Table of Contents
  • Volume 61 Issue 4

    Cover illustration: Cover illustration: The four maps show geographic patterns of standardized effect sizes of mean family stem age and mean family phylogenetic fuse for angiosperm assemblages in China. See Figure 3 of the article by Qian and Deng, pp. 563–571 in this issue, for details. The background photo shows an alpine vegetation community on the Qinghai-Tibetan Plateau. Photo by Tao Deng. Designed by Tao Deng & Hong Qian.
      
      Issue Information
    • 2023, 61 (4): 0–4
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    • Editorial
    • Song Ge, Jun Wen
      2023, 61 (4): 561–562
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    • Research Articles
    • Hong Qian and Tao Deng
      2023, 61 (4): 563–571
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      Deep evolutionary histories can play an important role in assembling species into communities, but few studies have explored the effects of deep evolutionary histories on species assembly of angiosperms (flowering plants). Here we explore patterns of family divergence and diversification times (stem and crown ages, respectively) and phylogenetic fuses for angiosperm assemblages in 100?×?100?km grid cells across geographic and ecological gradients in China. We found that both family stem and crown ages of angiosperm assemblages are older in southeastern China with warm and humid climates than in northwestern China with cold and dry climates; these patterns are stronger for family stem ages than for family crown ages; families in colder and drier climates are more closely related across the family-level angiosperm phylogeny; and family phylogenetic fuses are, on average, longer for angiosperm assemblages in warm and humid climates than in cold and dry climate. We conclude that the fact that deep evolutionary histories, which were measured as family stem and crown ages and family phylogenetic fuses in this study, have shown strong geographic and ecological patterns suggests that deep evolutionary histories of angiosperms have profound effects on assembling angiosperm species into ecological communities.
    • Geographic patterns of standardized effect sizes of mean family stem age and mean family phylogenetic fuse. Standardized effect sizes of mean family stem age (a) and mean family phylogenetic fuse (b) based on the data of the presence or absence of each family in each assemblage; standardized effect sizes of mean family stem age (c) and mean family phylogenetic fuse (d) based on the data of species richness of each family in each assemblage.
    • Nan Song and Heng Zhang
      2023, 61 (4): 572–586
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      Hemiptera is the largest order of non-holometabolous insects. Recent phylogenetic studies have contributed greatly to our understanding of the phylogeny of the group. However, parts of the hemipteran tree remain unclear. In this study, we have sequenced transcriptomes from five sternorrhynchan species, four exemplars from Aphididae, and one from Monophlebidae. The addition of published transcriptome data produced alignments of >105?000 amino acids from 224 hemipteran taxa in 30 superfamilies covering all suborder and infraorder clades. The data confirmed the Sternorrhyncha superfamily relationship of Aleyrodoidea?+?(Psylloidea?+?[Coccoidea?+?Aphidoidea]). In addition, phylogenomic analyses supported a basal split of Sternorrhyncha and the remaining hemipteran lineages. The Fulgoromorpha formed a sister clade to Cicadomorpha, with strong support. Therefore, our results supported the monophyly of Auchenorrhyncha. The Coleorrhyncha was robustly resolved as a sister group of Auchenorrhyncha, contrary to the concept of the Heteropterodea. Within Heteroptera, the Enicocephalomorpha, Dipsocoromorpha, and Gerromorpha were resolved as the earliest branching groups, while the Nepomorpha, Leptopodomorpha, Cimicomorpha, and Pentatomomorpha were the successive sister groups to the remaining clades in the series. The Cimicomorpha and Pentatomomorpha were sister groups that occupied the most apical branches of the heteropteran tree. Although some phylogenetic estimates yielded conflicting hypotheses for superfamily relationships of Cicadomorpha, Four-cluster Likelihood Mapping (FcLM) analyses revealed a greater phylogenetic signal for the relationship of Membracoidea?+?(Cicadoidea?+?Cercopoidea).
    • Maximum likelihood tree inferred from the supermatrix of 240-taxa_50% using IQ-TREE under an LG model. Numbers around nodes are ultrafast bootstrap values (left) from maximum likelihood analysis and posterior probability values (right) from Bayesian inference analysis. “-” indicates the nodes unsupported by ExaBayes analysis based on the same dataset. In this tree, Sternorrhyncha was sister to all other hemipteran lineages. The Aleyrodoidea was sister to a clade (Psylloidea + [Coccoidea + Aphidoidea]). The monophyly of Auchenorrhyncha was strongly supported. The Coleorrhyncha was robustly resolved as a sister group of Auchenorrhyncha.
    • Lu‐Shui Zhang, Chun‐Lin Chen, Xing‐Xing Mao, and Jian‐Quan Liu
      2023, 61 (4): 587–598
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      Many species of the subfamily Polygonoideae are economically important. However, phylogenetic relationships and taxonomic treatments of these species remain disputed. In this study, we used highly orthologous nuclear genes and plastome sequence variation extracted from transcriptomes from 98 species of 26 genera of Polygonaceae mainly from the subfamily Polygonoideae to construct a robust phylogeny. We discerned six successively diverged and well-defined clades and both nuclear and plastome phylogenies are highly consistent with each other in this subfamily. Phylogenetic relationships between all clades and subclades were well resolved within Polygonoideae. Our analyses revealed that the shrub tribe Atraphaxideae and the herbal genera Polygonum, Persicaria, and Fallopia are polyphyletic. The sampled species of Polygonoideae started to diversify around the Cretaceous/Tertiary boundary (70?Ma) when the global climate exhibited large oscillations. Further origins of more herbal and woody species were found to have clearly increased during later climatic oscillations. We found that woody habits, especially shrubs, originated multiple times from ancestral herbs in this subfamily. Local dry climates may have favored such habit shifts from ancestral herbs. Our results deepen our understanding of evolutionary diversification of Polygonoideae.
    • In this study, we used highly orthologous nuclear genes and plastome sequence variation extracted from transcriptomes from 98 species of 26 genera of Polygonaceae mainly from the subfamily Polygonoideae to construct a robust phylogeny. We discerned six successively diverged and well-defined clades and both nuclear and plastome phylogenies are highly consistent with each other in this subfamily. Phylogenetic relationships between all clades and subclades were well resolved within Polygonoideae. Our analyses revealed that the shrub tribe Atraphaxideae and the herbal genera Polygonum, Persicaria, and Fallopia are polyphyletic. The sampled species of Polygonoideae started to diversify around the Cretaceous/Tertiary boundary (70 Ma) when the global climate exhibited large oscillations. Further origins of more herbal and woody species were found to have clearly increased during later climatic oscillations. We found that woody habits, especially shrubs, originated multiple times from ancestral herbs in this subfamily. Local dry climates may have favored such habit shifts from ancestral herbs. Our results deepen our understanding of evolutionary diversification of Polygonoideae.
    • Li‐Jun Yan, Zhi‐Guo Zhu, Pei Wang, Chao‐Nan Fu, Xi‐Jin Guan, Philip Kear, Chun‐Zhi Zhang, and Guang‐Tao Zhu
      2023, 61 (4): 599–612
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      Common potato (Solanum tuberosum L.) and its wild relatives belong to Solanum section Petota. This section's phylogeny and species delimitation are complicated due to various ploidy levels, high heterozygosity, and frequent interspecific hybridization. Compared to the nuclear genome, the plastid genome is more conserved, has a haploid nature, and has a lower nucleotide substitution rate, providing informative alternative insights into the phylogenetic study of section Petota. Here, we analyzed 343 potato plastid genomes from 53 wild and four cultivated species. The diversity of sequences and genomes was comprehensively analyzed. A total of 24 species were placed in a phylogenetic tree based on genomic data for the first time. Overall, our results not only confirmed most existing clades and species boundaries inferred by nuclear evidence but also provided some distinctive species clade belonging and the maternally inherited evidence supporting the hybrid origin of some species. Furthermore, the divergence times between the major potato clades were estimated. In addition, the species discriminatory power of universal barcodes, nuclear ribosomal DNA, and whole and partial plastid genomes and their combinations were thoroughly evaluated; the plastid genome performed best but had limited discriminatory power for all survey species (40%). Overall, our study provided not only new insights into phylogeny and DNA barcoding of potato but also provided valuable genetic data resources for further systematical research of Petota.
    • Potato and its wild relatives belong to Solanum section Petota. In this study, the phylogeny of Petota was confirmed on clade level based on 343 plastid genomes. The species identification rates of universal barcodes, nuclear ribosomal DNA, whole plastid genomes, and hypervariable regions were thoroughly evaluated.
    • Chi‐Chuan Chen, Jaakko Hyvönen, and Harald Schneider
      2023, 61 (4): 613–626
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      The fern family Polypodiaceae, with over 1600 species, is not only one of the most species-rich families of ferns, but also a major contributor to the vascular epiphytic diversity throughout the tropics. Although the vast majority of species belonging to this family prefer to grow as epiphytes, several species colonize successfully rheophytic, lithophytic, and even terrestrial habitats. Here, we explore the hypothesis that non-epiphytic habitat preferences, including terrestrial growth, evolved secondarily with epiphytes being the plesiomorphic habitat preference. The results of phylogenetic analyses, based on dense taxon sampling and four chloroplast DNA regions, were integrated with divergence time estimates and ancestral character state reconstructions to test these predictions. Both fossils and secondary calibration data were incorporated to obtain divergence time estimations. The results support the prediction of multiple transitions from epiphytic/lithophytic to terrestrial/rheophytic habitats occurring mainly in the Microsoreae lineage. The change in niche preferences coincides with niche colonization opportunities created by climatic fluctuations and geographical changes during the Oligocene and Miocene periods.
    • This image shows the phylogenetic relationship of Oleandraceae, Davalliaceae, and Polypodiaceae with six subfamilies, the Oleandraceae selected as an outgroup. With the integration of the growth form dataset, distribution of terrestrial species within Polypodiaceae is displayed. A clade marked with blue color indicates Leptochilus-MG5, in which the terrestrial species is dominant. This change in niche preferences is associated with competition in upper forest canopy habitats, influenced by the geographical change or climate fluctuations during the Oligocene–Miocene periods.
    • Ying‐Xue Jiao, Xiao‐Fan He, Rui Song, Xue‐Meng Wang, Han Zhang, Reziya Aili, Yue‐Hui Chao, Yu‐Hua Shen, Long‐Xi Yu, Tie‐Jun Zhang, and Shan‐Gang Jia
      2023, 61 (4): 627–642
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      Medicago is a genus of legumes (Fabaceae) that resemble common clovers with pinnately trifoliate leaves and spirally coiled seed pods, and Medicago sativa is a famous forage crop throughout the world. In this study, we systematically assembled the complete plastid genomes of 18 Medicago species, representing 35 Medicago accessions, whose genome size ranged from ~119 to 125?kb, and identified one novel inverted repeat (IR) in two accessions of Medicago soleirolii (PI537242 and PI537243), albeit of no IRs in the most accessions. We built a phylogenetic tree based on common protein-coding sequences of 55 Medicago accessions in 38 species, which were placed into five clades with a divergence since 9.37 million years ago. Global alignment revealed independent genome evolution events, including eight inversions in nine species and four intron losses (ILs) in 10 species, among which four inversions and two ILs have not been reported previously. Within 109–111 unique genes, ndhA, rpl2, and ycf3 were under positive selection in 54 Medicago accessions. Finally, by aligning chloroplast genes against the nuclear genome assembly of M. sativa cultivar “Zhongmu No.1”, we found that a large number of chloroplast gene fragments were horizontally transferred to nuclear chromosomes in alfalfa, especially on the chr3:47518422–48722257 coordinates of chromosome 3. Our comprehensive exploration of Medicago chloroplast genomes provided insights for the understanding of Medicago diversity and their genomic evolution events.
    • 1. Eight inversions were identified in 10 species, among which four inversions have not been reported previously.
      2. A novel inverted repeat of ~0.3 kb was discovered in Medicago soleirolii in the present study.
      3. Large numbers of chloroplast gene fragments were horizontally transferred to nuclear chromosomes in alfalfa.
    • Chapa Manawaduge, Gillian Brown, Catherine L. Simmons, Matthew J. Phillips, and Susan Fuller
      2023, 61 (4): 643–656
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      Conservation decision-making and prioritization of management actions for taxa at high risk of extinction require a clear understanding of systematic relationships and species boundaries. Taxonomic uncertainty surrounds two threatened species of native olive (genus Notelaea) endemic to Australia. Notelaea ipsviciensis is known from only one small population and is listed as critically endangered under Australian and Queensland legislation, while Notelaea lloydii is listed as vulnerable due to its restricted distribution in south-east Queensland, Australia. Notelaea ipsviciensis occurs in sympatry with N. lloydii, at the only site where it is found, and exhibits intermediate morphology between N. lloydii and Notelaea ovata raising speculation that it is a natural hybrid of the two. To address this uncertainty, we have reconstructed the molecular phylogeny of the genus Notelaea using both single-gene sequencing and single nucleotide polymorphism (SNP) data. While the chloroplast DNA markers were not found to be informative, the genome-wide SNPs used in this study have unequivocally resolved the long-standing question of the hybrid origin of N. ipsviciensis. Results of the phylogenetics and hybrid analyses of SNP data show that N. ipsviciensis is a natural hybrid of N. lloydii and N. ovata, and the taxon's circumscription needs to be re-evaluated. Our results also revealed unclear species boundaries among numerous other Notelaea species, including the threatened N. lloydii, indicating that further investigation and taxonomic revision may be necessary.
    • Threatened species conservation relies on having knowledge of the systematic relationships between species and clearly resolved species boundaries. This study investigated the systematics of the two least studied threatened native olive species (Notelaea ipsviciensis and Notelaea lloydii) in Australia. Results of the phylogenetics and hybrid analyses of genomic data confirm that N. ipsviciensis is a natural hybrid of N. lloydii and Notelaea ovata, and the taxon's circumscription needs to be re-evaluated. Furthermore, the molecular phylogeny consequently raises doubts about the current circumscription of some other species of Notelaea, including the threatened species N. lloydii.
    • Marie Claire Veranso‐Libalah, Hannah Mertes, Robert Douglas Stone, Luo Chen, Tariq Stévart, Frank Almeda, Xander M. van der Burgt, and Gudrun Kadereit
      2023, 61 (4): 657–681
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      The tribe Sonerileae in tropical Africa and Madagascar is a morphologically diverse lineage that consists of 239 species in 10 genera. In this study, we present the first in-depth phylogenetic analysis of African Sonerileae to test monophyly of the currently recognized genera. Phylogenetic analyses were performed using sequence data from two nuclear (nrITS and nrETS) and three plastid loci (accD-psaI, ndhF and psbK-psbL). Sampling consisted of 140 accessions including 64 African, 27 Malagasy, 46 Asian, and three neotropical Sonerileae together with a broad outgroup sampling (105 spp.). Phylogenetic relationships were inferred using maximum likelihood and Bayesian inference approaches, and a careful reassessment of morphological characters was carried out. Our results neither support the monophyly of the Old World nor African Sonerileae. The monospecific African genus Benna is partially supported as sister to Phainantha, one of the basal neotropical lineages, while African and Malagasy Medinilla are nested among the SE Asian genera. Gravesia (116 spp.), the most species-rich and morphologically diverse genus in Madagascar, is recovered as monophyletic. The African genera of Sonerileae Calvoa, Dicellandra, and Preussiella form well-supported clades. In contrast, Amphiblemma (including Amphiblemma molle) and Cincinnobotrys s.l. (including Cincinnobotrys felicis) are not monophyletic. To accommodate the caulescent C. felicis we propose reinstatement of the monospecific genus Bourdaria. For the distinctive A. molle a new genus Mendelia is described. Calvoa hirsuta is designated here as the type of genus Calvoa, lectotypes are designated for Medinilla engleri and Veprecella lutea, and a neotype is designated for Preussiella kamerunensis.
    • The first in-depth phylogenetic analysis of African Sonerileae to test monophyly of the currently recognized genera is here presented. Phylogenetic analyses neither support monophyly of the Old World nor African Sonerileae. The genera Calvoa, Dicellandra, Gravesia, and Preussiella form well-supported clades while Amphiblemma and Cincinnobotrys are not monophyletic. The monospecific genus Bourdaria is reinstated and a new genus Mendelia is described to accommodate Cincinnobotrys felicis and Amphiblemma molle, respectively.
    • Tian‐Tian Xue, Xu‐Dong Yang, Qin Liu, Fei Qin, Wen‐Di Zhang, Steven B. Janssens, and Sheng‐Xiang Yu
      2023, 61 (4): 682–697
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      A significant fraction of higher plants in China are threatened due to dramatic landscape transformation and increasing climate change. However, the conservation effectiveness of threatened higher plants (THPs) and their response to climate change are still underexplored to date. Based on the latest list of THPs in China, we obtained 102?593 occurrence records with latitude and longitude for 3858 THPs. By integrating the distribution patterns of three biodiversity indexes (i.e., species richness, species complementarity, and weighted endemism) and 10 plant categories, we identified hotspots for THPs and calculated the conservation effectiveness of nature reserves. We then selected 1959 THPs to project the shift of species richness and range sizes under climate change (representative concentration pathway [RCP] 2.6 and RCP 8.5). In total, 16 hotspot areas covering 7.38% of Chinese land area and containing 91.73% of THPs were identified. Current nature reserves protected 35.05% of hotspots, 73.07% of all THPs, and 56.64% of narrow-ranged species. By the 2070s, the species richness of THPs were predicted to decrease in Southeast and Central China, and 42.42% (RCP 2.6) and 51.40% (RCP 8.5) of the 1959 THPs would confront habitat contraction. Future conservation efforts should focus on the conservation gaps and carry out targeted conservation for THPs with narrow distribution range. In order to cope with climate change, the hotspots with relatively low species loss can serve as important areas to contain current species diversity and the areas with high species gain offer opportunities for ex-situ conservation of THPs.
    • Different threatened higher plants (THPs) are not congruent with each other in distribution. Mountainous areas in South China are important for conserving the diversity of THPs. Hotspots in Northwest China are characterized by unique species composition. Nature reserves are not effectively preserving THPs with narrow distribution range. Species richness of THPs will significantly shift in Southeast China under climate change.
    • Gui‐Lin Wu, De‐Xiang Chen, Zhang Zhou, Qing Ye, Andrés Baselga, Hui Liu, Yin Wen, and Shou‐Qian Nong
      2023, 61 (4): 698–708
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      Diversification rates are critically important for understanding patterns of species richness among clades. However, the effects of climatic niche width on plant diversification rates remain to be elucidated. Based on the phylogenetic, climatic, and distributional information of angiosperms in China, a total of 26?906 species from 182 families were included in this study. We aimed to test relationships between diversification rate and climatic niche width and climatic niche width related variables (including climatic niche divergence, climatic niche position, geographic extent, and climatic niche evolutionary rate) using phylogenetic methods. We found that climatic niche divergence had the largest unique contribution to the diversification rate, while the unique effects of climatic niche width, climatic niche position, geographic extent, and climatic niche evolutionary rate on the diversification rate were negligible. We also observed that the relationship between diversification rate and climatic niche divergence was significantly stronger than the null assumption (artefactual relationship between diversification and clade-level climatic niche width by sampling more species). Our study supports the hypothesis that wider family climatic niche widths explain faster diversification rates through a higher climatic niche divergence rather than through higher geographic extent, higher climatic niche evolutionary rate, or separated climatic niche position. Hence, the results provide a potential explanation for large-scale diversity patterns within families of plants.
    • We sought to identify whether the niche divergence or niche conservatism hypothesis can explain why some clades have higher species richness than others for angiosperms of China. We tested the hypotheses based on the phylogenetic, climatic, and distributional data of the angiosperms in China, a total of 26 906 species from 182 families. We found that wider family niche widths drive faster diversification rates through a higher niche divergence, rather than geographic extent or niche position. Therefore, our study strongly supported niche divergence hypothesis. The results provide a potential explanation for large-scale diversity patterns within families of plants.
    • Zhang‐Jian Shan, Qian Zhang, Dan‐Xiao Peng, Jian‐Fei Ye, Lan Cao, Zhi‐Duan Chen, and Hai‐Ning Qin
      2023, 61 (4): 709–718
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      Over-harvesting, habitat loss and fragmentation, and biological invasions have led to a sharp decline in wild medicinal plants population in China, where they are an essential component of traditional medicine and used widely. The current national list of protected medicinal materials, the State Key-protected Wild Medicinal Species List (SKPWMSL), which has not been revised for 30 years, is in urgent need of an update. This study proposes a new scoring system with seven indicators that set the conservation priorities of threatened medicinal plants. The advantages of our approach include: (i) quantitative methods with high repeatability and comparability; and (ii) consideration of the evolutionary history of medicinal species. After assessing 911 threatened medicinal angiosperms in China, we identified 112 species as key medicinal plants for conservation priority (KMPCP). We suggest promoting the SKPWMSL with KMPCP as a supplement and update. Meanwhile, our scoring system will improve the future setting of conservation priority and can be extended to other countries or regions.
    • We propose a new robust and highly repeatable scoring system that includes seven indicators (i.e., threatened and conservation status, evolutionary distinctiveness, harvesting parts, medicinal usage, ecological value, and number of uses) to set the conservation priorities of threatened medicinal plants. To our knowledge, this is the first time that the evolutionary distinctiveness is considered for setting conservation priorities of medicinal plants.
    • Ada Chornelia and Alice C. Hughes
      2023, 61 (4): 719–732
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      Bats provide an excellent casestudy for studying evolution due to their remarkable flight and echolocation capabilities. In this study, we sought to understand the phenotypic evolution of key traits in Rhinolophidae (horseshoe bats) using phylogenetic comparative methods. We aimed to test the phylogenetic signals of traits, and evaluated the best-fit evolutionary models given the data for each trait considering different traits may evolve under different models (i.e., Brownian Motion [BM], Ornstein-Uhlenbeck [OU], and Early Burst [EB]) and reconstruct ancestral character states. We examined how phenotypic characters are associated with echolocation calls and minimum detectable prey size. We measured 34 traits of 10 Asian rhinolophids species (187 individuals). We found that the majority of traits showed a high phylogenetic signal based on Blomberg′s K and Pagel′s λ, but each trait may evolve under different evolutionary models. Sella traits were shown to evolve under stabilizing selection based on OU models, indicating sella traits have the tendency to move forward along the branches toward some medial value in equilibrium. Our findings highlight the importance of sella characters in association with echolocation call emissions in Rhinolophidae, as calls are important for spatial cognition and also influence dietary preferences. Minimum detectable prey size in Rhinolophidae was associated with call frequency, bandwidth, call duration, wingspan, and wing surface area. Ultimately, understanding trait evolution requires sensitivity due to the differential selective pressures which may apply to different characteristics.
    • Noseleaf characters showed relatively high phylogenetic signals and are highly correlated with call frequency at maximum energy. We showed the deviation of the allometry rule in Rhinolophidae, and suggested that noseleaf characters are better predictors for acoustic calls compared to body size. Furthermore, we also found that each character may evolve with different evolutionary models, interestingly, sella characters (the protruding tissue above the nostril) were shown to evolve under aOrnstein-Uhlenbeck (OU) trait evolution model. This suggested that disparities within these characteristics between species evolved during equilibrium, or stabilizing selection, and other intermediate forms or variance within species may be eliminated by natural selection or other pressures. Our result provides pilot information which valuable for future studies on the evolution of key functional traits
Editors-in-Chief
Song Ge
Jun Wen
Impact Factor
3.7
JCR 2022 IF ranking: 60/238 (Plant Sciences, top 25%, Q2 quartile)
Journal Abbreviation: J Syst Evol
ISSN: 1674-4918 (Print)
1759-6831 (Online)
CN: 11-5779/Q
Frequency: Bi-monthly

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