends southwards into alpine sites. In the Ural Mountains, its peripheral distribution overlaps with that of its sexually reproducing, diploid relative Saxifraga sibirica, and fertile polyploids of more or less intermediate appearance are found in this overlap zone. We used amplified fragment length polymorphism (AFLP) analysis to address the potential impact of interspecific gene flow on genetic diversity in the peripheral populations. A total of 149 plants from 17 populations along a 1650 km south-north gradient were analysed for 253 markers. The results suggest that three Middle Ural populations containing fertile and morphologically more or less intermediate plants have been affected by hybridization. All of these plants formed a strongly supported (100%) group with S. cernua in a neighbour-joining tree, but their AFLP phenotypes assigned either to S. cernua or to artificial (simulated) F(1) hybrids between S. cernua and S. sibirica in multilocus assignment tests. The three populations were highly diverse with virtually every plant representing a distinct AFLP phenotype, providing additional evidence for formation of later-generation hybrids and/or backcrossing to S. cernua. In contrast, other peripheral populations of S. cernua were typically monoclonal, suggesting that hybridization with S. sibirica can increase genetic diversity in S. cernua at its southern periphery.     

Reproductive strategies in some arctic Saxifraga (Saxifragaceae), with emphasis on the narrow endemic S. svalbardensis and its parental species

Arctic saxifrages show conspicuous reproductive and chromosomal variation. We examined sexual and asexual traits in 43 phytotron-cultivated Svalbard populations of nine species, including the endemic, supposedly entirely asexual and aneupolyploid S. svalbardensis and its parental species, S. cernua and S. rivularis. All species were self-compatible hermaphrodites with low pollen/ovule ratios, including the strongly protandrous S. cernua , which previously has been reported as self-incompatible with an androdioecious mating system. Spontaneous selfing resulted in considerable seed set in several species and a few seeds in S. svalbardensis and S. cernua; hand-selfing and cross-pollination often increased seed set in the two latter species. Self-fertilized seeds of S. svalbardensis and S. cernua were viable and developed into normal, vigorous plants. Saxifraga rivularis and its close relative S. hyperborea were strongly autogamous. The bulbil-reproducing S. svalbardensis and S. cernua showed extreme variation in fertility, probably because of frequent aneuploidy. Many plants of S. cernua were fully fertile, suggesting that although natural seed set rarely has been observed, sexual reproduction is frequent enough to maintain its previously reported high levels of clonal diversity. Some plants of S. svalbardensis were also fairly fertile. This species may have considerable evolutionary potential; sexual events can lead to increasingly fertile genets with euploid chromosome numbers.     

Same parental species, but different taxa: molecular evidence for hybrid origins of the rare endemics Saxifraga opdalensis and S. svalbardensis (Saxifragaceae)

Saxifraga opdolensis was described from Oppdal in southern Norway and hypothesized to have originated as the hybrid S. cernua × rivularis or to have been derived from a S . cernua-like progenitor. We tested these alternative hypotheses using uni- and biparentally inherited molecular markers observed in S. opdalensis and its putative parental species at the type locality: PCR-RFLPs (restriction fragment length polymorphisms in amplified fragments of chloroplast DNA; cpDNA), sequences of the cpDNA intron trnL and the spacer tmL-tmF , and RAPDs (random amplified polymorphic DNAs). The data provided unambiguous support for the hybrid hypothesis. The cpDNA analyses distinguished two well-differentiated chloroplast genomes, one in S. opdalensis and S. rivularis , and another in S. cernua . The majority of the RAPD markers showed distinct additivity in S. opdalensis relative to its postulated parental species. Thus, S. opdalensis has probably originated from a hybrid with S. rivularis as the maternal parent and S. cernua as the paternal parent. We also included S. svalbaardnsis in the present study because previous molecular analyses of Svalbard material have shown that this species had probably also originated as a hybrid between S. cernua and S. rivularis . The chloroplast genome of S. svalbaardnsis was identical to that of S. opdalensis , but the two species differed in many RAF'D markers. Although these two endemics probably have been derived from the same hybrid combination, they are morphologically and genetically distinct and should be referred to separate species. Differences between such independently originated hybrid taxa may result from intraspecific variation in their parental taxa. Saxufraga cernua comprises, for example, several highly divergent evolutionary lineages.     

GENETIC DIVERSITY IN THE MONOECIOUS HYDROPHILE CERATOPHYLLUM (CERATOPHYLLACEAE)

This study surveys genetic variation in two clonal, monoecious, water-pollinated species that differ in their extent of sexuality and distributional range. Electrophoresis was used to quantify allozyme variability in 12 Wisconsin populations of the widespread Ceratophyllum demersum and the rare C. echinatum. Electrophoretic data indicate that populations of both species have low levels of sexual recombination, low levels of variation, and are structured genetically like inbreeding terrestrial plants. Ceratophyllum populations differ from “typical” clonal terrestrial plants by lower genetic diversity, lower proportions of multiclonal populations, and fewer genotypes per population. In two populations where sexual recombination is documented, heterozygosity is low with significant deficiencies. Monoecy in Ceratophyllum may be related to historical evolutionary factors, whereas vegetative reproduction has a greater influence on the genetic population structure of extant populations. The low genetic identity between C. demersum and C. echinatum supports their recognition as distinct species.     

为什么自然条件下沙地柏种群以无性更新为主

 基于调查、实验和文献资料,作者探讨了沙地柏(Sabina vulgaris)种群的繁殖特征和更新特征。1)根原基是沙地柏进行营养繁殖的结构基础,由根原基发育而成的不定根能够吸收充足的土壤资源供给萌生苗需要,因此定居的萌生苗能够独立维持。2)沙地柏同时具有营养繁殖和有性繁殖,因而沙地柏种群可通过无性更新和有性更新实现维持。3)种子质量极差、萌发率低和实生苗存活率低是制约沙地柏种群有性更新的3个“瓶颈”。4)沙地柏的营养繁殖力强、萌生苗存活率高、繁殖体的产生途径多样,这使自然条件下沙地柏种群以无性更新为主。     

Molecular evidence for polyploid origins in Saxifraga (Saxifragaceae): the narrow arctic endemic S. svalbardensis and its widespread allies

The recently described polyploid Saxifraga svalbardensis is endemic to the arctic archipelago of Svalbard. We investigated relationships among four closely related species of Saxifraga in Svalbard and tested three previously proposed hypotheses for the origin of S. svalbardensis: (1) differentiation from the morphologically and chromosomally variable polyploid S. cernua; (2) hybridization between the diploid S. hyperborea and S. cernua; and (3) hybridization between the tetraploid S. rivularis and S. cernua. Fifteen populations were analyzed using random amplified polymorphic DNAs (RAPDs) and nucleotide sequences of the chloroplast gene matK and the internal transcribed spacers (ITS) of nuclear ribosomal DNA (rDNA). RAPD and matK data suggest that S. svalbardensis has originated from a hybrid with S. rivularis as the maternal parent and S. cernua as the paternal parent, possibly a single time, whereas ITS data could not be used to discriminate among the hypotheses. The data also suggest that the diploid S. hyperborea is a progenitor of the tetraploid S. rivularis. The four populations examined of S. svalbardensis were virtually identical for RAPD and ITS markers, whereas S. cernua showed high levels of variation, suggesting that the latter polyploid either has formed recurrently or has undergone considerable differentiation since its origin.     

Population genetic structure of native versus naturalized sympatric shrub willows (Salix; Salicaceae)

Vegetative propagation of an introduced species can contribute significantly to its ability to spread and become naturalized, potentially in competition with native species. This study focused on the naturalization of a willow shrub, Salix purpurea, which was introduced to the United States from Europe and is commonly sympatric with the native shrub willow, S. eriocephala. Both species are capable of vegetative and sexual reproduction, but little is known about their relative frequency, nor the impact of clonal propagation on population-level genetic diversity. We analyzed genotypes at several microsatellite loci in 993 individuals belonging to 30 subpopulations of S. eriocephala and 28 subpopulations of S. purpurea in areas of sympatry across three watersheds to compare their genetic diversity and genetic structure. Our results revealed six subpopulations of S. purpurea containing plants with identical multilocus genotypes, while clonal individuals were rare among S. eriocephala populations. These species are dioecious with relatively high levels of heterozygosity, but S. eriocephala had much higher allelic diversity and genotypic diversity than did S. purpurea. These results strongly suggest that vegetative propagation has contributed to the naturalization of S. purpurea and has resulted in higher levels of genetic differentiation among S. purpurea populations than among native S. eriocephala populations.     

Female sterility in Ulmus minor (Ulmaceae): a hypothesis invoking the cost of sex in a clonal plant

A high incidence of individuals with low seed set was found in two populations of the field elm Ulmus minor, a European tree that reproduces sexually and via vegetative propagation through root sprouting. One population was a seminatural stand, while the other was established by artificial propagation of genotypes sampled widely across Spain. The low seed set in both populations was due to both pre- and post-zygotic factors, the importance of which vary between genotypes. These factors included gynoecial malformations that produced a non-ovulated pistil, early gynoecial necrosis (i.e., necrosis before any opportunities for pollination), and seed abortion. Female sterility gave rise to two classes of individuals: trees that were largely female-sterile but dispersed normal quantities of viable pollen, and trees that dispersed both normal pollen and substantial numbers of seeds. Reduced production of protein-rich seeds may increase the resource availability for clonal propagation, helping to maintain female-sterile individuals with hermaphrodites.     

Microhabitat distribution of two Florida scrub endemic plants in comparison to their habitat-generalist congeners

Habitat-specialist species may be restricted to a narrower range of microhabitats than habitat-generalist species. We addressed this hypothesis by comparing microhabitats of two pairs of congeners that differ in habitat specificity and co-occur in one distinct habitat type, Florida rosemary scrub. We characterized microhabitats of rosemary scrub specialists, Polygonella basiramia and Lechea cernua, their habitat-generalist congeners, Polygonella robusta and Lechea deckertii, and random points in the rosemary scrub habitat. Plants of both habitat specialists occurred in microhabitats with significantly more bare sand than plants of habitat-generalist species and random points. Plants of all four species occurred in microhabitats that were farther from dominant shrubs, Ceratiola and Quercus spp., than random points. Seedlings of both habitat specialists grew larger in bare sand microhabitats, whereas ground lichens and litter did not affect seedling growth of the habitat generalists. As the time since fire increases, bare sand cover decreases, Ceratiola density increases, Quercus density remains constant, and shrubs become taller. Physical characteristics, such as soil temperature, soil carbon, and soil moisture, differ slightly with respect to microhabitat. Our results suggest that P. basiramia and L. cernua are specialized on bare sand microhabitats that characterize their preferred habitat, rosemary scrub. Microhabitat specialization may limit the distribution of these rare species.     

Variation within some Nordic Saxifraga species (Saxifragaceae)

By means of chemometric and morphometric analyses, the North and South Norwegian populations of Saxifraga paniculata are shown to be different from each other. No difference could be found between one North Norwegian and Icelandic populations of S. paniculata ; the name var. neogaea should not be used for the Icelandic plants. The enigmatic S . × blyttii from Western Norway is most probably an aberrant form of S. paniculata. S. hyperborea and S. rivularis were compared by means of morphometric analyses and a study of pollen size. Presence/ absence of stolons and pollen size are apparently sufficient as differential characters. No Icelandic material could be determined as S. hyperborea . By means of morphometric analyses, S × opdalensis was compared to its putative parents, S. cernua and S. rivularis , and shown to be well separated from them.     

Amplified fragment length polymorphism versus random amplified polymorphic DNA markers: clonal diversity in Saxifraga cernua

Random amplified polymorphic DNA (RAPD) markers are sensitive to changes in reaction conditions and may express polymorphisms of nongenetic origin. Taxa with variable chromosome numbers are particularly challenging cases, as differences in DNA content may also influence marker reproducibility. We addressed these problems by comparing RAPD and amplified fragment length polymorphism (AFLP) analyses of clonal identity and relationships in a chromosomally variable arctic plant, the polyploid Saxifraga cernua, which has been thought to be monoclonal over large geographical distances. Fifty-seven plants from four Greenland populations were analysed using a conservative scoring approach. In total, 26 AFLP and 32 RAPD multilocus phenotypes (putative clones) were identified, of which 21 were identical and each of the remaining five AFLP clones was split into two to three very similar RAPD clones. This minor difference can be explained by sampling error and stochastic variation. The pattern observed in Greenland corroborates our previous results from Svalbard, suggesting that rare sexual events in S. cernua are sufficient to maintain high levels of clonal diversity even at small spatial scales. We conclude that although AFLP analysis is superior in terms of efficiency, RAPDs may still be used as reliable markers in small low-tech laboratories.     

Reproductive challenges of a rare grass,Calamagrostis porteri subsp. insperata (Swallen) C. Greene: implications for habitat restoration

Background: Habitat management for reproductively challenged rare species is a problem when there is insufficient knowledge of their autecology. This study investigated reproductive failure in the rare grass Calamagrostis porteri ssp. insperata (Swallen) C. Greene (Reed bentgrass). Does the management recommendation of high light stimulate clonal growth, flowering, and seed production? Location: Shawnee National Forest, IL, USA, and in a greenhouse and an experimental garden at Southern Illinois University, Carbondale, IL, USA. Methods: Clones obtained from the three known Illinois populations were grown in a glasshouse under experimental light and soil moisture treatments. After 3 years, plants from the high light treatment were planted outside in an experimental garden where the light treatments were maintained for two more years. In the field, vegetative and flowering tiller density, canopy cover, and associated biotic and abiotic variables including abundance of co‐occurring plant species were monitored for 5 years. The overhead tree canopy was cleared over a portion of one population. Results: In the glasshouse, plants increased in size under high light and moist soil, and there were size differences among populations. Sixty‐six per cent (20 of 30) of the genets flowered when planted outdoors under full sunlight but did not produce seed. In the field, flowering only occurred in Calamagrostis growing in the cleared area, but no seed were produced. The plants in the flowering population were smaller than plants in the other two populations. The herbaceous community associated with Calamagrostis in the open diverged from the communities remaining under the shade. Conclusions: This study highlights the difficulty of managing reproductively challenged rare species. Calamagrostis populations can be managed to enhance clonal growth, but establishment of new populations would require translocation of vegetative material as it is highly unlikely that seed can be obtained.     

Female sterility associated with increased clonal propagation suggests a unique combination of androdioecy and asexual reproduction in populations of Cardamine amara (Brassicaceae)

Background and Aims The coexistence of hermaphrodites and female-sterile individuals, or androdioecy, has been documented in only a handful of plants and animals. This study reports its existence in the plant species Cardamine amara (Brassicaceae), in which female-sterile individuals have shorter pistils than seed-producing hermaphrodites.Methods Morphological analysis, in situ manual pollination, microsatellite genotyping and differential gene expression analysis using Arabidopsis microarrays were used to delimit variation between female-sterile individuals and hermaphrodites.Key Results Female sterility in C. amara appears to be caused by disrupted ovule development. It was associated with a 2.4- to 2.9-fold increase in clonal propagation. This made the pollen number of female-sterile genets more than double that of hermaphrodite genets, which fulfils a condition of co-existence predicted by simple androdioecy theories. When female-sterile individuals were observed in wild androdioecious populations, their ramet frequencies ranged from 5 to 54 %; however, their genet frequencies ranged from 11 to 29 %, which is consistent with the theoretically predicted upper limit of 50 %.Conclusions The results suggest that a combination of sexual reproduction and increased asexual proliferation by female-sterile individuals probably explains the invasion and maintenance of female sterility in otherwise hermaphroditic populations. To our knowledge, this is the first report of the coexistence of female sterility and hermaphrodites in the Brassicaceae.     

Sex after all: high levels of diversity detected in the arctic clonal plant Saxifraga cernua using RAPD markers

Arctic plants in general and arctic clonal plants in particular have often been assumed to contain low levels of genetic diversity. We used RAPDs (random amplified polymorphic DNAs) to investigate genetic diversity in the arctic-alpine Saxifraga cernua , which mainly reproduces clonally via bulbils, at three spatial scales in Svalbard: (i) 'macroscale', between two sites 11 km apart; (ii) 'mesoscale', along two crossing transects at each site; and (iii) 'microscale', within a 3 × 3 m square at each site. Thirteen putative clones (RAPD phenotypes) were distinguished among 93 ramets based on 38 RAPD markers. The genetic diversity ( D ; mean 0.52, range 0.10–0.81) and evenness ( E ; mean 0.42, range 0.00–0.82) were at the same level as in clonal plants in general. However, the diversity strongly depended on site and spatial scale. Several clones were highly divergent and clustered independently of site in UPGMA and PCO analyses. In an analysis of molecular variance ( AMOVA ), most of the variation (59%) was found within sites. Mantel tests revealed no correlation between spatial and genetic distance within sites. Our results suggest that occasional sexual reproduction as well as clonal migration via bulbil dispersal play a significant role in the treeless arctic environment, where S. cernua is widespread and locally very abundant. In contrast, Bauert et al. ( Molecular Ecology 7, 1519–1527) found no genetic variation within populations or regions of the Alps, where the species has highly isolated occurrences.     

Effects of local density on insect visitation and fertilization success in the narrow-endemic Centaurea corymbosa (Asteraceae)

Plant population size and density can influence the interactions between plants and pollinators, and affect plant reproductive success. We investigated the effects of local conspecific density on insect visitation and fertilization success in the rare, cliff-dwelling, self-incompatible Centaurea corymbosa , in which fecundity plays a key role in variation in population growth rate among years and among the six extant populations. Plant size, capitulum size, the abundance of co-flowering species, and the weather conditions were added as covariates in the analyses. Over three populations and two years (1995 and 2002), fertilization rate was positively related to the number of flowering conspecifics within 10 m. Fertilization rate varied among populations, but this variation over all six populations in 2002 could not be attributed to differences in population size. Observations in one population in 2003 showed that there was no difference in insect visitation per capitulum between plants in sparse vs dense patches whilst plants from sparse patches suffered reduced fertilization rate. Visitation and fertilization rates were not affected by plant size and the abundance of co-flowering species, but weather conditions at the time of flowering had a strong effect on both variables. Capitulum size had a positive effect on visitation rate, but an effect on fertilization rate only in 1995 and 2002 and not in 2003. Our results suggest that pollen limitation affects fertilization rate in C. corymbosa due to limited compatible mate availability rather than pollinator limitation. They agree with previous genetic results derived from paternity analysis. Whether or not the benefits of local spatial agregation to reproductive success result in increased individual fitness will depend on the relative reduction in survival of vegetative stages due to intra-specific competition.     

Regeneration capacity of oceanic-montane liverworts: implications for community distribution and conservation

Abstract

Scotland’s mountains are home to a rare and unique liverwort community, ‘the oceanic-montane liverwort-rich heath’, but its component species are absent from regions where they could potentially thrive. Many biological characteristics of these species are unknown, making it impossible to explain the reasons for their rarity; however, they have not been observed to produce sporophytes within Britain. We use ex situ cultivation of whole liverworts and fragments, and in situ cultivation of fragments, to assess the growth rate and the potential for vegetative reproduction of several species. Most of the species grew from both fragments and as whole plants, indicating that the rarity of the liverwort heath is not due to poor powers of regeneration. We propose that growth rate and the potential to regenerate from fragments are important factors structuring the liverwort heath community, at least locally. Furthermore, this study demonstrates that there is potential for ex situ conservation of rare liverwort species, in situ enhancement of existing populations, and creation of new ones.     

Seedling demography in an alpine ecosystem

Seedling establishment has long been believed to be rare on alpine tundra because of predicted life history trade-offs, the clonality of alpine species, and the harshness of the alpine climate. Contrary to the idea that seedlings are rare on alpine tundra, a 4-yr demographic study of seedlings at Niwot Ridge, Colorado, USA, found seedlings at high densities, particularly in wetter plant communities. Higher germination densities were associated with higher soil moistures both across communities and across time. Mortality of seedlings was highest in the first year and decreased in subsequent years. Species' abundances differed between seedling and adult populations. Many forbs that lacked vegetative reproduction were significantly more abundant among seedling populations, and many monocots and clonal forbs were more abundant among adult populations. In a comparison with published demographic rates, seedling recruitment and mortality rates of Niwot Ridge species fell above or within rates for a wide range of perennial species. Therefore, germination and seedling establishment stages are no more limiting to sexual reproduction in alpine plants than in other perennial plants.     

Lack of reproductive plasticity in alpine Saxifraga rhomboidea (Saxifragaceae)

Phenotypic plasticity is predicted to be highly advantageous to alpine plants because of the extreme spatial and temporal heterogeneity of the alpine environment. Alpine plants are constrained in their vegetative plasticity by developmental preformation, however, they have the potential for reproductive plasticity via abortion of reproductive structures. Saxifraga rhomboidea Greene (Saxifragaceae) is a perennial of alpine dry meadows. It lacks a mechanism for vegetative reproduction, therefore, seed production is particularly important in maintaining populations of this species. Using long-term fertilized plots at the Niwot Ridge Long-Term Ecological Research Site, we ask if fertilization has changed the abundance of S. rhomboidea , and if S. rhomboidea has shown a plastic reproductive response to nutrient additions. Despite a pronounced increase in community biomass, we found that abundance of S. rhomboidea was not affected by fertilization. Moreover, S. rhomboidea showed no plastic response in seed to ovule ratio or other reproductive measures. Other arctic and alpine species have shown the potential for plasticity of maternal investment; S. rhomboidea appears to lack this reproductive plasticity. We suggest that morphological constraints may limit plasticity in this species.     

Vegetative composition and community structure associated with beaver ponds in Canaan valley, West Virginia, USA

Beavers (Castor canadensis) can cause dramatic changes in vegetative composition and diversity. Although alterations by beaver have been studied extensively, little attention has been paid to the effects beaver impoundments have on rare plants. Effective conservation of riparian and wetland rare plant species must consider the responses of vegetation to changes in hydrology that can occur when beaver populations are present. The goal of this research was to establish the occurrence of locally rare plant species, examine community composition, and analyze vegetative community structure of vegetation associated with beaver ponds in Canaan Valley, West Virginia, USA. Species richness and diversity were similar between plots located inside beaver ponds and adjacent to beaver ponds (P > 0.05). Although no significant difference in rare plant species was detected among pond ages, the oldest ponds (>56 years) had twice as many rare species as the youngest ponds (≤6 years). The youngest ponds had higher overall mean species richness (S) than ponds 7–56 years old (P < 0.05), but S returned to similar levels in the oldest ponds. Of the 15 rare species observed, most were classified as obligate (9) or facultative wetland (4) species. The youngest ponds contained the fewest number of rare species. Multiple response permutation procedure (MRPP) analyses of community structure detected no relation between community composition and either pond age or size. However, both MRPP and non-metric multi-dimensional scaling showed proximity to pond was important in herbaceous community structure. Wetlands in beaver ponds also were shown to be distinct from adjacent wetland areas. Conservation of existing beaver populations is necessary so that the entire spectrum of pond ages is available for the maintenance of rare plant species and communities.     

Abundances of thrips on plants in vegetative and flowering stages are related to plant volatiles

Western flower thrips (WFTs), Frankliniella occidentalis Pergrande, and onion thrips (OTs), Thrips tabaci Lindeman, are two cosmopolitan insect pests of agricultural and horticultural plants. Understanding the occurrence and development of thrips on plants is crucial for identifying suitable plants that can be used for developing a “push-pull” strategy against thrips. In this study, the dynamics of WFTs and OTs on plants (Allium fistulosum L., Medicago sativa L., Luffa cylindrica (L.) Roem., Ocimum basilicum L., and Schizonepeta tenuifolia (Benth.) Briq.) were investigated for two consecutive years (2018–2019). Throughout the survey, the abundances of both thrips species were strongly associated with plant species and plant phenology; both thrips species were present at relatively high densities on M. sativa but very low densities on O. basilicum and S. tenuifolia. Populations of both thrips species greatly increased during plant flowering. A Y-tube olfactory test was used to study the effects of plant volatiles in mediating thrips behaviour and showed that volatiles of M. sativa were attractive to both thrips species whether emitted by the plant in the vegetative or flowering stage, while volatiles of O. basilicum and S. tenuifolia were repellent to thrips. Additionally, because of the presence of a high number of floral chemicals, both thrips species exhibited a greater preference for volatiles emitted by plants in the flowering period over those emitted by plants in the vegetative period. Our observations indicate that plant species and flowering status play an important role in the abundance dynamics of thrips and that the volatiles of flowering plants attract thrips more strongly than volatiles emitted by vegetative plants. These findings can facilitate the screening of attractive/unattractive plants for developing push-pull strategies to control thrips.