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  • Lang Li (李朗), Bing Liu (刘冰), Yu Song (宋钰), Hong-Hu Meng (孟宏虎), Xiu-Qin Ci (慈秀芹), John G. Conran, Rogier P.J. de Kok, Pedro Luís Rodrigues de Moraes, Jun-Wei Ye (叶俊伟), Yun-Hong Tan (谭运洪), Zhi-Fang Liu (刘志芳), Marlien van der Merwe, Henk van der Werff, Yong Yang (杨永), Jens G. Rohwer, Jie Li (李捷)
    Plant Diversity. 2025, 47(03): 341-364.
    https://doi.org/10.1016/j.pld.2025.04.001
    Over the past two decades, our understanding of Lauraceae, a large family of woody plants, has undergone significant advances in phylogeny, taxonomy, and biogeography. Molecular systematic studies have elucidated the basic relationships within the family with plastid phylogenomic analyses providing robust support for deep-level relationships between Lauraceae lineages, leading to the recognition of nine tribes: Hypodaphnideae, Cryptocaryeae, Cassytheae, Neocinnamomeae, Caryodaphnopsideae, Mezilaureae, Perseeae, Laureae, and Cinnamomeae, with Mezilaureae validated here. Nuclear genomes and comparative genomics studies have also clarified aspects of the family’s evolutionary history and metabolic diversity. Taxonomic studies have focused mainly on the most diverse regions, e.g., tropical Asia, tropical America, and Africa (Madagascar), with six new genera described and five reinstated since the last major overview of the family. The extensive fossil record suggests that Lauraceae diversified globally during the Late Cretaceous and Early Cenozoic. Biogeographic studies indicate that different lineages of the family are sorted into Gondwanan and Laurasian lineages, with patterns resulting from the disruption of boreotropical flora and multiple long-distance dispersal events. Phylogeographic studies, predominantly from East Asia, have shown patterns of in situ survival and demographic stability or expansion during the Quaternary. Nevertheless, many systematic relationships within the family remain unresolved and further research is needed into the complex biogeographic history and ecological roles of Lauraceae. A multifaceted approach integrating genomic studies, field work, morphological and ecological investigations is therefore needed to understand the evolution and diversity of this ecologically and economically significant plant family.
  • Wei Gu, Ting Zhang, Shui-Yin Liu, Qin Tian, Chen-Xuan Yang, Qing Lu, Xiao-Gang Fu, Heather R. Kates, Gregory W. Stull, Pamela S. Soltis, Douglas E. Soltis, Ryan A. Folk, Robert P. Guralnick, De-Zhu Li, Ting-Shuang Yi
    Plant Diversity. 2024, 46(06): 683-697.
    https://doi.org/10.1016/j.pld.2024.07.001
    The angiosperm family Elaeagnaceae comprises three genera and ca. 100 species distributed mainly in Eurasia and North America. Little family-wide phylogenetic and biogeographic research on Elaeagnaceae has been conducted, limiting the application and preservation of natural genetic resources. Here, we reconstructed a strongly supported phylogenetic framework of Elaeagnaceae to better understand inter- and intrageneric relationships, as well as the origin and biogeographical history of the family. For this purpose, we used both nuclear and plastid sequences from Hyb-Seq and genome skimming approaches to reconstruct a well-supported phylogeny and, along with current distributional data, infer historical biogeographical processes. Our phylogenetic analyses of both nuclear and plastid data strongly support the monophyly of Elaeagnaceae and each of the three genera. Elaeagnus was resolved as sister to the well-supported clade of Hippophae and Shepherdia. The intrageneric relationships of Elaeagnus and Hippophae were also well resolved. High levels of nuclear gene tree conflict and cytonuclear discordance were detected within Elaeagnus, and our analyses suggest putative ancient and recent hybridization. We inferred that Elaeagnaceae originated at ca. 90.48 Ma (95% CI = 89.91-91.05 Ma), and long-distance dispersal likely played a major role in shaping its intercontinentally disjunct distribution. This work presents the most comprehensive phylogenetic framework for Elaeagnaceae to date, offers new insights into previously unresolved relationships in Elaeagnus, and provides a foundation for further studies on classification, evolution, biogeography, and conservation of Elaeagnaceae.
  • Fangdong Geng (耿方东), Miaoqing Liu (刘苗青), Luzhen Wang (王璐珍), Xuedong Zhang (张雪栋), Jiayu Ma (马佳雨), Hang Ye (叶航), Keith Woeste, Peng Zhao (赵鹏)
    Plant Diversity. 2025, 47(03): 365-381.
    https://doi.org/10.1016/j.pld.2025.04.002
    Intraspecific genetic variance and gene flow can support the adaptive evolution of species challenged by climate shifts or novel environmental conditions. Less well understood is how genome organization and gene flow interact in closely related species during evolutionary divergence and differentiation. Here we conducted genomic footprint analyses to determine how three species of Pterocarya (P. stenoptera, P. hupehensis, and P. macroptera), which are sympatric but occupy different elevational niches, adapted to the heterogeneous environment of the Qinling-Daba Mountains, China. We identified candidate genes for environmental adaptation (i.e., PIEZO1, WRKY39, VDAC3, CBL1, and RAF), and also identified regions of gene introgression between P. hupehensis and P. macroptera that show lower genetic load and higher genetic diversity than the rest of their genomes. The same introgressed regions are notably situated in areas of minimal genetic divergence yet they are characterized by elevated recombination rates. We also identified candidate genes within these introgressed regions related to environmental adaptation (TPLC2, CYCH;1, LUH, bHLH112, GLX1, TLP-3, and ABC1). Our findings have thus clarified the important role of gene flow in ecological adaptation and revealed genomic signatures of past introgression. Together, these findings provide a stronger theoretical basis for understanding the ecological adaptation and conservation of Quaternary relict woody plants in East Asia.
  • Jiagang Zhao, Shufeng Li, Jian Huang, Wenna Ding, Mengxiao Wu, Tao Su, Alexander Farnsworth, Paul J. Valdes, Linlin Chen, Yaowu Xing, Zhekun Zhou
    Plant Diversity. 2025, 47(01): 1-12.
    https://doi.org/10.1016/j.pld.2024.07.004
    Evergreen broad-leaved forests (EBLFs) are widely distributed in East Asia and play a vital role in ecosystem stability. The occurrence of these forests in East Asia has been a subject of debate across various disciplines. In this study, we explored the occurrence of East Asian EBLFs from a paleobotanical perspective. By collecting plant fossils from four regions in East Asia, we have established the evolutionary history of EBLFs. Through floral similarity analysis and paleoclimatic reconstruction, we have revealed a diverse spatio-temporal pattern for the occurrence of EBLFs in East Asia. The earliest occurrence of EBLFs in southern China can be traced back to the middle Eocene, followed by southwestern China during the late Eocene-early Oligocene. Subsequently, EBLFs emerged in Japan during the early Oligocene and eventually appeared in central-eastern China around the Miocene. Paleoclimate simulation results suggest that the precipitation of wettest quarter (PWetQ, mm) exceeding 600 mm is crucial for the occurrence of EBLFs. Furthermore, the heterogeneous occurrence of EBLFs in East Asia is closely associated with the evolution of the Asian Monsoon. This study provides new insights into the occurrence of EBLFs in East Asia.
  • Kai-Yun Chen, Jin-Dan Wang, Rui-Qi Xiang, Xue-Dan Yang, Quan-Zheng Yun, Yuan Huang, Hang Sun, Jia-Hui Chen
    Plant Diversity. 2025, 47(02): 178-188.
    https://doi.org/10.1016/j.pld.2024.09.004
    The genus Salix is a common component of the Northern Hemisphere dendroflora with important ecological and economic value. However, taxonomy and systematics of Salix is extremely difficult and relationships between main lineages, especially deep phylogenies, remain largely unresolved. In this study, we used genome-skimming, plastome assembly, and single-copy orthologs (SCOs) from 66 Salix accessions, along with publicly available plastome and sequence read archive (SRA) datasets to obtain a robust backbone phylogeny of Salix, clarify relationships between its main lineages, and gain a more precise understanding of the origin and diversification of this species-rich genus. The plastome and SCO datasets resolved Salix into two robust clades, with plastome-based phylogenies lacking inner resolution and SCO offering fully resolved phylogenies. Our results support the classification of Salix into five subgenera: Salix, Urbaniana, Triandrae, Longifoliae and Vetrix. We observed a significant acceleration in the diversification rate within the Chamaetia-Vetrix clade, while Salix exhibited increased rates of diversification spanning from the early Oligocene to the late Miocene. These changes coincided with contemporaneous tectonic and climate change events. Our results provide a foundation for future systematic and evolutionary studies of Salix. Additionally, we showed that genome skimming data is an efficient, rapid, and reliable approach for obtaining extensive genomic data for phylogenomic studies, enabling the comprehensive elucidation of Salix relationships.
  • Haoxing Xie, Kaifeng Xing, Jun Zhou, Yao Zhao, Jian Zhang, Jun Rong
    Plant Diversity. 2025, 47(02): 214-228.
    https://doi.org/10.1016/j.pld.2024.07.009
    Subtropical evergreen broad-leaved trees are usually vulnerable to freezing stress, while hexaploid wild Camellia oleifera shows strong freezing tolerance. As a valuable genetic resource of woody oil crop C. oleifera, wild C. oleifera can serve as a case for studying the molecular bases of adaptive evolution to freezing stress. Here, 47 wild C. oleifera from 11 natural distribution sites in China and 4 relative species of C. oleifera were selected for genome sequencing. “Min Temperature of Coldest Month” (BIO6) had the highest comprehensive contribution to wild C. oleifera distribution. The population genetic structure of wild C. oleifera could be divided into two groups: in cold winter (BIO6 ≤ 0 °C) and warm winter (BIO6 > 0 °C) areas. Wild C. oleifera in cold winter areas might have experienced stronger selection pressures and population bottlenecks with lower Ne than those in warm winter areas. 155 single-nucleotide polymorphisms (SNPs) were significantly correlated with the key bioclimatic variables (106 SNPs significantly correlated with BIO6). Twenty key SNPs and 15 key copy number variation regions (CNVRs) were found with genotype differentiation > 50% between the two groups of wild C. oleifera. Key SNPs in cis-regulatory elements might affect the expression of key genes associated with freezing tolerance, and they were also found within a CNVR suggesting interactions between them. Some key CNVRs in the exon regions were closely related to the differentially expressed genes under freezing stress. The findings suggest that rich SNPs and CNVRs in polyploid trees may contribute to the adaptive evolution to freezing stress.
  • Zheng-Yu Zuo, Germinal Rouhan, Shi-Yong Dong, Hong-Mei Liu, Xin-Yu Du, Li-Bing Zhang, Jin-Mei Lu
    Plant Diversity. 2025, 47(01): 34-52.
    https://doi.org/10.1016/j.pld.2024.07.010
    Dryopteridaceae are the largest fern family and include nearly 20% of extant fern diversity, with 24 currently recognized genera. Recognition and delineation of genera within this family have varied greatly. The three-subfamily classification of Dryopteridaceae was based primarily on molecular phylogenetic relationships but lacked morphological evidence, and the phylogenetic relationships of the subfamilies and genera of Dryopteridaceae are only partially resolved. A comprehensive and robust phylogeny is urgently needed. The heterogeneous morphology of the current members of Dryopteridaceae makes the family and its subfamilies difficult to define by single morphological characteristics or even character combinations. We carried out phylogenetic analyses to reconstruct a highly supported phylogeny of Dryopteridaceae. Our analyses recovered 24 strongly supported clades grouped into seven major clades of Dryopteridaceae. Seven morphological characters including habit, rhizome shape, frond morphology, rachis-costae architecture, appendages on stipe base and lamina, and soral arrangement were found to be informative for identifying different major clades and clades in Dryopteridaceae. Based on phylogenetic reconstruction and morphological analysis, we presented an updated infra-familial classification of Dryopteridaceae with seven subfamilies and 24 genera including four newly proposed subfamilies (Ctenitidoideae, Lastreopsidoideae, Pleocnemioideae, and Polystichopsidoideae). Morphological character combinations of each subfamily are summarized, and a key is provided. Most genera sensu PPG I are recognized, with Stigmatopetris reclassified into Dryopteridoideae and Arthrobotrya considered a synonym of Teratophyllum. A new genus Pseudarachniodes is introduced. This revised classification will serve as a foundational framework for future investigations on taxonomy, biogeography, and diversification of the most species-rich Dryopteridaceae in ferns.
  • Tian-Rui Wang, Xin Ning, Si-Si Zheng, Yu Li, Zi-Jia Lu, Hong-Hu Meng, Bin-Jie Ge, Gregor Kozlowski, Meng-Xiao Yan, Yi-Gang Song
    Plant Diversity. 2025, 47(01): 53-67.
    https://doi.org/10.1016/j.pld.2024.07.008
    Understanding the ecological adaptation of tree species can not only reveal the evolutionary potential but also benefit biodiversity conservation under global climate change. Quercus is a keystone genus in Northern Hemisphere forests, and its wide distribution in diverse ecosystems and long evolutionary history make it an ideal model for studying the genomic basis of ecological adaptations. Here we used a newly sequenced genome of Quercus gilva, an evergreen oak species from East Asia, with 18 published Fagales genomes to determine how Fagaceae genomes have evolved, identify genomic footprints of ecological adaptability in oaks in general, as well as between evergreen and deciduous oaks. We found that oak species exhibited a higher degree of genomic conservation and stability, as indicated by the absence of large-scale chromosomal structural variations or additional whole-genome duplication events. In addition, we identified expansion and tandem repetitions within gene families that contribute to plant physical and chemical defense (e.g., cuticle biosynthesis and oxidosqualene cyclase genes), which may represent the foundation for the ecological adaptation of oak species. Circadian rhythm and hormone-related genes may regulate the habits of evergreen and deciduous oaks. This study provides a comprehensive perspective on the ecological adaptations of tree species based on phylogenetic, genome evolutionary, and functional genomic analyses.
  • Min-Fei Jin, Xiang-Hai Cai, Gao Chen
    Plant Diversity. 2025, 47(02): 169-177.
    https://doi.org/10.1016/j.pld.2024.07.006
    Mimetic seeds attract birds to disperse seeds mainly by mimicking fleshy fruits or arillate seeds, however, they provide little nutritive reward for bird dispersers. The key characteristics of mimetic seeds are conspicuous seed color, hard seed coat, certain toxic secondary metabolites, and perhaps smooth waxy layer. In this review, we discuss the global distribution of mimetic seeds, the interaction of mimetic seeds with bird dispersers, and secondary metabolites that underlie key characteristics of mimetic seeds. Mimetic-seed species mainly occur in the tropics, with large numbers distributed along coastal areas. The interaction between mimetic-seed species and bird dispersers can be antagonistic, mutualistic, or both. These interactions are generally established by conspicuous visual cues and hard tactile cues from mimetic seeds. The formation and variation of key characteristics of mimetic seeds may contribute to the metabolism of several kind of secondary compounds. Here, we also discuss mimetic-seed dispersal in the context of an evolutionary game, and propose several aspects of mimetic-seed dispersal that remain unstudied. While this review is based on preliminary findings and does not account for other potential influencing factors such as climate, it is expected to contribute to an improved understanding of mimetic-seed dispersal.
  • Gulbar Yisilam, Enting Zheng, Chuanning Li, Zhiyong Zhang, Ying Su, Zhenzhou Chu, Pan Li, Xinmin Tian
    Plant Diversity. 2025, 47(02): 201-213.
    https://doi.org/10.1016/j.pld.2025.01.001
    The black wolfberry (Lycium ruthenicum; 2n = 2x = 24) is an important medicinal plant with ecological and economic value. Its fruits have numerous beneficial pharmacological activities, especially those of anthocyanins, polysaccharides, and alkaloids, and have high nutritional value. However, the lack of available genomic resources for this species has hindered research on its medicinal and evolutionary mechanisms. In this study, we developed the telomere-to-telomere (T2T) nearly gapless genome of L. ruthenicum (2.26 Gb) by integrating PacBio HiFi, Nanopore Ultra-Long, and Hi-C technologies. The assembled genome comprised 12 chromosomes with 37,149 protein-coding genes functionally annotated. Approximately 80% of the repetitive sequences were identified, of which long terminal repeats (LTRs) were the most abundant, accounting for 73.01%. The abundance of LTRs might be the main reason for the larger genome of this species compared to that of other Lycium species. The species-specific genes of L. ruthenicum were related to defense mechanisms, salt tolerance, drought resistance, and oxidative stress, further demonstrating their superior adaptability to arid environments. Based on the assembled genome and fruit transcriptome data, we further constructed an anthocyanin biosynthesis pathway and identified 19 candidate structural genes and seven transcription factors that regulate anthocyanin biosynthesis in the fruit developmental stage of L. ruthenicum, most of which were highly expressed at a later stage in fruit development. Furthermore, 154 potential disease resistance-related nucleotide-binding genes have been identified in the L. ruthenicum genome. The whole-genome and proximal, dispersed, and tandem duplication genes in the L. ruthenicum genome enriched the number of genes involved in anthocyanin synthesis and resistance-related pathways. These results provide an important genetic basis for understanding genome evolution and biosynthesis of pharmacologically active components in the Lycium genus.
  • Qinfeng Guo, Hong Qian, Shenhua Qian
    Plant Diversity. 2025, 47(02): 255-263.
    https://doi.org/10.1016/j.pld.2024.11.005
    The future distribution of invading species depends on the climate space available and certain life history traits that facilitate invasion. Here, to predict the spread potential of plant species introduced in North America north of Mexico (NAM), we compiled distribution and life history data (i.e., seed size, life form, and photosynthetic pathways) for 3021 exotic plant species introduced to NAM. We comparatively examined the species’ range size and climate space in both native and exotic regions and the role of key life history traits. We found that large climate space for most exotic plants is still available in NAM. The range sizes in global exotic regions could better predict the current range sizes in NAM than those in global native regions or global native plus exotic regions. C3 species had larger ranges on average than C4 and CAM plants, and herbaceous species consistently showed stronger relationships in range size between native and exotic regions than woody species, as was the case within the C3 species group. Seed size was negatively related to range size both in native regions and in NAM. However, seed size surprisingly showed a positive correlation with global exotic range size and no correlation with the current actual global (native plus exotic) range size. Our findings underline the importance of species’ native distribution and life history traits in predicting the spread of exotic species. Future studies should continue to identify potential climate space and use underappreciated species traits to better predict species invasions under changing climate.
  • Yuxuan Jiang, Fuli Wu, Xiaomin Fang, Haitao Wang, Yulong Xie, Cuirong Yu
    Plant Diversity. 2025, 47(02): 244-254.
    https://doi.org/10.1016/j.pld.2025.01.004
    The utilization of palynological data for plant diversity reconstructions offers notable advantages in addressing the discontinuity of plant fossils in the stratigraphic record. However, additional studies of modern processes are required to validate or refine the accuracy of diversity results obtained from palynological data. In this study, we used a modern pollen dataset of China to compare the accuracy of plant diversity reconstructions using five different palynological diversity indices (i.e., the pollen species number, Berger–Parker index, Simpson diversity index, Hill index, and Shannon–Wiener index) over a large spatial scale. We then identified climate factors that are most strongly correlated with these patterns of plant diversity. We found that the index that most accurately reflects plant diversity is the Shannon–Wiener index. Our analyses indicated that the most effective indices at reflecting plant diversity are the Shannon–Wiener index and Berger–Parker index. Numerical analysis revealed that palynological diversity (measured using the Shannon–Wiener index) was strongly correlated with climatic parameters, in particular average temperature in the coldest month and annual precipitation, suggesting these factors may be primary determinants of plant diversity distribution. We also found that a threshold value of the normalized Shannon–Wiener index (NH = 0.4) approximately aligns with the contour line specifying 400 mm annual precipitation, serving as a rudimentary indicator for assessing plant diversity in arid versus humid climates. This study suggests that pollen diversity indices have remarkable potential for quantitatively reconstructing paleoclimatic parameters.
  • Amos Kipkoech, Ke Li, Richard I. Milne, Oyetola Olusegun Oyebanji, Moses C. Wambulwa, Xiao-Gang Fu, Dennis A. Wakhungu, Zeng-Yuan Wu, Jie Liu
    Plant Diversity. 2025, 47(02): 229-243.
    https://doi.org/10.1016/j.pld.2024.11.004
    Integrative data from plastid and nuclear loci are increasingly utilized to resolve species boundaries and phylogenetic relationships within major angiosperm clades. Debregeasia (Urticaceae), an economically important genus, presents challenges in species delimitation due to its overlapping morphological traits and unstable taxonomic assignments. Here, we analyzed 14 morphological traits and generated 12 data matrices from the plastomes and nrDNA using genome skimming from the nine recognized morphospecies to clarify species boundaries and assess barcode performance in Debregeasia. We also used a universal set of 353 nuclear genes to explore reticulate evolution and biogeographic history of Debregeasia. Plastomes of Debregeasia exhibited the typical quadripartite structure with conserved gene content and marginal independent variations in the SC/IR boundary at inter- and intra-specific levels. Three Debregeasia species were non-monophyletic and could not be discerned by any barcode; however, ultra-barcodes identified the remaining six (67%), outperforming standard barcodes (56%). Our phylogenetic analyses placed Debregeasia wallichiana outside the genus and suggested six monophyletic clades in Debregeasia, although the placement between Debregeasia hekouensis and Debregeasia libera varied. There was extensive trait overlap in key morphologically diagnostic characters, with reticulation analysis showing potentially pervasive hybridization, likely influenced by speciation patterns and overlaps between species ranges. We inferred that Debregeasia crown diversification began at ca. 12.82 Ma (95% HPD: 11.54-14.63 Ma) in the mid-Miocene within Australia, followed by vicariance and later long-distance dispersal, mainly out of southern China. Our findings highlight the utility of genomic data with integrative lines of evidence to refine species delimitation and explore evolutionary relationships in complex plant lineages.
  • Jingyang He, Wenjing Yang, Qinghui You, Qiwu Hu, Mingyang Cong, Chao Tian, Keping Ma
    Plant Diversity. 2025, 47(03): 403-414.
    https://doi.org/10.1016/j.pld.2025.02.001
    Biological collections are critical for the understanding of species distributions and for formulating biodiversity conservation strategies. However, biological collections are susceptible to various biases, including the “road-map effect”, meaning that the geography of biological collections can be influenced by road networks. Here, using species occurrence records derived from 921,233 plant specimens, we quantified the intensity of the “road-map effect” on floristic collections of China, and investigated its relationships with various environmental and socio-economic variables. Species occurrence records mainly distributed in major mountain ranges, while lowlands were underrepresented. The distance of species occurrence records to the nearest road decreased from 19.54 km in 1960s to 3.58 km in 2010s. These records showed significant clustering within 5 km and 10 km buffer zones of roads. The road density surrounding these records was significantly higher than that in random patterns. Collectively, our results confirmed a significant “road-map effect” in the floristic collections of China, and this effect has substantially intensified from the 1960s to the 2010s, even after controlling for the impact of road network expansion. Topographic, climatic and socio-economic variables that determine regional species diversity, vegetation cover and human impact on vegetation played crucial roles in predicting the intensity of the “road-map effect”. Our findings indicate that biological surveys have become increasingly dependent on road networks, a trend rarely reported in published studies. Future floristic surveys in China should prioritize the lowland areas that have experienced stronger human disturbances, as well as remote areas that may harbor more unique and rare species.
  • Kuiling Zu, Fusheng Chen, Yaoqi Li, Nawal Shrestha, Xiangmin Fang, Shahid Ahmad, Ghulam Nabi, Zhiheng Wang
    Plant Diversity. 2024, 46(06): 774-782.
    https://doi.org/10.1016/j.pld.2023.07.007
    Flowering phenology of plants, which is important for reproductive growth, has been shown to be influenced by climate change. Understanding how flowering phenology responds to climate change and exploring the variation of this response across plant groups can help predict structural and functional changes in plant communities in response to ongoing climate change. Here, we used long-term collections of 33 flowering plant species from the Gongga Mountains (Mt. Gongga hereafter), a biodiversity hotspot, to investigate how plant flowering phenology changed over the past 70 years in response to climate change. We found that mean flowering times in Mt. Gongga were delayed in all vegetation types and elevations over the last 70 years. Furthermore, flowering time was delayed more in lowlands than at high elevations. Interestingly, we observed that spring-flowering plants show earlier flowering times whereas summer/autumn plants show delayed flowering times. Non-synchronous flowering phenology across species was mainly driven by changes in temperature and precipitation. We also found that the flowering phenology of 78.8% plant species was delayed in response to warming temperatures. Our findings also indicate that the magnitude and direction of variation in plant flowering times vary significantly among species along elevation gradients. Shifts in flowering time might cause trophic mismatches with co-occurring and related species, affecting both forest ecosystem structure and function.
  • Liansheng Xu, Zhuqiu Song, Tian Li, Zichao Jin, Buyun Zhang, Siyi Du, Shuyuan Liao, Xingjie Zhong, Yousheng Chen
    Plant Diversity. 2025, 47(01): 21-33.
    https://doi.org/10.1016/j.pld.2024.10.003
    Saussurea is one of the largest and most rapidly evolving genera within the Asteraceae, comprising approximately 520 species from the Northern Hemisphere. A comprehensive infrageneric classification, supported by robust phylogenetic trees and corroborated by morphological and other data, has not yet been published. For the first time, we recovered a well-resolved nuclear phylogeny of Saussurea consisting of four main clades, which was also supported by morphological data. Our analyses show that ancient hybridization is the most likely source of deep cytoplasmic-nuclear conflict in Saussurea, and a phylogeny based on nuclear data is more suitable than one based on chloroplast data for exploring the infrageneric classification of Saussurea. Based on the nuclear phylogeny obtained and morphological characters, we proposed a revised infrageneric taxonomy of Saussurea, which includes four subgenera and 13 sections. Specifically, 1) S. sect. Cincta, S. sect. Gymnocline, S. sect. Lagurostemon, and S. sect. Strictae were moved from S. subg. Saussurea to S. subg. Amphilaena, 2) S. sect. Pseudoeriocoryne was moved from S. subg. Eriocoryne to S. subg. Amphilaena, and 3) S. sect. Laguranthera was moved from S. subg. Saussurea to S. subg. Theodorea.
  • Shuran Yao, Weigang Hu, Mingfei Ji, Abraham Allan Degen, Qiajun Du, Muhammad Adnan Akram, Yuan Sun, Ying Sun, Yan Deng, Longwei Dong, Haiyang Gong, Qingqing Hou, Shubin Xie, Xiaoting Wang, Jinzhi Ran, Bernhard Schmid, Qinfeng Guo, Karl J. Niklas, Jianming Deng
    Plant Diversity. 2025, 47(02): 273-281.
    https://doi.org/10.1016/j.pld.2024.09.007
    Studies on plant diversity are usually based on the total number of species in a community. However, few studies have examined species richness (SR) of different plant life forms in a community along large-scale environmental gradients. Particularly, the relative importance (RIV) of different plant life forms in a community and how they vary with environmental variables are still unclear. To fill these gaps, we determined plant diversity of ephemeral plants, annual herbs, perennial herbs, and woody plants from 187 sites across drylands in China. The SR patterns of herbaceous plants, especially perennial herbs, and their RIV in plant communities increased with increasing precipitation and soil nutrient content; however, the RIV of annual herbs was not altered along these gradients. The SR and RIV of ephemeral plants were affected mainly by precipitation seasonality. The SR of woody plants had a unimodal relationship with air temperature and exhibited the highest RIV and SR percentage in plant communities under the harshest environments. An obvious shift emerged in plant community composition, SR and their critical impact factors at 238.5 mm of mean annual precipitation (MAP). In mesic regions (> 238.5 mm), herbs were the dominant species, and the SR displayed a relatively slow decreasing rate with increasing aridity, which was mediated mainly by MAP and soil nutrients. In arid regions (< 238.5 mm), woody plants were the dominant species, and the SR displayed a relatively fast decreasing rate with increasing aridity, which was mediated mainly by climate variables, especially precipitation. Our findings highlight the importance of comparative life form studies in community structure and biodiversity, as their responses to gradients differed substantially on a large scale.
  • Han-Ning Duan, Yin-Zi Jiang, Jun-Bo Yang, Jie Cai, Jian-Li Zhao, Lu Li, Xiang-Qin Yu
    Plant Diversity. 2024, 46(06): 713-722.
    https://doi.org/10.1016/j.pld.2024.06.003
    Genome skimming has dramatically extended DNA barcoding from short DNA fragments to next generation barcodes in plants. However, conserved DNA barcoding markers, including complete plastid genome and nuclear ribosomal DNA (nrDNA) sequences, are inadequate for accurate species identification. Skmer, a recently proposed approach that estimates genetic distances among species based on unassembled genome skims, has been proposed to effectively improve species discrimination rate. In this study, we used Skmer to identify species based on genomic skims of 47 individuals representing 10 out of 13 species of Schima (Theaceae) from China. The unassembled reads identified six species, with a species identification rate of 60%, twice as high as previous efforts that used plastid genomes (27.27%). In addition, Skmer was able to identify Schima species with only 0.5×sequencing depth, as six species were well-supported with unassembled data sizes as small as 0.5 Gb. These findings demonstrate the potential for Skmer approach in species identification, where nuclear genomic data plays a crucial role. For taxonomically difficult taxa such as Schima, which have diverged recently and have low levels of genetic variation, Skmer is a promising alternative to next generation barcodes.
  • Yuan Wang, Ji Suonan, Kun Liu, Yanni Gao, Sihao Zhu, Qian Liu, Ning Zhao
    Plant Diversity. 2025, 47(01): 159-165.
    https://doi.org/10.1016/j.pld.2024.10.001
    Alpine plants possess unique traits to adapt alpine environments. Whether leaf trait relationships of alpine plants can be captured by the two trait dimensions of organ size and resource economics is unknown. We hypothesized that, beyond the trait dimensions of leaf size and resource economics, non-structured carbohydrates (NSC) would reflect a dimension of cold-tolerance in alpine plants. To test this hypothesis, we measured 12 leaf traits critical to leaf construction and growth in 143 species across 7 sites ranging from alpine steppes to alpine meadows along an environmental gradient on the Tibetan Plateau. Furthermore, a cold resistance experiment was conducted at one of these sites to estimate the lethal temperature causing 50% frost damage (LT50) of 11 alpine species. The majority of variations in 12 leaf traits of alpine plants were captured by three trait axes, in which leaf carbon (LCC) and NSC (including leaf starch; LSC and leaf soluble sugars; LSS) were clustered in a new dimension (PC3) beyond leaf size and structure, and resource economics. Although LCC, LSC and LSS all showed negative correlations with mean annual temperature, a significant negative correlation was only found between LSS and LT50. It indicated that PC3 was able to reflect the cold-tolerance of alpine plants to some extent, in which LSS was the most critical trait. The storage and transformation of NSC under stressful conditions could reflect a dimension of long-term metabolic adaptation and cold-tolerance, which is an extension of the resource-utilization strategy beyond construction cost and growth.
  • Zengke Zhang, Wensheng Chen, Zengyan Li, Wentao Ren, Ling Mou, Junyong Zheng, Tian Zhang, Hantang Qin, Liyi Zhou, Bile Sai, Hang Ci, Yongchuan Yang, Shekhar R. Biswas, Enrong Yan
    Plant Diversity. 2025, 47(02): 300-310.
    https://doi.org/10.1016/j.pld.2024.12.001
    The island rule, a general pattern of dwarfism in large species to gigantism in small species on islands relative to mainland, is typically seen as a macroevolutionary phenomenon. However, whether the ecological processes associated with abiotic and biotic factors generate a pattern of plant size variation similar to the island rule remains unknown. We measured plant height for 29,623 individuals of 50 common woody plant species across 43 islands in the Zhoushan Archipelago (8500 years old and yet to undergo major evolutionary adaptation) and the adjacent mainlands in China. We found pronounced variations in plant height, similar to those of the island rule. Interestingly, islands with low resource availability, such as low soil organic matter content and low precipitation, had a high degree of dwarfism; islands experiencing high environmental stress, such as high soil pH, had a high degree of dwarfism; and islands experiencing less plant–plant competition had a high degree of gigantism. The magnitude of plant dwarfism was higher on small and remote islands than on larger and nearer islands. These results highlight the importance of ecological processes associated with abiotic and biotic conditions in shaping the island rule-like patterns of plant size variation. Since our studied archipelago is too young to undergo major evolution, ecological processes likely played a prominent role in generating the observed pattern, challenging the notion that the evolutionary process is the dominant factor underlying the island rule. Future studies on the island rule need to perform experiments to disentangle evolutionary from ecological mechanisms.
  • Javier Hernández-Velasco, José Ciro Hernández-Díaz, Sergio Leonel Simental-Rodríguez, Juan P. Jaramillo-Correa, David S. Gernandt, José Jesús Vargas-Hernández, Ilga Porth, Roos Goessen, M. Socorro González-Elizondo, Matthias Fladung, Cuauhtémoc Sáenz-Romero, José Guadalupe Martínez-Ávalos, Artemio Carrillo-Parra, Eduardo Mendoza-Maya, Arnulfo Blanco-García, Christian Wehenkel
    Plant Diversity. 2025, 47(03): 415-428.
    https://doi.org/10.1016/j.pld.2024.12.006
    The presence of heterozygous individuals in a population is crucial for maintaining genetic diversity, which can positively affect fitness and adaptability to environmental changes. While inbreeding generally reduces the proportion of heterozygous individuals in a population, polyploidy tends to increase the proportion. North American Populus tremuloides is one of the most widely distributed and ecologically important tree species in the Northern Hemisphere. However, genetic variation in Mexican populations of P. tremuloides, including the genetic signatures of their adaptation to a variety of environments, remains largely uncharacterized. The aim of this study was to analyze how inbreeding coefficient (FIS) and ploidy are associated with clonal richness, population cover, climate and soil traits in 91 marginal to small, isolated populations of this tree species throughout its entire distribution in Mexico. Genetic variables were determined using 36,810 filtered SNPs derived from genome re-sequencing. We found that FIS was approximately between 0 and -1, indicating an extreme heterozygosity excess. One key contributor to the observed extreme heterozygosity excess was asexual reproduction, although ploidy levels cannot explain this excess. Analysis of all neutral SNPs showed that asexual reproduction was positively correlated with observed heterozygosity (Ho) but negatively correlated with expected heterozygosity (He). Analysis of outlier SNPs also showed that asexual reproduction was positively correlated with Ho and negatively correlated with He, although this latter correlation was not significant. These findings support the presence of a Meselson effect.
  • Kai Chen, Yan-Chun Liu, Yue Huang, Xu-Kun Wu, Hai-Ying Ma, Hua Peng, De-Zhu Li, Peng-Fei Ma
    Plant Diversity. 2025, 47(03): 382-393.
    https://doi.org/10.1016/j.pld.2025.03.002
    The sugarcane subtribe Saccharinae (Andropogoneae, Poaceae) was established in 1846, but its delimitation has long been debated. Moreover, the relationships among the genera of Saccharinae remain unclear, and there is no consensus on whether Pseudosorghum, a small genus in tropical Asia with only two species, should be included. Here, we performed phylogenomic analyses using whole plastomes (69 of them newly sequenced) from 132 individuals, representing 65 species in 19 related genera. We also built trees with nuclear ribosomal DNA sequences. Our results justify the inclusion of Pseudosorghum, likely also the Eulalia Clade III, in Saccharinae. Furthermore, both morphological and molecular analyses support merging the two Pseudosorghum species. The backbone relationships of the Saccharinae phylogeny were highly supported with four polyphyletic clades of Miscanthus and the inclusion of Narenga and Tripidium rufipilum in Saccharum. Pseudosorghum is moderately supported as sister to the Miscanthus Clade I, while the remaining Tripidium species could be excluded from the subtribe. Saccharinae is estimated to have originated ~3.73 million years ago in East Asia, followed by intercontinental dispersals. Our study provides a comprehensive phylogenetic framework for future taxonomic revisions of this economically important subtribe.
  • Jianchao Liang, Zhifeng Ding, Ganwen Lie, Zhixin Zhou, Zhixiang Zhang, Huijian Hu
    Plant Diversity. 2025, 47(02): 264-272.
    https://doi.org/10.1016/j.pld.2024.07.011
    Understanding how and why assemblage dissimilarity changes along spatial gradient is a great challenge in ecology, because answers to these questions depend on the analytical types, dimensions, and components of beta diversity we concerned. To obtain a comprehensive understanding of assemblage dissimilarity and its implications for biodiversity conservation in the Himalayas, we explored the elevational patterns and determinants of beta diversity and its turnover and nestedness components of pairwise and multiple types and taxonomic and phylogenetic dimensions simultaneously. Patterns of beta diversity and their components of different types and dimensions were calculated based on 96 sampling quadrats along an 1800-5400 m elevational gradient. We examined whether and how these patterns differed from random expectations using null models. Furthermore, we used random forest methods to quantify the role of environmental variables representing climate, topography, and human disturbance in determining these patterns. We found that beta diversity and its turnover component, regardless of its types and dimensions, shown a hump-shaped elevational patterns. Both pairwise and multiple phylogenetic beta diversity were remarkably lower than their taxonomic counterpart. These patterns were significantly less than random expectation and were mostly associated with climate variables. In summary, our results suggested that assemblage dissimilarity of seed plants was mostly originate from the replacement of closely related species determined by climate-driven environmental filtering. Accordingly, conservation efforts should better cover elevations with different climate types to maximalize biodiversity conservation, rather than only focus on elevations with highest species richness. Our study demonstrated that comparisons of beta diversity of different types, dimensions, and components could be conductive to consensus on the origin and mechanism of assemblage dissimilarity.
  • Xue Wang, Xinrui Liu, Shuang Chen, Jiang Zhu, Yanqi Yuan, Rong Zhu, Kaixi Chen, Xue Yang, Xiaochun Wang, Weiyi Mo, Ruili Wang, Shuoxin Zhang
    Plant Diversity. 2025, 47(02): 291-299.
    https://doi.org/10.1016/j.pld.2024.09.008
    Root anatomical traits play an important role in understanding the link between root physiological function and ecological process. To determine how plants change root anatomical traits to adapt to distinct environments, we measured four key root anatomical traits—stele diameter (SD), cortex thickness (CT), root diameter (RD), and the stele to root diameter ratio (SDRD)—of first-order roots of 82 species collected from different vegetation zones along a 2000 m altitudinal gradient on the northern slope of Taibai Mountain. Compared with other altitudes, plants located in temperate birch and fir forests had thinner SD, CT, RD, and SDRD. We found that elevational variation in root anatomical traits could largely be explained by phylogenetic taxonomy (clade). In addition, changes in SD were driven by soil bulk density, whereas variations in CT and RD were influenced by soil available nitrogen. When phylogenetic factors were removed from our analysis, allometric relationships between RD and root anatomical traits (SD and CT) were observed across different altitudes. Our study reveals the influence of phylogeny and environment on the elevational variation in root anatomical traits and further supports the allometric relationship between root anatomical traits (SD and CT) and RD. These findings enhance our understanding of the evolutionary and adaptive mechanisms of root anatomical structures, providing a basis for predicting how root anatomical traits respond to global changes.
  • Yanlei Liu, Kai Chen, Lihu Wang, Xinqiang Yu, Chao Xu, Zhili Suo, Shiliang Zhou, Shuo Shi, Wenpan Dong
    Plant Diversity. 2025, 47(01): 115-126.
    https://doi.org/10.1016/j.pld.2024.10.002
    DNA barcoding has been extensively used for species identification. However, species identification of mixed samples or degraded DNA is limited by current DNA barcoding methods. In this study, we use plant species in Juglandaceae to evaluate an assembly-free reads accurate identification (AFRAID) method of species identification, a novel approach for precise species identification in plants. Specifically, we determined (1) the accuracy of DNA barcoding approaches in delimiting species in Juglandaceae, (2) the minimum size of chloroplast dataset for species discrimination, and (3) minimum amount of next generation sequencing (NGS) data required for species identification. We found that species identification rates were highest when whole chloroplast genomes were used, followed by taxon-specific DNA barcodes, and then universal DNA barcodes. Species identification of 100% was achieved when chloroplast genome sequence coverage reached 20% and the original sequencing data reached 500,000 reads. AFRAID accurately identified species for all samples tested after 500,000 clean reads, with far less computing time than common approaches. These results provide a new approach to accurately identify species, overcoming limitations of traditional DNA barcodes. Our method, which uses next generation sequencing to generate partial chloroplast genomes, reveals that DNA barcode regions are not necessarily fixed, accelerating the process of species identification.
  • Tingting Deng, Qingqing Du, Yan Zhu, Simon A. Queenborough
    Plant Diversity. 2025, 47(02): 282-290.
    https://doi.org/10.1016/j.pld.2025.01.003
    Herbaceous plants are an essential component of forest diversity and driver of ecosystem processes. However, because the growth forms and life-history strategies of herbaceous plants differ from those of woody plants, it is unclear whether the mechanisms that drive patterns plant diversity and community structure in these two plant groups are the same. In this study, we determined whether herb and woody plant communities have similar patterns and drivers of alpha- and beta-diversity. We compared species richness, distribution, and abundance of herbs to woody seedlings in a 20-ha Donglingshan warm-temperate forest (Donglingshan FDP), China. We also determined whether variation in patterns of species richness and composition are better explained by environmental or spatial variables. Herbaceous plants accounted for 72% of all species (81 herbaceous, 31 woody) recorded. Alpha- and beta-diversity were higher in herbs than in woody seedlings. Although alpha-diversity of herbs and woody seedlings was not correlated across the site, the local-site contributions to beta-diversity for herbs and woody seedlings were negatively correlated. Habitat type explained slightly more variation in herb community composition than in woody seedling composition, with the highest diversity in the low-elevation slope. Environmental variables explained the variation in species richness and composition more in herbaceous plants than in woody seedlings. Our results indicate that different mechanisms drive variation in the herb and woody seedling communities, with herbs exhibiting greater environmental sensitivity and habitat dependence. These findings contribute to the better understanding of herbaceous plant diversity and composition in forest communities.
  • Yongli Wang, Yan-Da Li, Shuo Wang, Erik Tihelka, Michael S. Engel, Chenyang Cai
    Plant Diversity. 2025, 47(01): 13-20.
    https://doi.org/10.1016/j.pld.2024.07.007
    Angiosperms experienced one of the most remarkable radiations of land plants and are now the dominant autotrophs in terrestrial ecosystems. Recent phylogenomic studies based on large-scale data from plastid, mitochondrial, or nuclear transcriptomes/genomes and increased taxon sampling have provided unprecedent resolution into the phylogeny of flowering plants. However, owing to ancient rapid radiations, the interrelationships among the five lineages of Mesangiospermae, the vast majority of angiosperms, remain contentious. Here we show that, although plastid and mitochondrial genomes lack sufficient phylogenetic signal for resolving deeper phylogeny, the relationships among five mesangiosperm lineages can be confidently resolved under better-fitting models using genome-scale data. According to our Bayesian cross-validation and model test in a maximum likelihood framework, site-heterogeneous models (e.g., CAT-GTR + G4, LG + C20 + F + G) outperform site-homogeneous or partition models often used in previous studies. Under site-heterogeneous models, the approximately unbiased test favored our preferred tree recovered from various datasets: Ceratophyllales (coontails) are robustly recovered as sister to monocots, and they together are sister to the clade comprising magnoliids, Chloranthales, and eudicots. Our phylogenomic analyses resolve the last enigma of the deeper phylogeny of angiosperms and emphasize the efficacy of modeling compositional heterogeneity in resolving rapid radiations of plants.
  • Pablo C. Guerrero, Jaime Martínez-Harms, Mary T.K. Arroyo, Deren Eaton, Beatriz M. Meriño, Antonio Varas-Myrik, Heidy M. Villalobos-Barrantes, Gastón O. Carvallo
    Plant Diversity. 2025, 47(03): 440-453.
    https://doi.org/10.1016/j.pld.2025.03.001
    Pollination niches, which encompass the factors influencing pollen exchange among angiosperms, are fundamental to understanding reproductive success and the intricate eco-evolutionary dynamics of plants. In this study, we investigated pollination niche shift among four sympatric Eriosyce cacti with restricted distributions in the South American Mediterranean region. Utilizing a comprehensive approach—including pollination niche analysis, pollinator color perception studies, reproductive output assessments, molecular phylogenetics, gene flow analyses, and species distribution modeling—we revealed a significant pollination niche displacement in E. chilensis and its variety, E. chilensis var. albidiflora. These taxa transitioned from a hummingbird-pollinated system prevalent in the Eriosyce sect. Neoporteria, characteristic of their sister species E. litoralis, to a bee-pollination strategy akin to that of the co-occurring species E. mutabilis. This shift highlights a simultaneous convergence of E. chilensis toward E. mutabilis and divergence from E. litoralis in pollination strategies, providing adaptive advantages by reducing pollen limitation and enhancing seed production. The morphological and flowering phenological similarities between E. chilensis and E. mutabilis suggest the evolution of a shared advertising display, potentially indicative of floral mimicry, wherein both species benefit from attracting shared bee pollinators. Genomic analyses reveal distinct pollinator-driven selection pressures, with E. chilensis/E. albidiflora exhibiting traits that promote reproductive isolation from E. litoralis, supporting a scenario of rapid speciation occurring within the past half of million years in the absence of geographic barriers. These findings underscore the pivotal role of pollinator interactions in shaping angiosperm speciation and biodiversity, highlighting their dynamic influence on ecological and evolutionary processes.
  • Ziwei Chen, Dongsheng Zhao, Siqi Deng, Yu Zhu, Ke Wang, Shunsheng Wang, Du Zheng
    Plant Diversity. 2025, 47(03): 394-402.
    https://doi.org/10.1016/j.pld.2024.09.006
    The expansion of road networks in recent decades has drawn considerable attention due to its impact on biodiversity in high-altitude ecosystems. Here, we conducted a comprehensive field survey to investigate the effects of road disturbance on plant diversity in alpine grasslands on the Tibetan Plateau. Our results indicate that road disturbance caused no significant changes in species richness, Shannon-Wiener’s diversity, or Simpson’s diversity, and the alteration in species composition was limited. These findings demonstrate the robust resistance of alpine grassland plant diversity to road disturbance. Plant diversity exhibited more resistance to road disturbance in regions with more hostile environments, such as plateau sub-frigid regions and alpine steppes. Our study suggests that road construction in the Tibetan Plateau poses limited risk to plant biodiversity.
  • Qi Zhang, Jiancan Du, Xiao Han, Yanru Hu
    Plant Diversity. 2024, 46(06): 791-803.
    https://doi.org/10.1016/j.pld.2024.05.003
    Salinity is a severe abiotic stress that affects plant growth and yield. Salinity stress activates jasmonate (JA) signaling in Arabidopsis thaliana, but the underlying molecular mechanism remains to be elucidated. In this study, we confirmed the activation of JA signaling under saline conditions and demonstrated the importance of the CORONATINE INSENSITIVE1 (COI1)-mediated JA signaling for this process. Phenotypic analyses reflected the negative regulation of JASMONATE ZIM-DOMAIN (JAZ) repressors during salinity stress-enhanced JA signaling. Mechanistic analyses revealed that JAZ proteins physically interact with ABSCISIC ACID-RESPONSIVE ELEMENT BINDING FACTOR1 (ABF1), AREB1/ABF2, ABF3, and AREB2/ABF4, which belong to the basic leucine zipper (bZIP) transcription factor family and respond to salinity stress. Analyses on the ABF3 overexpression plants and ABF mutants indicated the positive role of ABF3 in regulating JA signaling under saline condition. Furthermore, ABF3 overexpression partially recovered the JA-related phenotypes of JAZ1-Δ3A plants. Moreover, ABF3 was observed to indirectly activate ALLENE OXIDE SYNTHASE (AOS) transcription, but this activation was inhibited by JAZ1. In addition, ABF3 competitively bind to JAZ1, thereby decreasing the interaction between JAZ1 and MYC2, which is the master transcription factor controlling JA signaling. Collectively, our findings have clarified the regulatory effects of ABF3 on JA signaling and provide new insights into how JA signaling is enhanced following an exposure to salinity stress.
  • Weidong Zhu, Jie Qian, Yingke Hou, Luke R. Tembrock, Liyun Nie, Yi-Feng Hsu, Yong Xiang, Yi Zou, Zhiqiang Wu
    Plant Diversity. 2025, 47(01): 127-135.
    https://doi.org/10.1016/j.pld.2024.11.001
    Single-stranded DNA-binding proteins (SSBs) play essential roles in the replication, recombination and repair processes of organellar DNA molecules. In Arabidopsis thaliana, SSBs are encoded by a small family of two genes (SSB1 and SSB2). However, the functional divergence of these two SSB copies in plants remains largely unknown, and detailed studies regarding their roles in the replication and recombination of organellar genomes are still incomplete. In this study, phylogenetic, gene structure and protein motif analyses all suggested that SSB1 and SSB2 probably diverged during the early evolution of seed plants. Based on accurate long-read sequencing results, ssb1 and ssb2 mutants had decreased copy numbers for both mitochondrial DNA (mtDNA) and plastid DNA (ptDNA), accompanied by a slight increase in structural rearrangements mediated by intermediate-sized repeats in mt genome and small-scale variants in both genomes. Our findings provide an important foundation for further investigating the effects of DNA dosage in the regulation of mutation frequencies in plant organellar genomes.
  • Hong Qian, Oriol Grau
    Plant Diversity. 2025, 47(01): 98-105.
    https://doi.org/10.1016/j.pld.2024.07.005
    Understanding the underlying mechanisms driving species assembly along elevational gradients is of great interest in ecology and biogeography. The Himalaya is one of the world’s hotspots of biodiversity, and the elevational gradient of the central Himalaya in Nepal is one of the longest elevational gradients in the world. Mosses are important constituents of vegetation in the Himalaya, and knowledge of geographic patterns and ecological causes of phylogenetic structure of mosses along elevational gradients in the Himalaya is critical to understanding the assembly of plant diversity in general, and moss diversity in particular, in the Himalaya. Here, we investigate the relationships of phylogenetic structure metrics reflecting different evolutionary depths with elevation and climatic variables representing mean temperature and precipitation conditions, climate extremes, and climate seasonality for mosses distributed along an elevational gradient spanning about 5000 m in the central Himalaya, Nepal. For a given climatic variable, different metrics of phylogenetic structure show different spatial and climatic patterns, but all phylogenetic metrics standardized for species richness show that phylogenetic dispersion in moss assemblages tend to increase with increasing elevation and decreasing temperature. The standardized effect size of mean pairwise distance of moss assemblages shows a triphasic (zig-zag) pattern, which is generally consistent with the triphasic patterns previously found in angiosperms and ferns along the same elevational gradient. Our study shows that temperature-related variables and climate seasonality variables are more important drivers of phylogenetic dispersion in mosses in Nepal, compared with precipitation-related variables and climate extreme variables, respectively.
  • Hong Qian, Shenhua Qian
    Plant Diversity. 2025, 47(01): 82-88.
    https://doi.org/10.1016/j.pld.2024.08.004
    Exploring the worldwide patterns of endemism and the processes that lead to the formation of high-endemism centers is crucial in biogeography. This study examines the geographic distribution and ecological influences on the endemism of liverworts across 390 regions worldwide. We assess phylogenetic endemism and relative phylogenetic endemism in relation to eleven environmental factors, which represent current and Quaternary climate variations, as well as topographic and environmental heterogeneity. Areas with higher endemism in liverworts tend to have higher temperatures, precipitation, and environmental heterogeneity, but lower temperature seasonality and lesser impacts from Quaternary climate changes. Regions exhibiting notably high endemism are predominantly found in tropical Asia, Madagascar, eastern Australia, and the Andes, while those with notably low endemism are generally in temperate Eurasia and North America, parts of Africa, and eastern South America. Centers of neo-endemism are mainly in southern Africa, whereas centers of paleo-endemism are in southern South America, tropical Asia, and New Zealand. Environment variability is a more significant predictor of phylogenetic endemism than current climate conditions, which are themselves more predictive than variables related to Quaternary climate changes. Nevertheless, these three types of explanatory variables combined explain only about one-third of the variance in phylogenetic endemism.
  • Xian Chen, Pyae Phyo Hein, Mengxue Shi, Fen Yang, Jun Yang, Yao Fu, Xuefei Yang
    Plant Diversity. 2025, 47(02): 311-322.
    https://doi.org/10.1016/j.pld.2024.09.011
    The global rise in animal protein consumption has significantly amplified the demand for fodder. A comprehensive understanding of the diversity and characteristics of existing fodder resources is essential for balanced nutritional fodder production. This study investigates the diversity and composition of fodder plants and identifies key species for cattle in Zhaotong City, Yunnan, China, while documenting indigenous knowledge on their usage and selection criteria. Ethnobotanical surveys were conducted in 19 villages across seven townships with 140 informants. Data were collected through semi-structured interviews, free listing, and participatory observation, and analyzed using Relative Frequency Citation. A total of 125 taxa (including 106 wild and 19 cultivated) were reported. The most cited family is Poaceae (27 taxa, 21.43%), followed by Asteraceae (17 taxa, 13.49%), Fabaceae (14 taxa, 11.11%), Polygonaceae (9 taxa, 7.14%) and Lamiaceae (4 taxa, 3.17%). The whole plant (66.04%) and herbaceous plants (84.80%) were the most used parts and life forms. The most cited species were Zea mays, Brassica rapa, Solanum tuberosum, Eragrostis nigra, and Artemisia dubia. Usage of diverse fodder resources reflects local wisdom in managing resource availability and achieving balanced nutrition while coping with environmental and climatic risks. Preferences for certain taxonomic groups are due to their quality as premier fodder resources. To promote integrated crop-livestock farming, we suggest further research into highly preferred fodder species, focusing on nutritional assessment, digestibility, meat quality impacts, and potential as antibiotic alternatives. Establishing germplasm and gene banks for fodder resources is also recommended.
  • Maochou Liu, Wenxiang Wu, Ke Wang, Xinshuai Ren, Xueqin Zhang, Lei Wang, Jing Geng, Bo Yang
    Plant Diversity. 2025, 47(01): 89-97.
    https://doi.org/10.1016/j.pld.2024.11.003
    Latitudinal patterns of tree β-diversity reveal important insights into the biogeographical processes that influence forest ecosystems. Although previous studies have extensively documented β-diversity within relatively small spatial extents, the potential drivers of β-diversity along latitudinal gradients are still not well understood at larger spatial extents. In this study, we determined whether tree β-diversity is correlated with latitude in forests of southeastern China, and if so, what ecological processes contribute to these patterns of tree β-diversity. We specifically aimed to disentangle the relative contributions from interspecific aggregation and environmental filtering across various spatial extents. We delineated regional communities comprising multiple nearby national forest inventory (NFI) plots around random focal plots. The number of NFI plots in a regional community served as a surrogate for spatial extent. We also used a null model to simulate randomly assembled communities and quantify the deviation (β-deviation) between observed and expected β-diversity. We found that β-diversity decreased along a latitudinal gradient and that this pattern was clearer at larger spatial extents. In addition, latitudinal patterns of β-deviation were explained by the degree of species spatial aggregation. We also identified environmental factors that drive β-deviation in these forests, including precipitation, seasonality, and temperature variation. At larger spatial extents, these environmental variables explained up to 84% of the β-deviation. Our results reinforce that ecological processes are scale-dependent and collectively contribute to the β-gradient in subtropical forests. We recommend that conservation efforts maintain diverse forests and heterogeneous environments at multiple spatial extents to mitigate the adverse effects of climate change.
  • Liang Zhang, Zhen-Long Liang, Xue-Ping Fan, Ngan Thi Lu, Xin-Mao Zhou, Hong-Jin Wei, Li-Bing Zhang
    Plant Diversity. 2024, 46(06): 698-712.
    https://doi.org/10.1016/j.pld.2024.08.005
    The Indo-Burma Biodiversity Hotspot is renowned for its rich biodiversity, including that of vascular plants. However, the fern diversity and its endemism in this hotspot have not been well understood and so far, the diversity of very few groups of ferns in this region has been explored using combined molecular and morphological approaches. Here, we updated the plastid phylogeny of the Java fern genus Leptochilus with 226 (115% increase of the latest sampling) samples across the distribution range, specifically those of three phylogenetically significant species, Leptochilus ovatus, L. pedunculatus, and L. pothifolius. We also reconstructed the first nuclear phylogeny of the genus based on pgiC gene data. Based on molecular and morphological evidence, we identified three new major clades and six new subclades, redefined three existing species, discovered a number of cryptic species of the genus, and elucidated the evolution of the three most variable characters. Our divergence time analyses and ancestral area reconstruction showed that Leptochilus originated in the Oligocene and diversified from early Miocene and 15 dispersal events from lower to higher latitudes are identified. The evolution of three most important morphological characters is analyzed in a context of the new phylogeny. Our analysis showed that 30 (59% of total 51) species of Leptochilus occur in Indo-Burma hotspot, 24 (80% of the 30 species) of which are endemic to this hotspot. We argue that the Indo-Burma hotspot should be recognized as a diversity hotspot for ferns.
  • Fu-Qiang Huang, Josep Peñuelas, Jordi Sardans, Scott L. Collins, Kai-Liang Yu, Man-Qiong Liu, Jiu-Ying Pei, Wen-Bin Ke, Jian-Sheng Ye
    Plant Diversity. 2025, 47(03): 479-488.
    https://doi.org/10.1016/j.pld.2025.02.006
    The primary mechanism driving plant species loss after nitrogen (N) addition has been often hypothesized to be asymmetric competition for light, resulting from increased aboveground biomass. However, it is largely unknown whether plants’ access to soil water at different depths would affect their responses, fate, and community composition under nitrogen addition. In a semiarid grassland exposed to 8-years of N addition, we measured plant aboveground biomass and diversity under four nitrogen addition rates (0, 4, 10, and 16 g m-2 year-1), and evaluated plant use of water across the soil profile using oxygen isotope. Aboveground biomass increased significantly, but diversity and shallow soil-water content decreased, with increasing rate of nitrogen addition. The water isotopic signature for both plant and soil water at the high N rate indicated that Leymus secalinus (a perennial grass) absorbed 7% more water from the subsurface soil layer (20-100 cm) compared to Elymus dahuricus (a perennial grass) and Artemisia annua (an annual forb). L. secalinus thus had a significantly larger biomass and was more abundant than the other two species at the high N rate but did not differ significantly from the other two species under ambient and the low N rate. Species that could use water from deeper soil layers became dominant when water in the shallow layers was insufficient to meet the demands of increased aboveground plant biomass. Our study highlights the importance of water across soil depths as key driver of plant growth and dominance in grasslands under N addition.
  • Yufeng Xu, Chenyang Xue, Xuezhi Wang, Lin Meng, Ying Gao, Mengyang Yu, Lin Geng, Meini Shao, Bo Qu
    Plant Diversity. 2025, 47(03): 499-510.
    https://doi.org/10.1016/j.pld.2024.10.005
    Rivers are crucial in the spread of invasive plants. Invasive plants alter their seed traits to adapt to environmental changes and promote invasion. Studying the trait changes in invasive plant seeds may improve the understanding of their propagation mechanisms along the river and provide appropriate control measures. In this study, seven Ambrosia trifida populations along the Liaohe River were used as study subjects. The results showed that the seven A. trifida populations were closely related and exhibited a certain gene exchange, but the absence of evidence of directed gene flow among populations did not confirm that rivers were the medium of seed dispersal of A. trifida. Along the Liaohe River, from top to bottom, the positive view area, length, width, perimeter, and thousand seed weight of A. trifida seeds showed an increasing trend. The total nitrogen and phosphorus contents in the river water of the A. trifida population in the lower reaches of the Liaohe River were higher than those at the other sites. Furthermore, along the river, from top to bottom, the available nitrogen, total nitrogen, total potassium, available potassium, and organic matter contents in the soil in which A. trifida populations grew showed significant increasing trends. River structure, water quality, and soil nutrients had direct and indirect effects on seed morphology. Soil total nitrogen, available potassium, and organic matter had significant positive effects on seed positive view area and perimeter, suggesting that the maternal effect played a critical role in shaping seed morphology. Our analysis showed that soil nutrients along the river may be the primary driver that governs changes in A. trifida seed traits.
  • Yu Feng, Chaochao Yan, Wen-Qin Tu, Yu-Mei Yuan, Jing-Bo Wang, Xiao-Juan Chen, Chang-Qiu Liu, Yundong Gao
    Plant Diversity. 2025, 47(02): 189-200.
    https://doi.org/10.1016/j.pld.2024.12.005
    Species boundaries are dynamic and constantly challenged by gene flow. Understanding the strategies different lineages adopt to maintain ecological and genetic distinctiveness requires employing an integrative species concept that incorporates data from a variety of sources. In this study, we incorporated genetic, ecological, and environmental evidence to assess the extent of speciation or evolutionary divergence within a monophyletic yet dimorphic group (i.e., clade Leucolirion consisting of six species) within the genus Lilium. This clade consists of two lineages that exhibit unexpectedly distinct perianth appearances: whitish trumpet (funnel form, encompassing four species) and orange recurved (reflex form, including two species), respectively, which are separated by completely different pollination syndromes. Transcriptome-based nuclear and plastome datasets indicate that these two lineages are isolated, with only weak ancient gene flow between them. Within each lineage, several taxa with incomplete isolation have diverged, as indicated by weak genetic structure, strong gene flow, and conflicts between nuclear and chloroplast phylogenies, especially in the trumpet lineage. Although these taxa are not entirely independent, our evidence indicates that they are diverging, with recent gene flow disappearing and multiple isolation strategies emerging, such as differences in flowering time and niche specialization. Taken together, our findings suggest that species divergence and maintenance in Lilium are driven by a combination of adaptive and non-adaptive processes, highlighting the complex interplay of historical climate changes, ecological adaptation, and gene flow in shaping biodiversity within this genus.
  • Hong Qian, Jian Wang, Shenhua Qian, Michael Kessler
    Plant Diversity. 2024, 46(06): 723-731.
    https://doi.org/10.1016/j.pld.2024.07.002
    Phylogenetic niche conservatism posits that species tend to retain ancestral ecological traits and distributions, which has been broadly tested for lineages originating in tropical climates but has been rarely tested for lineages that originated and diversified in temperate climates. Liverworts are thought to originate in temperate climates. Mean lineage age reflects evolutionary history of biological communities. Here, using regional liverwort floras across a long latitudinal gradient from tropical to arctic climates in North America, we test the age-component of the temperate niche conservatism hypothesis. Mean genus age (MGA) was estimated for each of 76 regional floras of liverworts. We related MGA to climatic variables for North America as a whole and for its eastern and western parts separately, and used variation partitioning analysis to assess the relative importance of temperature- versus precipitation-related variables and of climate extremes versus seasonality on MGA. We found that older genera of liverworts tend to concentrate in humid regions of intermediate temperatures in the range of 10 ℃-20 ℃, from which liverworts have adapted to and diversified into more arid, colder, and hotter regions, supporting the temperate niche conservatism hypothesis. We also found that across North America the MGA of liverwort assemblages is more strongly affected by precipitation-related variables than by temperature-related variables, and is more strongly affected by climate extremes than by climate seasonality. Geographic patterns of the MGA of liverworts are consistent with the temperate niche conservatism hypothesis, rather than the tropical niche conservatism hypothesis, the latter of which is broadly supported by angiosperms.
  • Zhiliang Yao, Xia Pan, Xin Yang, Xiaona Shao, Bin Wang, Yun Deng, Zhiming Zhang, Qiaoming Li, Luxiang Lin
    Plant Diversity. 2025, 47(01): 106-114.
    https://doi.org/10.1016/j.pld.2024.08.003
    Patterns and drivers of species-genetic diversity correlations (SGDCs) have been broadly examined across taxa and ecosystems and greatly deepen our understanding of how biodiversity is maintained. However, few studies have examined the role of canopy structural heterogeneity, which is a defining feature of forests, in shaping SGDCs. Here, we determine what factors contribute to α- and β-species-genetic diversity correlations (i.e., α- and β-SGDCs) in a Chinese subtropical forest. For this purpose, we used neutral molecular markers to assess genetic variation in almost all adult individuals of the dominant tree species, Lithocarpus xylocarpus, across plots in the Ailaoshan National Natural Reserve. We also quantified microhabitat variation by quantifying canopy structure heterogeneity with airborne laser scanning on 20 1-ha subtropical forest plots. We found that species α-diversity was negatively correlated with genetic α-diversity. Canopy structural heterogeneity was positively correlated with species α-diversity but negatively correlated with genetic α-diversity. These contrasting effects contributed to the formation of a negative α-SGDC. Further, we found that canopy structural heterogeneity increases species α-diversity and decreases genetic α-diversity by reducing the population size of target species. Species β-diversity, in contrast, was positively correlated with genetic β-diversity. Differences in canopy structural heterogeneity between plots had non-linear parallel effects on the two levels of β-diversity, while geographic distance had a relatively weak effect on β-SGDC. Our study indicates that canopy structural heterogeneity simultaneously affects plot-level community species diversity and population genetic diversity, and species and genetic turnover across plots, thus driving α- and β-SGDCs.
  • Xin Wang, Qing-Hong Feng, Zhi-Hua Zeng, Zhi-Qiang Zhang, Jie Cai, Gao Chen, De-Zhu Li, Hong Wang, Wei Zhou
    Plant Diversity. 2025, 47(01): 136-147.
    https://doi.org/10.1016/j.pld.2024.09.009
    Evolutionary transitions from sexual to asexual reproduction should have significant influences on genetic divergence and polymorphism at the genome level. Plant lineages with diverse reproductive systems provide opportunities to investigate this question using comparative approaches and studies of molecular evolution. We investigated evidence for differences among the transcriptomes of 19 Dioscorea species (wild yams) with diverse reproductive systems. These included sexual species, those that propagate primarily by bulbils, and those with mixed sexual and asexual reproductive modes. We examined how transitions between these reproductive systems affected between-species divergence and within-species polymorphism. Primarily asexual species exhibited a reduced efficacy of natural selection and accumulation of deleterious mutations for both divergence and polymorphism. In contrast, species with mixed reproductive strategies involving both seed and clonal reproduction showed no evidence of an increased fixation of harmful mutations at the divergence level, while an accumulation of genetic load present in polymorphism was evident. Our study indicates that the genetic consequences of evolutionary transitions from sexual to predominantly clonal reproduction is likely to depend on both the duration and extent of asexuality occurring in populations.
  • Jianxiang Yang (杨建翔), Guojing Shen (申国境), Jianqiang Wu (吴建强)
    Plant Diversity. 2025, 47(03): 511-521.
    https://doi.org/10.1016/j.pld.2025.03.003
    The parasitic dodder (Cuscuta, Convolvulaceae) species have wide ranges of hosts. However, some plants, including the cultivated tomato (Solanum lycopersicum), have different degrees of resistance to Cuscuta. The cultivated tomato plants activate a strong hypersensitive response (HR) where Cuscuta haustoria penetrate stems of cultivated tomato, but the underlying mechanisms by which the cultivated tomato perceives Cuscuta and activates resistance remain unclear. In this study, we show that the phytohormones jasmonic acid (JA) and salicylic acid (SA) in cultivated tomato stems were highly induced by Cuscuta australis parasitization. Genetic analyses and experiments of supplementation of JA or SA indicated that the JA and SA pathway not only are both required for activation of HR against Cuscuta parasitization but also function in non-HR-based resistance. The Cuscuta Receptor 1 (CuRe1), which is a leucine-rich repeat receptor-like protein, and suppressor of BAK1-interacting receptor kinase (SOBIR1) and SOBIR1-like, two adaptor kinases, are also important for HR-based and non-HR-based resistance. Importantly, we found that the JA and SA pathway both transcriptionally regulate CuRe1. However, in the cure1 mutants, JA and SA levels were still normally induced by C. australis parasitization. We propose a linear model that an unknown receptor perceives Cuscuta parasitization and thus triggers accumulation of JA and SA, which in turn induce the transcription of CuRe1, and CuRe1 and SOBIR1/SOBIR1-like thereby activate HR-based and non-HR-based resistance to Cuscuta. This study underscores the important roles of hormone signaling and resistance (R) genes in host plant-parasitic plant interactions.
  • Zhi-Li Zhou (周知里), Tial C. Ling, Jian-Li Zhao (赵建立), Xin-Zhi Wang (王欣之), Lin-Lin Wang (王林林), Li Li (李莉), Wen-Jing Wang (王雯婧), Dong-Rui Jia (贾东瑞), Zhi-Kun Wu (吴之坤), Xu-Dong Sun (孙旭东), Yong-Ping Yang (杨永平), Yuan-Wen Duan (段元文)
    Plant Diversity. 2025, 47(03): 429-439.
    https://doi.org/10.1016/j.pld.2025.03.004
    Change of flower color can readily lead to a shift in pollinators, potentially causing pollinator mediated reproductive isolation or even speciation. Here, we examined the ecological and evolutionary consequences of flower color polymorphism in Roscoea cautleoides, an alpine ginger with sympatric distribution of purple- and yellow-flowered plants. Variations in pollinator visitation and specialization to the flower color contributed greatly to pre-zygotic reproductive isolation, with post-zygotic isolation also observed in reciprocal pollination. Yellow-flowered plants evolved independently from purple-flowered plants in two populations due to the absence of anthocyanins, as supported by metabolic, expression, and genetic analysis. Despite early genetic divergence between the two-flower-colored plants, highly differentiated genes were associated with reproduction and stress, while highly selective genes were enriched in stress. Our results suggest that parallel loss of anthocyanins leads to flower color polymorphism in different populations of R. cautleoides, with pollinator preference contributing to reproductive isolation and subsequent genetic differentiation, indicating the process of incipient speciation triggered by flower color changes with sympatric distribution.
  • José Luiz Alves Silva, Alexandre Souza, Angela Pierre Vitória
    Plant Diversity. 2024, 46(06): 732-743.
    https://doi.org/10.1016/j.pld.2024.06.004
    Functional diversity (FD) reflects within- and between-site variation of species traits (α- and β-FD, respectively). Understanding how much data types (occurrence-based vs. abundance-weighted) and spatial scales (sites vs. regions) change FD and ultimately interfere with the detection of underlying geoclimatic filters is still debated. To contribute to this debate, we explored the occurrence of 1690 species in 690 sites, abundances of 1198 species in 343 sites, and seven functional traits of the Atlantic Forest woody flora in South America. All FD indices were sensitive and dependent on the data type at both scales, with occurrence particularly increasing α richness and dispersion (occurrence > abundance in 80% of the sites) while abundance increased β total, β replacement, and α evenness (abundance > occurrence in 60% of the sites). Furthermore, detecting the effect of geoclimatic filters depended on the data type and was scale-dependent. At the site scale, precipitation seasonality and soil depth had weak effects on α- and β-FD (max. R2 = 0.11). However, regional-scale patterns of α richness, dispersion, and evenness strongly mirrored the variation in precipitation seasonality, soil depth, forest stability over the last 120 kyr, and cation exchange capacity (correlations > 0.80), suggesting that geoclimatic filters manifest stronger effects at the regional scale. Also, the role of edaphic gradients expands the idea of biogeographical filters beyond climate. Our findings caution functional biogeographic studies to consider the effect of data type and spatial scale before designing and reaching ecological conclusions about the complex nature of FD.
  • Sen-Tao Lyu, Ting-Ting Zou, Qi-Lin Jiang, Xiao-Fan Wang
    Plant Diversity. 2024, 46(06): 783-790.
    https://doi.org/10.1016/j.pld.2023.03.009
    Andromonoecy is a rare sexual system in plants. The function of additional male flowers in andromonoecious species has been widely discussed; however, few studies have taken offspring fitness into account. In addition, little is known about the mechanisms that maintain andromonoecy in autogamous species. In this study, we compared morphology, pollinator preference, pollen production and export, siring ability, natural siring success, hundred seed dry weight, and seed germination rates between male and hermaphroditic flowers in an endangered autogamous andromonoecious species, Sagittaria guayanensis. Male flowers, which are larger than hermaphroditic flowers, required fewer resources to produce. Pollinators visited male flowers more frequently than they visited hermaphroditic flowers. In addition, pollen production and export were higher in male flowers. Hand pollination demonstrated that siring ability did not differ between flower type. However, the natural siring success of male flowers was triple that of hermaphroditic flowers. The seeds sired by male flowers performed better than those sired by hermaphroditic flowers, with greater dry weight and higher germination rate. In conclusion, male flowers may be superior pollen donors for outcrossing. The maintenance of andromonoecy in S. guayanensis may result from the better performance of male flowers in male function compared to that of hermaphroditic flowers.
  • Miaomiao Shi, Ping Liang, Zhonglai Luo, Yu Zhang, Shiran Gu, Xiangping Wang, Xin Qian, Shuguang Jian, Kuaifei Xia, Shijin Li, Zhongtao Zhao, Tieyao Tu, Dianxiang Zhang
    Plant Diversity. 2025, 47(02): 337-340.
    https://doi.org/10.1016/j.pld.2025.01.002
  • Jing Chen, Jingjing Cao, Binglin Guo, Meixu Han, Zhipei Feng, Jinqi Tang, Xiaohan Mo, Junjian Wang, Qingpei Yang, Yuxin Pei, Yakov Kuzyakov, Junxiang Ding, Naoki Makita, Xitian Yang, Haiyang Zhang, Yong Zhao, Deliang Kong
    Plant Diversity. 2025, 47(03): 466-478.
    https://doi.org/10.1016/j.pld.2025.02.004
    Nutrient acquisition through symbiotic ectomycorrhizal fungi is carbon (C) costly but fundamental for plant growth, community, and ecosystem functioning. Here, we examined the functions of roots and mycorrhiza with respect to nutrient uptake after artificially inducing C limitation-seven months after girdling of an ectomycorrhizal tree, Pinus taeda. Root physiological activity (measured as root nitrogen content and root exudation) declined after girdling and was accompanied with 110% and 340% increases in mycorrhizal colonization and extramatrical hyphal length, respectively. Fungi colonizing roots switched to a community characterized by higher C efficiency (lower C cost) of nutrient acquisition (CENA, the amount of nutrient acquisition per unit C cost) and lower network complexity, indicating a tradeoff between CENA and stability of the fungal community. Root transcriptome analysis suggested a shift in metabolic pathways from a tricarboxylic acid cycle decomposition of carbohydrate to lipid biosynthesis to maintain closer associations with mycorrhiza for nutrient cycling after the girdling. By integrating multi-level evidence, including root transcriptome, fungal composition, and network complexity data, we demonstrate an increased dependence on mycorrhiza for nutrient acquisition under the C limitation condition, which is likely due to a shift to fungal community with higher CENA at the cost of lower stability.