Table of Contents

01 January 2022, Volume 60 Issue 1

Cover illustration: Left to right, top to bottom: Magnolia lucida, M. viridula, M. champaca, M. championii, M. odora, M. salicifolia, M. grandis, M. liliifera, M. foveolata, M. grandifl ora, M. biondii, M. martini. See Dong et al., pp. 1–15 in this issue. Photographs by Ya-Ling Wang. Designed by Shan-Shan Dong.

  

Research Articles
• 

  Plastid and nuclear phylogenomic incongruences and biogeographic implications of Magnolia s.l. (Magnoliaceae)

  Shan-Shan Dong, Ya-Ling Wang, Nian-He Xia, Yang Liu, Min Liu, Lian Lian, Na Li, Ling-Fei Li, Xiao-An Lang, Yi-Qing Gong, Lu Chen, Ernest Wu, and Shou-Zhou Zhang

  J Syst Evol. 2022, 60(1): 1-15.
  https://doi.org/10.1111/jse.12727

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  Magnoliaceae, an assemblage of early diverged angiosperms, comprises two subfamilies, speciose Magnolioideae with approximately 300 species in varying numbers of genera and monogeneric Liriodendroideae with two species in Liriodendron L. This family occupies a pivotal phylogenetic position with important insights into the diversification of early angiosperms, and shows intercontinentally disjunct distribution patterns between eastern Asia and the Americas. Widespread morphological homogeneity and slow substitution rates in Magnolia L. s.l. resulted in poorly supported phylogenetic relationships based on morphology or molecular evidence, which hampers our understanding of the genus’ temporal and spacial evolution. Here, based on the newly generated genome skimming data for 48 Magnolia s.l. species, we produced robust Magnolia phylogenies using genome-wide markers from both plastid genomes and single nucleotide polymorphism data. Contrasting the plastid and nuclear phylogenies revealed extensive cytonuclear conflicts in both shallow and deep relationships. ABBA-BABA and PhyloNet analyses suggested hybridization occurred within sect. Yulania, and sect. Magnolia, which is in concordance with the ploidy level of the species in these two sections. Divergence time estimates and biogeographic reconstruction indicated that the timing of the three tropical Magnolia disjunctions coincided with the mid-Eocene cooling climate and/or late Eocene climate deterioration, and two temperate disjunctions occurred much later, possibly during the warm periods of the Miocene, hence supporting the boreotropical flora concept of Magnolia s.l.

  Magnoliaceae comprises two subfamilies, speciose Magnolioideae with about 300 species in varying numbers of genera and monogeneric Liriodendroideae with two species in Liriodendron. Widespread morphological homogeneity and slow substitution rates in Magnolia s.l. resulted in poorly supported phylogenetic relationships based on morphology or molecular evidences. Here, based on the newly generated genome skimming data for 48 Magnolia s.l. species, we produced robust Magnolia phylogenies using genome-wide markers from both plastid genomes and SNP data. Contrasting the plastid and nuclear phylogenies revealed extensive cytonuclear conflicts in both shallow and deep relationships. Split tree network and ABBA-BABA analyses suggested hybridization occurred within sect. Yulania, and sect. Magnolia, which is in concordance with the ploidy level of the species in these two sections, and further substantiated by PhyloNet and JML analyses. Divergence time estimates and biogeographic reconstruction indicated that the timing of the three tropical Magnolia disjunctions coincided with the Mid-Eocene cooling climate and/or Late-Eocene climate deterioration, and two temperate disjunctions occurred much later, possibly during the warm periods of Miocene, hence supporting the boreotropical flora concept of Magnolia s.l.
Issue Information
• Issue Information

  J Syst Evol. 2022, 60(1): 1-4.
  https://doi.org/10.1111/jse.12764

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Letter to the Editor
• Exploring the impact of RNA editing on mitochondrial phylogenetic analyses in liverworts, an early land plant lineage

  Shan-Shan Dong, Hong-Lei Li, Bernard Goffinet, and Yang Liu

  J Syst Evol. 2022, 60(1): 16-22.
  https://doi.org/10.1111/jse.12706

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  Plant mitochondrial protein-coding genes are slow evolving, and therefore are less subject to substitution saturation, and therefore perhaps more suitable for resolving deep relationships of high taxonomic categories. Plant mitochondrial genes hold, however, hundreds of RNA editing sites, involving mostly non-synonymous substitutions in the first and second codon positions, which has been reported to affect phylogenetic reconstructions. We have previously identified ca. 4700 mitochondrial RNA editing sites within a group of liverworts representing the ordinal diversity of liverworts, allowing us to critically evaluate the impacts of RNA editing sites on phylogenetic reconstructions in liverworts. Our phylogenetic inferences are mostly congruent on topology inferred from the original mitochondrial gene dataset, dataset with RNA editing sites corrected, and dataset with RNA editing sites excluded. The RNA editing site excluded dataset recovered a topology identical to that of the RNA editing site corrected dataset, supporting the sister relationship of Ptilidium and Jungermanniales, whereas the original dataset supported the sister relationship of Ptilidium and Jungermanniidae. The controversial placements of Ptilidium could be explained by site-wise log-likelihood analysis, as the majority of liverwort RNA editing sites supported the sister relationship of Ptilidium and Jungermanniidae, hence the correction or exclusion of the RNA editing sites changed the tree topology and supported the sister relationship of Ptilidium and Jungermanniales. Our study shows that RNA editing sites potentially impact phylogenetic analyses, suggesting that both genome and transcriptome derived data should be used with caution for phylogenetic reconstruction with genes hosting vast numbers of RNA editing sites such as plant organellar genes.
Research Articles
• Unravelling evolutionary relationships between epifoliar Meliolaceae and angiosperms

  Xiang-Yu Zeng, Rajesh Jeewon, Sinang Hongsanan, Kevin D. Hyde, and Ting-Chi Wen

  J Syst Evol. 2022, 60(1): 23-42.
  https://doi.org/10.1111/jse.12643

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  Epifoliar fungi are a group of poorly studied fungal symbionts that coinhabit the surface of living plants. Meliolaceae is the largest group of epifoliar fungi and has been considered as obligate parasites. We investigated the taxonomy of Meliolaceae and the coevolutionary events with their host plants using time-calibrated cophylogeny based on large subunit, small subunit, and internal transcribed spacer (ITS) sequence data obtained from 17 different fungal taxa and rbcL, ITS, and trnH-psbA sequence data from their corresponding hosts. Nine new fungal species are introduced in this paper and Appendiculella is synonymized under Asteridiella. The dominant coevolutionary events during the Cretaceous and Cenozoic are cospeciation and host shift, respectively. We hypothesize that the evolutionary history of epifoliar fungi can be divided into three major periods: origins of families, formations of genera, and diversification of species. The rise of angiosperms prompted the evolution of modern epifoliar fungi and the diversification of orders of Angiospermae fostered the formation of epifoliar fungal genera. Phylogenetically, epifoliar fungal genera can be delimited according to their coevolutionary patterns and divergent periods.
• 

  Morphological divergence and the Quaternary speciation of Actaea purpurea (Ranunculaceae) and its relatives

  Xiao-Peng Chang, Jiu-Dong Zhang, Xiao-Fei Li, Lei Huang , Xian-Hua Tian, Yi Ren, and Jian-Qiang Zhang

  J Syst Evol. 2022, 60(1): 43-54.
  https://doi.org/10.1111/jse.12667

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  To understand the process and mechanism of speciation, a detailed analysis of origin and demographic history of recently diverged species pairs is necessary. Here, we investigate the evolutionary history of Actaea purpurea (P.K. Hsiao) J. Compton and its closest relatives, A. japonica Thunb. and A. biternata (Siebold and Zuccarini) Prantl. We aim to estimate important parameters of the divergence event, and to lay the foundation for further investigation of the speciation mechanism of this system. Floral and vegetative traits were measured and analyzed. Genetic structure, divergence history, and historical gene flow were also inferred from the plastid and single nucleotide polymorphism data. Floral traits were divergent, and a strong match between pollinator and floral traits was revealed. Genetically the two species were also well diverged, and the time of divergence was dated to the Pleistocene. The demographic modelling results suggest that A. purpurea had continuous limited gene flow with A. japonica and A. biternata since divergence. More work is now needed to confirm that floral trait divergence was selected by pollinators, as well as to understand how pollinator isolation acts in conjunction with other reproductive barriers to reduce gene flow between the two species.

  Plastid and single nucleotide polymorphism data reveals that divergence and speciation of Actaea purpurea occurred in the Pleistocene with limited gene flow with A. japonica and A. biternata. Morphological data also supports the divergence. Actaea purpurea has purple and yellow flowers, in contrast with white in other congeners, and was mainly pollinated by a wasp species. More work is now needed to confirm that floral trait divergence was selected by pollinators, and unravel how pollinators triggered divergence and speciation.
• 

  Phylogenetic relationships within the Mexican genus Bakerantha (Hechtioideae, Bromeliaceae) based on plastid and nuclear DNA: Implications for taxonomy

  Katya J. Romero-Soler, Ivón M. Ramírez-Morillo, Eduardo Ruiz-Sanchez, Claudia T. Hornung-Leoni, Germán Carnevali, and Néstor Raigoza

  J Syst Evol. 2022, 60(1): 55-72.
  https://doi.org/10.1111/jse.12674

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  Bakerantha is one of the three genera of subfamily Hechtioideae (Bromeliaceae). This genus was re-established recently, and it currently contains four species (B. caerulea, B. lundelliorum, B. purpusii, and B. tillandsioides), which are distributed throughout the central region of Mexico. Bakerantha tillandsioides has the widest geographical distribution of the four species, and some populations currently referred to it do not match the species description. In this study, we used extensive sampling (81 accessions) of four plastid regions (matK, rpl32-trnL, rps16-trnK, and ycf1) and the nuclear PRK gene to reconstruct the phylogenetic relationships and delimit the species boundaries in Bakerantha. Our results confirm the monophyly of Bakerantha, and the species delimitation analysis supports five evolutionary lineages within Bakerantha, showing that B. tillandsioides is nonmonophyletic as currently circumscribed. Diagnostic characters and coherent geographical distributions support the five lineages. On the basis of our results, we describe and illustrate B. hidalguense as a new species and provide evidence that B. caerulea is morphologically and ecologically different from B. tillandsioides, with which it has been confused in the past. Additionally, we provide a morphological key to the Bakerantha species.

  The combination of plastid and nuclear markers, and the phylogenetic analysis confirm the monophyly of Bakerantha (Bromeliaceae). The species delimitation analyses showed the existence of five lineages within the genus, all with diagnostic characters and coherent geographical distributions. One of these lineages is here described as a new species, B. hidalguense.
• Comparative analysis of complete plastid genome reveals powerful barcode regions for identifying wood of Dalbergia odorifera and D. tonkinensis (Leguminosae)

  Ming Qin, Cheng-Jie Zhu, Jun-Bo Yang, Mohammad Vatanparast, Rowan Schley, Qiang Lai, Dan-Yan Zhang, Tie-Yao Tu, Bente B. Klitgård, Shi-Jin Li, and Dian-Xiang Zhang

  J Syst Evol. 2022, 60(1): 73-84.
  https://doi.org/10.1111/jse.12598

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  Dalbergia odorifera T. C. Chen (Leguminosae), a rare and endangered tree species endemic to Hainan Island of China, produces the most expensive and rarest wood in China. The wood characteristics of D. odorifera are remarkably similar to those of D. tonkinensis (a much less sought-after species from Vietnam), and the DNA from wood is often highly degraded, making it very difficult to identify the two species using anatomical features or DNA barcoding based on regular DNA markers. To solve the confusion of identifying wood reliably from the two species, we built and analyzed the plastome library of 26 samples from 18 Dalbergia species, of which 12 samples from eight closely related species of D. odorifera are newly sequenced in this study. Phylogenomic analysis suggested that the relationships among the 26 samples are mostly well resolved, and conspecific individuals from different populations of D. odorifera and D. tonkinensis clustered together. Between the plastid genomes of the two species, we identified 129 indels and 114 single nucleotide polymorphisms. By assessing a subset of 20 nucleotide polymorphisms and 10 indels using 37 population-level samples (20 samples of D. odorifera and 17 samples of D. tonkinensis), we recovered eight species-specific barcode regions that could be suitable for identifying the wood D. odorifera and D. tonkinensis. To examine their utility in wood identification, we amplified the eight DNA barcodes using six wood samples and recovered an amplification success rate of 83.3%, demonstrating a reliable method for precise wood identification of the two species.
• 

  Population genetic structure and classification of cultivated and wild pea (Pisum sp.) based on morphological traits and SSR markers

  Rong Liu, Yu-Ning Huang, Tao Yang, Jin-Guo Hu, Hong-Yan Zhang, Yi-Shan Ji, Dong Wang, Guan Li, Chen-Yu Wang, Meng-Wei Li, Xin Yan, and Xu-Xiao Zong

  J Syst Evol. 2022, 60(1): 85-100.
  https://doi.org/10.1111/jse.12710

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  Pea (Pisum sativum L.) is an important legume crop that is widely grown worldwide for human consumption and livestock feed. Despite extensive studies, the population genetic structure and classification of cultivated and wild pea (Pisum sp.) remain controversial. To characterize patterns of genetic and morphological variation and investigate the classification of Pisum, we conducted comprehensive population genetic analyses for 323 accessions from cultivated and wild pea, representing three species of Pisum and utilizing 34 morphological traits and 87 polymorphic simple sequence repeat markers. First, we identified three distinct genetic groups among all samples. Group I was primarily composed of Pisum fulvum, Pisum abyssinicum, and some wild P. sativum accessions, whereas Groups II and III consisted of the two genetic groups under P. sativum that represented different geographic distributions of cultivated pea. Analyses of morphological variation revealed significant differences among the three species. Second, among pea germplasms representing eight taxa of Pisum, P. fulvum and P. abyssinicum possessed unique genetic backgrounds and morphological characteristics, corroborating their independent species status. The intraspecific subdivisions of P. sativum described by some authors were not supported in this study, with the exception of several genotypes of P. sativum subsp. elatius that clustered with P. fulvum and P. abyssinicum. Finally, we confirmed that the Chinese pea germplasm was genetically distinct and could be divided into two genetic groups, each of which included both spring-sowing and autumn-sowing ecotypes. These results provide a robust foundation for understanding pea domestication and the utilization of wild genetic resources of pea.

  Comprehensive population genetic analyses for 286 accessions from cultivated and wild pea within Pisum and utilizing 34 morphological traits and 87 polymorphic SSR markers. Three distinct genetic groups were characterized: Group I was composed of P. fulvum, P. abyssinicum and some wild-type P. sativum; Groups II and III consisted of two cultivated groups under P. sativum and represented different geographic distributions (A, B, C). Significant morphological differences were identified among P. fulvum, P. abyssinicum, and P. sativum (D).
• 

  Large-scale diversity reassessment, evolutionary history, and taxonomic revision of the green macroalgae family Udoteaceae (Bryopsidales, Chlorophyta)

  Laura Lagourgue and Claude E. Payri

  J Syst Evol. 2022, 60(1): 101-127.
  https://doi.org/10.1111/jse.12716

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  Udoteaceae is a morphologically diverse family of the order Bryopsidales. Despite being very widespread geographically, this family is little known as compared with the closely related Halimedaceae or Caulerpaceae. Using the most extensive Udoteaceae collection to date and a multilocus genetic data set (tufA, rbcL, and 18S rDNA), we reassessed the species diversity of the family, as well as the phylogenetic relationships, the diagnostic morphoanatomical characters, and evolutionary history of its genera, toward a proposed taxonomic revision. Our approach included a combination of molecular and morphological criteria, including species delimitation methods, phylogenetic reconstruction, and mapping of trait evolution. We successfully delimited 62 species hypotheses, of which 29 were assigned (existing) species names and 13 represent putative new species. Our results also led us to revise the genera Udotea s.s., Rhipidosiphon s.s., and Chlorodesmis s.s., to validate the genus Rhipidodesmis, and to propose three new genera: Glaukea gen. nov., Ventalia gen. nov., and Udoteopsis gen. nov. We also identified two large species complexes, which we refer to as the “Penicillus–Rhipidosiphon–Rhipocephalus–Udotea complex” and the “Poropsis–Penicillus–Rhipidodesmis complex”. Using a time-calibrated phylogeny, we estimated the origin of the family Udoteaceae at Late Triassic (ca. 216 Ma), whereas most of the genera originated during Paleogene. Our morphological inference results indicated that the thallus of the Udoteaceae ancestor was likely entirely corticated and calcified, composed of a creeping axis with a multisiphonous stipe and a pluristromatic flabellate frond. The frond shape, cortication, and calcification are still symplesiomorphies for most extant Udoteaceae genera and represent useful diagnostic characters.

  ● A reassessment of the Udoteaceae diversity with an unexpected species is carried out and genus richness is highlighted.
  ● A taxonomic revision at the genus level is proposed, and the relevance of diagnostic morphoanatomical characters is re-evaluated.
  ● The origin of the family Udoteaceae is estimated at about 216 Ma (Late Triassic), whereas most of the extant species originated in the Paleogene.
• 

  Multiple Pleistocene refugia and recent diversification for Streptocarpus ionanthus (Gesneriaceae) complex: Insights from multiple molecular sources

  Cornelius M. Kyalo, Ling-Yun Chen, Mathias Lema, Itambo Malombe, Guang-Wan Hu, and Qing-Feng Wang

  J Syst Evol. 2022, 60(1): 128-143.
  https://doi.org/10.1111/jse.12658

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  Streptocarpus ionanthus (Gesneriaceae) is endemic to Tanzania and Kenya, distributed in Tanga, Morogoro, and Kilifi regions. The species houses nine subspecies characterized by complex morphotypes and poorly understood evolutionary relationships, and thus is an ideal model for investigating evolutionary dynamics over time. Using multiple methods, we sought to test our hypothesis that the infraspecific taxa in Str. ionanthus are slightly variable and evolving populations. We first examined the genetic diversity, population differentiation, and phylogeographic structure among the populations of Str. ionanthus using both chloroplast and nuclear markers. We then estimated the divergence time of Str. ionanthus lineages and modeled past and future distribution. Despite Str. ionanthus exhibiting bottleneck events across its range, the populations maintain relatively high genetic diversity attributed to historical population admixture or local adaptation arising from habitat heterogeneity. The phylogeographic and genetic structure revealed a high connection among the Usambara mountains populations, while molecular dating suggested most diversification of haplotypes began ~1.32–0.18 million years ago and intensified toward the present, a conclusion of recent diversification. Phylogenetic relationship of Str. ionanthus cpDNA haplotypes revealed five main lineages with unique haplotypes that could be suggestive of past isolated refugia during the Pleistocene climate shifts. According to niche modeling, the stability of suitable areas during the Last Glacial Maximum (LGM) offered protective micro-habitats that have preserved the genetic diversity of Str. ionanthus to date. In conclusion, our findings suggest a complex Str. ionanthus with slightly variable lineages or populations attributed to multiple refugia and on the verge of divergence.

  Streptocarpus ionanthus (Gesneriaceae) is a morphologically diverse complex with low molecular variation. This study investigated the genetic diversity and phylogeographic structure among the populations of Str. ionanthus. Phylogeography and genetic structure analyses revealed poor structuring, weak genetic clusters, and high haplotype connection. Several slightly variable lineages with unique haplotypes were discovered, suggestive of multiple historical refugia.
• 

  Ancestral gene duplications in mosses characterized by integrated phylogenomic analyses

  Bei Gao, Mo-Xian Chen, Xiao-Shuang Li, Yu-Qing Liang, Dao-Yuan Zhang, Andrew J. Wood, Melvin J. Oliver, and Jian-Hua Zhang

  J Syst Evol. 2022, 60(1): 144-159.
  https://doi.org/10.1111/jse.12683

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  Mosses (Bryophyta) are a key group occupying an important phylogenetic position in land plant (embryophyte) evolution. The class Bryopsida represents the most diversified lineage, containing more than 95% of modern mosses, whereas other classes are species‐poor. Two branches with large numbers of gene duplications were elucidated by phylogenomic analyses, one in the ancestry of all mosses and another before the separation of the Bryopsida, Polytrichopsida, and Tetraphidopsida. The analysis of the phylogenetic progression of duplicated paralogs retained on genomic syntenic regions in the Physcomitrella patens genome confirmed that the whole‐genome duplication events WGD1 and WGD2 were re‐recognized as the ψ event and the Funarioideae duplication event, respectively. The ψ polyploidy event was tightly associated with the early diversification of Bryopsida, in the ancestor of Bryidae, Dicranidae, Timmiidae, and Funariidae. Together, four branches with large numbers of gene duplications were unveiled in the evolutionary past of P. patens. Gene retention patterns following the four large‐scale duplications in different moss lineages were analyzed and discussed. Recurrent significant retention of stress‐related genes may have contributed to their adaption to distinct ecological environments and the evolutionary success of this early‐diverging land plant lineage.

  A graphical representation of the main highlights of the manuscript entitled “Ancestral gene duplications in mosses characterized by integrated phylogenomic analyses” by Gao et al.
Review
• 

  Genomic architectural variation of plant mitochondria—A review of multichromosomal structuring

  Zhi-Qiang Wu, Xue-Zhu Liao, Xiao-Ni Zhang, Luke R. Tembrock, and Amanda Broz

  J Syst Evol. 2022, 60(1): 160-168.
  https://doi.org/10.1111/jse.12655

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  Since the endosymbiont origin from α-Proteobacteria, mitochondrial genomes have undergone extremely divergent evolutionary trajectories among eukaryotic lineages. Compared with the relatively compact and conserved animal mitochondrial genomes, plant mitochondrial genomes have many unique features, especially their large and complex genomic arrangements. The sizes of fully sequenced plant mitochondrial genomes span over a 100-fold range from 66 kb in Viscum scurruloideum to 11 000 kb in Silene conica. In addition to the typical circular structure, some species of plants also possess linear, and even multichromosomal, architectures. In contrast with the thousands of fully sequenced animal mitochondrial genomes and plant plastid genomes, only around 200 fully sequenced land plant mitochondrial genomes have been published, with many being only draft assemblies. In this review, we summarize some of the known novel characteristics found in plant mitochondrial genomes, with special emphasis on multichromosomal structures described in recent publications. Finally, we discuss the future prospects for studying the inheritance patterns of multichromosomal plant mitochondria and examining architectural variation at different levels of taxonomic organization—including at the population level.

  A simplified schematic diagram from an insect (A) and plant species (B) with multichromosomal mt genomes.
Research Articles
• Functional divergence of Populus MYB158 and MYB189 gene pair created by whole genome duplication

  Peng-Fei Jiang, Hui Xu, Chao-Nan Guan, Xiao-Xia Wang, Ai-Min Wu, Yan-Jing Liu, and Qing-Yin Zeng

  J Syst Evol. 2022, 60(1): 169-185.
  https://doi.org/10.1111/jse.12595

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  Whole genome duplication (WGD) provides new genetic material for genome evolution. After a WGD event, some duplicates are lost, while other duplicates still persist and evolve diverse functions. A particular challenge is to understand how this diversity arises. This study identified two WGD-derived duplicates, MYB158 and MYB189, from Populus tomentosa. Populus MYB158 and MYB189 had expression divergence. Populus tomentosa overexpressing MYB158 or MYB189 had similar phenotypes: creep growth, decreased width of xylem and secondary cell wall thickness. Compared to wild-type, neither myb158 mutant nor myb158 myb189 double mutant showed obvious phenotypic variation in P. tomentosa. Although MYB158 and MYB189 proteins could repress the same structural genes involved in lignin, cellulose, and xylan biosynthesis, the two proteins had their own specific regulatory targets. Populus MYB158 could act as the upstream regulator of secondary cell wall NAC master switch and directly represses the expression of the SND1-B2 gene. Taken together, Populus MYB158 and MYB189 have retained similar functions in negatively regulating secondary cell wall biosynthesis, but have evolved partially distinct functions in direct regulation of NAC master switch, with MYB158 playing a more crucial role. Our findings provide new insights into the evolutionary and functional divergence of WGD-derived duplicate genes.
• Pollinator shifts along an elevation gradient mediate different response in self-pollination in heterostylous Primula nivalis

  Aysajan Abdusalam, Wan-Jin Liao, Zhi-Qiang Zhang, and Qing-Jun Li

  J Syst Evol. 2022, 60(1): 186-195.
  https://doi.org/10.1111/jse.12639

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  Pollinator shift and its influence on floral traits have been well documented to explain the diversity of angiosperms, but such effects are still less known at the intraspecific level, especially the responses of different morphs of distylous plants. We hypothesized that the pollen transfer efficiency would decrease if the pollinator shifted from a long-tongued to short-tongued insect across populations, and plants would evolve towards selfing in response to the stressed pollination environment. Given the gender specialization between flower morphs, the long styled (L-morph) plants would increase female reproduction. Our study showed that the short-tongued Bombus tianshanicus, the most-visit pollinator in high elevation populations of distylous Primula nivalis, was less efficient in pollen transfer than long-tongued Bombylius major. The plants evolved to promote selfing through reducing the anther–stigma separation and increasing intramorph self-compatibility. The hand pollination experiment showed that after intramorph selfing, the fruit set and seed set increased with increasing elevation. Moreover, anther and stigma were closer in the L-morph than in the short styled (S-morph) plants, and the L-morphs showed higher pollen transfer efficiency than the S-morphs. Along with increased self-compatibility, the fruit set and seed set of L-morph plants were significantly higher than those of S-morph plants. We described the pollinator shifts along an elevation gradient in a distylous plant and the response of plants by promoting selfing, which confirmed our hypothesis and supported the pollinator-shift model. Our study also highlighted the different response in self-compatibility between flower morphs.
• Equivocal evidence for a change in balance between selfing and pollinator-mediated reproduction in annual Brassicaceae growing along a rainfall gradient

  Eleanor V. J. Gibson-Forty, Katja Tielbörger, and Merav Seifan

  J Syst Evol. 2022, 60(1): 196-207.
  https://doi.org/10.1111/jse.12637

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  Because many plants ensure their reproductive success using some level of self-fertilization, it is predicted that the balance between pollinator-mediated pollen transfer and autonomous selfing will be correlated with habitat harshness and pollinator activity. To study these correlations, we used three annual species that differ in their self-incompatibility and compared their reproductive success in the form of fruit production and seed viability, with and without pollinators, along a rainfall gradient and in relation to pollinator activity. We found that pollinator activity decreased with increasing aridity. Although the reproductive success in the absence of pollinators fitted the species’ level of self-incompatibility, no clear trend was detected between pollinator visits or site harshness and relative reproductive success. Only one of the species, Sinapis alba, showed a consistent increase in autonomous selfing with both increased aridity and decreased pollinator visits, a finding that was supported by an increase in the viability of seeds produced in the absence of pollinators. Nevertheless, unlike our prediction, the absolute number of seeds produced per S. alba flower increased with aridity. Our overall findings suggest that despite decreasing pollinator abundance and diversity with increasing aridity, the reproductive success of annual species, even of those with high self-incompatibility, remains relatively high. Although there may be several mechanisms governing these results, the overall constant seed production despite environmental uncertainties may be part of the mechanism by which plants in arid ecosystems buffer environmental variations.
• 

  Production and evolution pattern of “fruity smell” aggregation pheromones in genus Drosophila

  Wu-Fan Zhang, Lu-Mei Liu, Shan He, Bin-Yan Lu, and Yi-Bo Luo

  J Syst Evol. 2022, 60(1): 208-219.
  https://doi.org/10.1111/jse.12648

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  Insects produce pheromones to serve a range of ecological functions throughout their lifetime. The chemical composition, production pattern, and interspecies specificity provide information for carrying out their function and biological significance. Several species of Drosophila produce a class of volatile esters considered as “fruity smells”; however, the production pattern and ecological functions of these “fruity smell” volatiles in genus Drosophila are poorly understood. Here, using the headspace solid-phase microextraction (SPME) method, we tested the production pattern of volatile pheromones in Drosophila immigrans and factors that possibly affected pheromone production, including mating, feeding conditions, age of adult flies, and geographical distribution. We also explored the evolution and production pattern of volatile pheromones in 14 species of genus Drosophila. Our result showed that male D. immigrans adult flies produce three male-specific volatile ester pheromones, which are also considered as “fruity smell” chemicals, in a relatively stable pattern. In addition, a series of “fruity smell” ester pheromones with similar structure and chemical properties were found to appear in the species of D. virilis and D. immigrans species group, but not in the species of D. melanogaster species group. The ester volatile pheromone production of male flies has a correspondence with the female's demand for host plants. Integrating the production and evolution pattern of these volatile chemicals, we inferred the interaction between insects and host plants reflected in the Drosophila “fruity smell” pheromones.

  We infer the production pattern of Drosophila “fruity smell” pheromones, including ethyl tiglate and ethyl hexanoate. These pheromones are found to appear in the species of D. virilis and D. immigrans species group, but not in the species of D. melanogaster species group. These male-specific volatile pheromones have a correspondence with the demand for host plant in female flies.
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  Fossil pinnae, sporangia, and spores of Osmunda from the Eocene of South China and their implications for biogeography and paleoecology

  Xiao-Yan Liu, Yong-Dong Wang, Li Wang, Ning Zhou, Li-Qin Li, and Jian-Hua Jin

  J Syst Evol. 2022, 60(1): 220-234.
  https://doi.org/10.1111/jse.12653

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  The genus Osmunda L. contains approximately 10 extant species widely distributed in tropical and temperate regions, with the greatest concentration of species in East and Southeast Asia. Osmunda is characterized by dimorphic or commonly hemidimorphic fronds with dimorphic pinnae. Its geological history has been traced back to the Triassic. Most records of the genus are based on rhizomes and rarely on pinnae bearing sporangia and spores. Here, we describe fossil pinnae, sporangia, and spores of Osmunda lignitum (Giebel) Stur recovered from the middle Eocene of the Changchang Formation in the Changchang Basin, Hainan Island and the Youganwo Formation in the Maoming Basin, Guangdong, South China. The fossils closely resemble the extant Osmunda banksiifolia (C. Presl) Kuhn of the subgenus Plenasium on the basis of their morphological and anatomical structures. The present occurrence of O. lignitum indicates subg. Plenasium flourished and extended from the high latitude regions such as Northeast China to the low latitude areas of South China during the middle Eocene. Large numbers of specimens described here also indicate that Osmunda was the dominant understory fern element beneath mixed evergreen broad-leaf angiosperm and gymnosperm forests living in a warm and humid environment.

  The lowest latitudinal occurrence of the genus Osmunda, O. lignitum (Giebel) Stur, is reported to closely resemble the extant O. banksiifolia (C. Presl) Kuhn of the subgenus Plenasium based on fossil pinnae, sporangia, and spores recovered from the middle Eocene of South China. This occurrence indicates subg. Plenasium flourished and extended from the high latitude regions to the low latitude areas of South China during the middle Eocene. Large numbers of specimens suggest that Osmunda was the dominant understory fern element beneath mixed evergreen broad-leaf angiosperm and gymnosperm forests living in a warm and humid environment.
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  J Syst Evol. 2022, 60(1): 235-236.
  https://doi.org/10.1111/jse.12822

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  The Journal of Systematics and Evolution would like to acknowledge and thank the following reviewers for their contributions in the period January 1–December 31 in 2021:

  Allen, Geraldine

  Appelhans, Marc

  Areces-Berazain, Fabiola

  Averianov, Alexander O.

  Bai, Wei-Ning

  Bard, Nicholas

  Bell, Charles

  Besnard, Guillaume

  Bisang, Irene

  Bitencourt, Cássia

  Bouckaert, Remco

  Brandrud, Marie Kristine

  Buellesbach, Jan

  Câmara, Paulo

  Cameron, Kenneth

  Cannon, Chuck

  Carta, Angelino

  Chen, Jun

  Chen, Xiao-Yong

  Chen, Zhi-Duan

  Christie, Kyle

  Corlett, Richard

  Dering, Monika

  Ding, Wen-Na

  Durka, Walter

  Ebersbach, Jana

  Edwards, Christine

  Engle-Wrye, Nick

  Eriksson, Torsten

  Escudero, Marcial

  Fady, Bruno

  Favre, Adrien

  Fazekas, Aron

  Ford, Kerry

  Franco, Fernando

  Gao, Jiang-Yun

  Gao, Lian-Ming

  Gauquelin, Thierry

  Ge, De-Yan

  Ge, Xue-Jun

  Gerrath, Jean

  Goertzen, Leslie

  Gong, Yan-Bing

  Gradstein, Robbert

  Harris, AJ

  Hoffmann, Matthias H.

  Howard, Cody

  Huang, Ji-Hong

  Huang, Jia-Xing

  Huang, Shuang-Quan

  Huarte, Roberto

  Hultine, Kevin R.

  Ickert-Bond, Stefanie

  Ignatov, Michael

  Ikeda, Hajime

  Jiang, Hong-En

  Jiménez-Mejías, Pedro

  Jin, Xiao-Hua

  Johnson, Steven D.

  Kainulainen, Kent

  Kandasamy, Dineshkumar

  Kang, Long-Li

  Kang, Ming

  Klahs, Phillip

  Klopper, Ronell

  Kooyman, Robert

  Ksepka, Daniel

  Kutanan, Wibhu

  Landrein, Sven

  Lepais, Olivier

  Li, Bo

  Li, Jianhua

  Li, Lin-Feng

  Li, Xin-Xin

  Li, Zhong-Hu

  Liang, Er-Yuan

  Liao, Wan-Jin

  Liu, Bo-Ling

  Liu, Xiao-Yan

  Liu, Zhi-Jin

  Liu, Zhi-Yong

  Lou, Hai-Yi

  Luo, A-Rong

  Luo, Yi-Bo

  Ma, Liang

  Ma, Yong-Peng

  Mao, Jian-Feng

  Mao, Kang-Shan

  Márquez-Corro, José Ignacio

  Martín-Bravo, Santiago

  McFarlane, Terry

  McNeal, Joel

  Michelangeli, Fabian A.

  Miller, Joseph

  Mondal, Mayukh

  Morgan, John

  Musser, Grace

  Nawal, Shrestha

  Oh, Sang-Hun

  Olofsson, Jill

  Ortiz, Edgardo

  Passalacqua, Nicodemo

  Pennell, Matthew

  Phillips, Matthew

  Pipes, Leonore

  Pócs, Tamás

  Qian, Hong

  Qiu, Ying-Xiong

  Ranker, Tom

  Razafimandimbison, Sylvain G.

  Rebollo, Roberto

  Ree, Richard

  Ren, Guang-Peng

  Ren, Zong-Xin

  Roalson, Eric

  Roeser, Martin

  Ruiz-Sanchez, Eduardo

  Saarela, Jeffery

  Salazar-Mendias, Carlos

  Sánchez-Vilas, Julia

  Sanmartin, Isabel

  Santos-Gally, Rocio

  Schlueter, Philipp

  Schneider, Harald

  Schwery, Orlando

  Shrestha, Nawal

  Shi, Tao

  Šmarda, Petr

  Sniderman, Kale

  Snow, Neil

  Song, Gang

  Song, Yi-Gang

  Soreng, Robert

  Sosa, Pedro A.

  Stark, Lloyd

  Stedje, Brita

  Su, Tao

  Su, Xu

  Sun, Hang

  Sun, Kun

  Sun, Miao

  Sun, Yan-Xia

  Sun, Yong-Shuai

  Sundue, Michael

  Thiv, Mike

  Thomas, Daniel

  Tian, Bao-Liang

  Tribble, Carrie

  Triplett, Jimmy

  Vaillancourt, Rene

  Vallès, Joan

  van der Merwe, Marlien

  Vanderpoorten, Alain

  Villar, Jose

  von Balthazar, Maria

  Vorontsova, Maria S.

  Walker, Joseph F.

  Wang, Bao-Sheng

  Wang, Heng-Chang

  Wang, Hongru

  Wang, Jing

  Wilson, Paul

  Wu, Yong-Jie

  Wu, Zhi-Qiang

  Xiang, Chun-Lei

  Xiang, Qiao-Ping

  Xie, Shu-Lian

  Xing, Yao-Wu

  Xu, Qing

  Xu, Xiao-Ting

  Yan, Yujing

  Yang, Shi-Xiong

  Yang, Ying-Bo

  Yu, Yan

  Zander, Richard

  Zhang, De-Zhi

  Zhang, Xiao-Ming

  Zhang, Zhi-Yong

  Zhao, Shi-Lei

  Zhou, Ren-Chao

  Zu, Pengjuan