of information on mating styles and reproductive fitness of theOpisthopappus , we cannot determine the likely causes for the heterozygote excess and further research is needed.

4.2 Landscape factors determine gene exhchange pattern of Opisthopappus species

Maintaining an adequate migration rate is usually beneficial because the populations of species experiencing limited gene flow would be susceptible to loss of genetic diversity or genetic variation due to inbreeding and drift (Bansept et al., 2021; Kottler et al., 2021). Lower genetic diversity or genetic variation may make species difficult to adapt to changing environmental conditions, leading to local extinctions (Matzrafi et al., 2021; Wellband et al., 2021).
The gene exchange degree within Opisthopappus species present a pattern mainly from north to south. The higher altitude of the northern in Taihang mountains may contribute to some extent to the genes spread of the Opisthopappus species. Meanwhile, the pollens and seeds ofOpisthopappus are small and light, and easily spread with wind. Each year during the breeding period of the Opisthopappus , the wind by Siberia weather in the Taihang Mountains could be an important factor that facilitate the gene transmission from north to south (Wadl et al., 2022).
Although there is gene exchange between two species and four groups, the gene exchange was extremely limited (Fig. 4, 5 and Table S1). This indicated the existence of a gene exchange barrier between O. longilobus and O. taihangensis or among groups (Fig. 6). Whilist, no obvious topographic obstacles were observed at the barrier between Opisthopappus species. Thus, it suggested that the factors hindering gene migration among populations might not be simple topographic landscape factors but other factors of Taihang Mountians landscape (Lu et al., 2021; Zhang et al., 2021). Taihang Mountains have many deep gullies and diverse landforms, which could result in diverse micro-landscape in different regions. This landscape heterogeneity could produce heterogeneous habitats for different populations of species and bring the spatial isolation for gene exchange.On the other hand, gene exchange/flow is often considered a source of maladaptation because it can limit genetic and phenotypic differentiation and reduce mean fitness in a population (Youngblut et al., 2015). Limited gene exchange/flow occurred in O. longilobus and O. taihangensis might be benifical, even when populations are geographically disparate, because gene flow can allow for the spread of environment-specific beneficial alleles that arise in a single population that could bring the differentiation and adaption (Ellstrand, 2014; Morente‐López et al., 2021).
Meanwhile, the gene exchange barrier was located at the juncture of the Hebei, Shanxi, and Henan provinces. The areas of this barrier were suspected to be closely related to human activities. As the flowers ofOpisthopappus are beautiful and have good medicinal value, human picking and planting activities may significantly influence migration rates (Blanco et al., 2021; Crispo et al., 2011; Liao et al., 2021).
Among the landscape factors of Taihang Mountains, eight factors were the key ones, mainly including six climatic factors and two human activity factors. These factors might be heterogeneous in different regions and form a climatic isolation and/or a corridor by human activites for the populations of species, and which indicating that these factors are important for the development and growth of O. longilobus andO. taihangensis (Ma et al., 2018; Shih et al., 2018).
For the two species, their flowering begin from June to September. And their seeds develop during September and November. During the development of breeding organs, they need accumulated temperature and precipitation, thus four climatic factors, average precipitation in August, average precipitation in October, average precipitation in November, and solar radiation in August, would play an important role. Except for the above factors, built-up land and rain-fed cultivated land factors also significantly impacted the Opisthopappus (Table 3). These two human activities are destructive to the environment that has a long-term occupation and large-scale deep-level transformation on the natural landscape especially physical and chemical properties of the land, which would result in the changes of land workability and soil PH.
Based on the results of Samßada and LFMM, average precipitation in November might have the greatest influence on the Opisthopappus(Shih et al., 2018). In the reciprocal transplant, it was found that too much or too little precipitation would lead to seed growth with cavitation. Therefore, it is speculated that the precipitation in November may be regarded as a ecological indicator for theOpisthopappus two species (An et al., 2020; Chen et al., 2021; Wang et al., 2010).
Meanwhile, among the pathways mainly enriched by the identified selected SNPs, MAPK signaling and plant hormone signal transduction pathways were significant ones. The two pathways are directly or indirectly related to resistance of plant. In the growth and development of Arabidopsis thaliana , it has been shown that MAPK signaling pathway was related to environmental response (Fengbo, 2019). Plant hormone signal transduction pathway has been shown to be related to the plant height through affecting the cell elongation (Haruta and Sussman, 2017). Plant height can affect the ability of plants to receive sunlight, which might be one reason why solar radiation in August is an important factor forOpisthopappus species (Wang et al., 2021a). Thus, it could be hypothesized that precipitation, solar light and soil PH might paly a signal role that activate or inhibit the relative gene express, thereby, bring the adaption of two species to the different landscapt in the Taihang Mountains.

Conclusion

Landscape heterogeneity often plays a vital role in the ability of organisms to disperse or not to disperse. Habitat fragmentation, both natural and artificial, adversely affects the connectivity and persistence of populations. Various landscape features of the Taihang Mountains could make the different environmental conditions and fragmentized habitat of each population or species ofOpisthopappus . With the changes of climate, and human activities, landscape heterogeneity lead to the barrier among Opisthopappusspecies and hinder the gene exchange among the population and promote the differentiation and adaptation among populations ever between species.

CRediT authorship contribution statement

Hao Zhang: Methodology, Writing – review & editing. Hang Ye: Methodology, Editing. Li Liu: Methodology. En Zang: Analysis, Investigation. Qiyang Qie: Sorting data, Investigation. Shan He: Investigation. Weili Hao: Analysis, Investigation. Yafei Lan: Investigation. Zhixia Liu: Investigation. Genlou Sun: Supervision, Review & editing. Yiling Wang: Conceptualization, Funding acquisition, Validation, Writing – review & editing.

Declaration of Competing Interest

The authors declare that they do not have any competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgments

This work was supported by the National Natural Science Foundation of China (31970358); Research Project of Shanxi Scholarship Council (2020-090); Natural Science Foundation of Shanxi Province of China (20210302124501); and Graduate Education nnovation Project of Shanxi (2020SY323).

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