Table of Contents

25 July 2016, Volume 54 Issue 4

Cover illustration: Current framework of rosid phylogeny is displayed as a big tree with sturdy trunk and wide-spreading branches. A series of snapshots of flowers or fruits are displayed on the top of each clade, reflecting their morphological characters and economic importance. Different colors represent 17 orders of Rosidae. The super-ordinal relationships within Rosidae are outlined, sketching out a basic structure of the tree trunk (please see Sun et al., pp. 363–391 in this issue). This cover image shapes like a fan in Chinese painting, attempting to reflect that this special issue is a major regional effort toward the whole tree of life project. [Detail] ...

  

Editorial
• The Tree of Life: China project

  Zhi-Duan Chen, An-Ming Lu, Shou-Zhou Zhang, Qing-Feng Wang, Zhong-Jian Liu, De-Zhu Li, Hong Ma, Jian-Hua Li, Douglas E. Soltis, Pamela S. Soltis, Jun Wen, China Phylogeny Consortium

  J Syst Evol. 2016, 54(4): 273-276.
  https://doi.org/10.1111/jse.12215

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Research Articles
• Tree of life for the genera of Chinese vascular plants

  Zhi-Duan Chen, Tuo Yang, Li Lin, Li-Min Lu, Hong-Lei Li, Miao Sun, Bing Liu, Min Chen, Yan-Ting Niu, Jian-Fei Ye, Zhi-Yong Cao, Hong-Mei Liu, Xiao-Ming Wang, Wei Wang, Jing-Bo Zhang, Zhen Meng, Wei Cao, Jian-Hui Li, Sheng-Dan Wu, Hui-Ling Zhao, Zhong-Jian Liu, Zhi-Yuan Du, Qing-Feng Wang, Jing Guo, Xin-Xin Tan, Jun-Xia Su, Lin-Jing Zhang, Lei-Lei Yang, Yi-Ying Liao, Ming-He Li, Guo-Qiang Zhang, Shih-Wen Chung, Jian Zhang, Kun-Li Xiang, Rui-Qi Li, Douglas E. Soltis, Pamela S. Soltis, Shi-Liang Zhou, Jin-Hua Ran, Xiao-Quan Wang, Xiao-Hua Jin, You-Sheng Chen, Tian-Gang Gao, Jian-Hua Li, Shou-Zhou Zhang, An-Ming Lu, China Phylogeny Consortium

  J Syst Evol. 2016, 54(4): 277-306.
  https://doi.org/10.1111/jse.12219

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  We reconstructed a phylogenetic tree of Chinese vascular plants (Tracheophyta) using sequences of the chloroplast genes atpB, matK, ndhF, and rbcL and mitochondrial matR. We produced a matrix comprising 6098 species and including 13?695 DNA sequences, of which 1803 were newly generated. Our taxonomic sampling spanned 3114 genera representing 323 families of Chinese vascular plants, covering more than 93% of all genera known from China. The comprehensive large phylogeny supports most relationships among and within families recognized by recent molecular phylogenetic studies for lycophytes, ferns (monilophytes), gymnosperms, and angiosperms. For angiosperms, most families in Angiosperm Phylogeny Group IV are supported as monophyletic, except for a paraphyletic Dipterocarpaceae and Santalaceae. The infrafamilial relationships of several large families and monophyly of some large genera are well supported by our dense taxonomic sampling. Our results showed that two species of Eberhardtia are sister to a clade formed by all other taxa of Sapotaceae, except Sarcosperma. We have made our phylogeny of Chinese vascular plants publically available for the creation of subtrees via SoTree (http://www.darwintree.cn/flora/index.shtml), an automated phylogeny assembly tool for ecologists.
• Embracing the pteridophyte classification of Ren-Chang Ching using a generic phylogeny of Chinese ferns and lycophytes

  Hong-Mei Liu

  J Syst Evol. 2016, 54(4): 307-335.
  https://doi.org/10.1111/jse.12184

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  The phylogenetic relationships of pteridophytes occurring in China were reconstructed using DNA sequences of the three plastid genes, atpA, atpB, and rbcL. The sampling comprised all genera of Chinese pteridophytes—including ferns and lycophytes—with the exception of four small genera. The effort to sample all recorded families and genera in a phylogenetic framework enabled the phylogenetic relationships of all Chinese pteridophytes to be addressed for the first time in a single phylogenetic hypothesis. The results provided strong evidence to support the continuing impact of Ren-Chang Ching's integrative classification of pteridophytes. Ten out of 11 orders accepted by Ching were consistent with the phylogeny, whereas four new orders were introduced to avoid paraphyletic taxa in the leptosporangiate ferns. Of the 63 families considered by Ching, 36 families were supported by molecular data, 22 of those had the same or nearly the same circumscription, and the remaining 14 families were supported but substantially revised. Twenty-eight small families were now accepted as synonyms. A consistent pattern was observed at the generic level. Among the 223 genera considered by Ching, 133 genera were recognized by the phylogeny, although some of them were substantially changed in the context of circumscription, and 90 were now accepted as synonyms. Three endemic genera were incorporated here for the first time in DNA-based phylogenetic analyses, namely Blechnidium, Saxiglossum, and Sinephropteris, which were shown to be nested in Blechnum, Pyrrosia, and Asplenium respectively. This paper tentatively accepts 40 families and 151 genera of ferns and lycophytes occurring in China; the importance of phylodiversity of Chinese pteridophytes is also briefly discussed.
• Accelerated evolution of early angiosperms: Evidence from ranunculalean phylogeny by integrating living and fossil data

  Wei Wang, David L. Dilcher, Ge Sun, Hong-Shan Wang, Zhi-Duan Chen

  J Syst Evol. 2016, 54(4): 336-341.
  https://doi.org/10.1111/jse.12090

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  The new discovery of angiosperm remains in the Jehol Biota of northeastern China contributes to our understanding of the origin and early evolution of flowering plants. The earliest eudicot genus with reproductive organs, Leefructus, was recently documented from the Lower Cretaceous Yixian Formation at 125.8–123.0 Ma, and was reconsidered to be close to the extant family Ranunculaceae based on gross morphology. However, this hypothesis has not been tested using a cladistic approach. To determine the possible allies of Leefructus within extant eudicots, we constructed a 66 morphological data matrix. Molecular and morphological analyses of extant Ranunculales combined with the fossil suggest that it has an affinity with the Ranunculaceae. The earliest fossil record of the eudicots is 127–125 Ma based on tricolpate pollen grains. Thus, we suggest a hypothesis that the basal eudicots might have experienced an accelerated evolution and diversification during the latest Barremian and earliest Aptian, leading to the stem groups of at least six extant families or lineages, 10–15 Myr earlier than currently documented. Angiosperms have undergone multiple uneven pulses of radiation since their origin. Many key character innovations occurred in different stages that could have triggered those radiations in concert with various biotic and abiotic factors.
• Phylogenetic tree of vascular plants reveals the origins of aquatic angiosperms

  Zhi-Yuan Du, Qing-Feng Wang, China Phylogeny Consortium

  J Syst Evol. 2016, 54(4): 342-348.
  https://doi.org/10.1111/jse.12182

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  Although aquatic plants are discussed as a unified biological group, they are phylogenetically well dispersed across the angiosperms. In this study, we annotated the aquatic taxa on the tree of vascular plants, and extracted the topology of these aquatic lineages to construct the tree of aquatic angiosperms. We also reconstructed the ancestral areas of aquatic families. We found that aquatic angiosperms could be divided into two different categories: the four aquatic orders and the aquatic taxa in terrestrial orders. Aquatic lineages evolved early in the radiation of angiosperms, both in the orders Nymphaeales and Ceratophyllales and among basal monocots (Acorales and Alismatales). These aquatic orders do not have any extant terrestrial relatives. They originated from aquatic habitats during the Early Cretaceous. Asia would have been one of the centers for early diversification of aquatic angiosperms. The aquatic families within terrestrial orders may originate from other areas besides Asia, such as America or Australia. The lineages leading to extant angiosperms diversified early in underexploited freshwater habitats. The four extant aquatic orders were relicts of an early radiation of angiosperm in aquatic environments. Their extinct ancestors might be aquatic early angiosperms.
• A molecular phylogeny of Chinese orchids

  Ming-He Li, Guo-Qiang Zhang, Si-Ren Lan, Zhong-Jian Liu, China Phylogeny Consortium

  J Syst Evol. 2016, 54(4): 349-362.
  https://doi.org/10.1111/jse.12187

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  We estimated the molecular phylogenetic relationships of the Chinese members of the family Orchidaceae. Within the Tree of Life for the Genera of Chinese Vascular Plants using atpB, rbcL, matK, ndhF, and matR, the currently delimited subfamilies, tribes, and subtribes are highly supported as monophyletic except for the perplexing Epidendroideae. Five genes (rbcL, matK, psaB, ycf1, and Xdh), which are more universally used in Orchidaceae, were further analyzed to reconstruct the phylogeny of Epidendroideae. The reconstructed trees were in strong agreement and showed significant support for the tribal and subtribal clades. Based on the highly supported circumscription and arrangement of the suprageneric levels in the Tree of Life and reconstructed trees, we have proposed a new phylogenetic classification of Chinese Orchidaceae that includes five subfamilies, 17 tribes, and 21 subtribes.
• Phylogeny of the Rosidae: A dense taxon sampling analysis

  Miao Sun, Rehan Naeem, Jun-Xia Su, Zhi-Yong Cao, J. Gordon Burleigh, Pamela S. Soltis, Douglas E. Soltis, Zhi-Duan Chen

  J Syst Evol. 2016, 54(4): 363-391.
  https://doi.org/10.1111/jse.12211

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  Rosidae, a clade of approximately 90 000 species of angiosperms, exhibits remarkable morphological diversity and extraordinary heterogeneity in habitats and life forms. Resolving phylogenetic relationships within Rosidae has been difficult, in large part due to nested radiations and the enormous size of the clade. Current estimates of phylogeny contain areas of poor resolution and/or support, and there have been few attempts to synthesize the available data into a comprehensive view of Rosidae phylogeny. We aim to improve understanding of the phylogeny of Rosidae with a dense sampling scheme using both newly generated sequences and data from GenBank of the chloroplast rbcL, atpB, and matK genes and the mitochondrial matR gene. We combined sequences from 9300 species, representing 2775 genera, 138 families, and 17 orders into a supermatrix. Although 59.26% of the cells in the supermatrix have no data, our results generally agree with previous estimates of Rosidae phylogeny and provide greater resolution and support in several areas of the topology. Several noteworthy phylogenetic relationships are recovered, including some novel relationships. Two families (Euphorbiaceae and Salvadoraceae) and 467 genera are recovered as non-monophyletic in our sampling, suggesting the need for future systematic studies of these groups. Our study shows the value of a botanically informed bioinformatics approach and dense taxonomic sampling for resolving rosid relationships. The resulting tree provides a starting point for large-scale analyses of the evolutionary patterns within Rosidae.
• Global versus Chinese perspectives on the phylogeny of the N-fixing clade

  Hong-Lei Li, Wei Wang, Rui-Qi Li, Jing-Bo Zhang, Miao Sun, Rehan Naeem, Jun-Xia Su, Xiao-Guo Xiang, Peter E. Mortimer, De-Zhu Li, Kevin D. Hyde, Jian-Chu Xu, Douglas E. Soltis, Pamela S. Soltis, Jianhua Li, Shou-Zhou Zhang, Hong Wu, Zhi-Duan Chen, An-Ming Lu

  J Syst Evol. 2016, 54(4): 392-399.
  https://doi.org/10.1111/jse.12201

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  There has been increasing interest in integrating a regional tree of life with community assembly rules in the ecological research. This raises questions regarding the impacts of taxon sampling strategies at the regional versus global scales on the topology. To address this concern, we constructed two trees for the nitrogen-fixing clade: (i) a genus-level global tree including 1023 genera; and (ii) a regional tree comprising 303 genera, with taxon sampling limited to China. We used the supermatrix approach and performed maximum likelihood analyses on combined matK, rbcL, and trnL-F plastid sequences. We found that the topology of the global and the regional tree of the N-fixing clade were generally congruent. However, whereas relationships among the four orders obtained with the global tree agreed with the accepted topology obtained in focused analyses with more genes, the regional topology obtained different relationships, albeit weakly supported. At a finer scale, the phylogenetic position of the family Myricaceae was found to be sensitive to sampling density. We expect that internal support throughout the phylogeny could be improved with denser taxon sampling. The taxon sampling approach (global vs. regional) did not have a major impact on fine-level branching patterns of the N-fixing clade. Thus, a well-resolved phylogeny with relatively dense taxon sampling strategy at the regional scale appears, in this case, to be a good representation of the overall phylogenetic pattern and could be used in ecological research. Otherwise, the regional tree should be adjusted according to the correspondingly reliable global tree.
• A supermatrix approach provides a comprehensive genus-level phylogeny for Gentianales

  Lei-Lei Yang, Hong-Lei Li, Lei Wei, Tuo Yang, Dai-Yong Kuang, Ming-Hong Li, Yi-Ying Liao, Zhi-Duan Chen, Hong Wu, Shou-Zhou Zhang

  J Syst Evol. 2016, 54(4): 400-415.
  https://doi.org/10.1111/jse.12192

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  Gentianales consist of Apocynaceae, Gelsemiaceae, Gentianaceae, Loganiaceae, and Rubiaceae, of which the majority are woody plants in tropical and subtropical areas. Despite extensive efforts in reconstructing the phylogeny of Gentianales based on molecular data, some interfamily and intrafamily relationships remain uncertain. We reconstructed the genus-level phylogeny of Gentianales based on the supermatrix of eight plastid markers (rbcL, matK, atpB, ndhF, rpl16, rps16, thetrnL-trnF region, and atpB-rbcL spacer) and one mitochondrial gene (matR) using maximum likelihood. The major clades and their relationships retrieved in the present study concur with those of previous studies. All of the five families of Gentianales are monophyletic with strong support. We resolved Rubiaceae as sister to the remaining families in Gentianales and showed support for the sister relationship between Loganiaceae and Apocynaceae. Our results provide new insights into relationships among intrafamilial clades. For example, within Rubiaceae we found that Craterispermeae were sister to Morindeae + (Palicoureeae + Psychotrieae) and that Theligoneae were sister to Putorieae. Within Gentianaceae, our phylogeny revealed that Gentianeae were sister to Helieae and Potalieae, and subtribe Lisianthiinae were sister to Potaliinae and Faroinae. Within Loganiaceae, we found Neuburgia as sister to Spigelieae. Within Apocynaceae, our results supported Amsonieae as sister to Melodineae, and Hunterieae as sister to a clade comprising Plumerieae + (Carisseae + APSA). We also confirmed the monophyly of Perplocoideae and the relationships among Baisseeae + (Secamonoideae + Asclepiadoideae).
• A comprehensive generic-level phylogeny of the sunflower family: Implications for the systematics of Chinese Asteraceae

  Zhi-Xi Fu, Bo-Han Jiao, Bao Nie, Guo-Jin Zhang, Tian-Gang Gao, China Phylogeny Consortium

  J Syst Evol. 2016, 54(4): 416-437.
  https://doi.org/10.1111/jse.12216

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  The sunflower family (Asteraceae) is the largest and the most diverse flowering plant family, comprising 24 000–30 000 species and 1600–1700 genera. In China, Asteraceae are also the largest family, with approximately 2336 indigenous species in 248 genera. In the past two decades, molecular phylogenetic analyses has contributed greatly to our understanding of the systematics of Asteraceae. Nevertheless, the large-scale analyses and knowledge about the relationships of Chinese Asteraceae at the generic level as a whole are far from complete due to difficulties in sampling. In this study, we presented a three-marker (rbcL, ndhF, and matK) phylogeny of Asteraceae, including 506 genera (i.e., approximately one-third of Asteraceae genera). The study sampled 200 Chinese genera (i.e., approximately 80% of Chinese Asteraceae genera). The backbones of the new phylogeny were largely congruent with earlier studies, with 13 subfamilies and 45 tribes recognized. Chinese Asteraceae were distributed in 7 subfamilies (Mutisioideae, Wunderlichioideae, Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, and Asteroideae) and 22 tribes (Mutiseae, Hyalideae, Cardueae, Pertyeae, Gymnarrheneae, Vernonieae, Cichorieae, Doroniceae, Senecioneae, Astereae, Anthemideae, Gnaphalieae, Calenduleae, Inuleae, Athroismeae, Helenieae, Coreopsideae, Neurolaeneae, Tageteae, Millieae, Eupatorieae, and Heliantheae). Chinese Asteraceae lacked 6 basal subfamilies and 23 tribes. Several previously ambiguous relationships were clarified. Our analyses also resolved some unplaced genera within Chinese Asteraceae. Finally, our phylogenetic tree was used to revise the classification for all genera of Chinese Asteraceae. In total, 255 genera, 22 tribes, and 7 subfamilies in China are recognized.
• Using nuclear genes to reconstruct angiosperm phylogeny at the species level: A casestudy with Brassicaceae species

  Liming Cai, Hong Ma

  J Syst Evol. 2016, 54(4): 438-452.
  https://doi.org/10.1111/jse.12204

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  Angiosperm phylogeny has been investigated extensively using organellar sequences; recent efforts using nuclear genes have also been successful in reconstructing angiosperm phylogenies at family or deeper levels. However, it is not clear whether nuclear genes are also effective in understanding relationships between species in a genus. Here we present a case study of phylogeny at generic and specific levels with nuclear genes, using Brassicaceae taxa as examples. Brassicaceae includes various crops and the model plant Arabidopsis thaliana. A recent study showed that nuclear genes can provide well-resolved relationships between tribes and larger lineages in Brassicaceae, but few species were included in any given genus. We present a phylogeny with multiple species in each of five genera within Brassicaceae for a total of 65 taxa, using three protein-coding nuclear genes, MLH1, SMC2, and MCM5, with up to approximately 10 200 base pairs (in both exons and introns). Maximum likelihood and Bayesian analyses of the separate gene regions and combined data reveal high resolution at various phylogenetic depths. The relationships between genera here were largely congruent with previous results, with further resolution at the species level. Also, we report for the first time the affinity of Cardamine rockii with tribe Camelineae instead of other Cardamine members. In addition, we report sequence divergence at three levels: across angiosperms, among Brassicaceae species, and between Arabidopsis ecotypes. Our results provide a robust species-level phylogeny for a number of Brassicaceae members and support an optimistic perspective on the phylogenetic utility of conserved nuclear data for relatively recent clades.
• Phylogeny and diversification of Chinese Araliaceae based on nuclear and plastid DNA sequence data

  Rong Li, Jun Wen

  J Syst Evol. 2016, 54(4): 453-467.
  https://doi.org/10.1111/jse.12196

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  Chinese Araliaceae consist of 20 genera and ca. 175 species. To assess the evolutionary relationships of Araliaceae and their biogeographic diversification in China, the phylogeny of Chinese Araliaceae was constructed by sampling 96 accessions representing 20 genera and 50 species of Chinese Araliaceae and 45 closely related taxa using sequences of the nuclear ribosomal internal transcribed spacer (ITS) region and six plastid regions (the ndhF gene, the trnL-trnF region, the rps16intron, the atpB-rbcL intergenic spacer, the rpl16 intron, and the psbA-trnH intergenic spacer). Phylogenetic analyses of the combined plastid and ITS data supported the results of the previously studies that the Chinese members of Araliaceae were scattered within the Asian Palmate group and the Aralia-Panax group withOsmoxylon at the base of core Araliaceae. The generic status of Pentapanax and Tupidanthus is not supported. Our analysis clearly places them in Aralia and AsianSchefflera, respectively. In a broader phylogenetic framework of Araliaceae, based on the fossil-calibrated Bayesian dating, Chinese Araliaceae was inferred to have originated in Asia and underwent a rapid radiation in its evolutionary history. Its diversification is hypothesized to have been driven largely by the orogenies in Asia during the Cenozoic. In China, the distribution pattern of the phylogenetic diversity of Araliaceae corresponds with its taxonomic diversity across the entire region.