Table of Contents
  • Volume 57 Issue 4

    Special issue: Celebrating Research Devoted to Seed-Free Land Plants

    Cover illustration: The images selected for the cover refl ect the focus of the special issue on the biology and phylogenetic history of seed-free land plants. Progress in our understanding of seed-free land plants has been largely achieved by testing established hypotheses using genomic evidence and/or fossil evidence. Until recently, the fossil record of some of these plants has been considered to be poor but exciting discoveries have challenged this assumption. Furthermore, studies using genomic evidence have rej [Detail] ...
    • Harald Schneider
      2019, 57 (4): 303–304
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    • Research Articles
    • Eva-Maria Sadowski, Leyla J. Seyfullah, Ledis Regalado, Laura E. Skadell, Alexander Gehler, Carsten Gröhn, Christel Hoffeins, Hans Werner Hoffeins, Christian Neumann, Harald Schneider, and Alexander R. Schmidt
      2019, 57 (4): 305–328
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      Diverse temperate forest types and a high atmospheric humidity have recently been suggested for the Eocene source area of Baltic amber. However, ferns are astonishingly rare as inclusions in this amber, which is in contrast to other seed‐free land plants, fungi, and lichens. Moreover, the identities of some of the few described putative fern taxa are dubious, and some fossils were even assigned to the Paleozoic seed fern genera Alethopteris, Pecopteris and to the form genus Sphenopteris containing Paleozoic and Mesozoic fern‐like leaf fossils. Here, we review previously described fern inclusions from Baltic amber and identify further fern‐like leaf inclusions as belonging to the extant angiosperm genus Comptonia (sweet ferns, Myricaceae). We conclude that only one taxon, Matonia striata (Matoniaceae), can with confidence be identified as a Polypodiopsida representative. Although “Pecopterishumboldtiana is so far only known as sterile foliage, its leaf morphology strongly suggests that also this taxon belongs to the Polypodiopsida rather than to any other tracheophyte lineage. We propose accommodating “Pecopterishumboldtiana in the new genus Berendtiopteris. “Alethopterisserrata and “Sphenopteris” phyllocladoides are not to be regarded as evidence of ferns from Baltic amber. Reinvestigation of the holotypes of these two taxa did not reveal to which tracheophyte lineages these fossils belong. We suggest that the scarcity of fern remains from Baltic amber may reflect both a relatively low fern diversity in the source area of the fossil resin, and an absence or rarity of epiphytic and climbing ferns as observed in modern temperate forest ecosystems.

    • Ledis Regalado, Alexander R. Schmidt, Patrick Müller, Lisa Niedermeier, Michael Krings, and Harald Schneider
      2019, 57 (4): 329–338

      Divergence time estimates suggest that most clades constituting the fern family Pteridaceae (Polypodiales) were in existence by the Early Cretaceous. However, fossil evidence to corroborate this remains exceedingly rare. Burmese amber is an important source of new information on the radiation of derived fern lineages during the Cretaceous Terrestrial Revolution. This study describes Heinrichsia cheilanthoides gen. et sp. nov., a fern with suggested affinities to Pteridaceae, based on fertile foliage portions preserved in Early Cretaceous (~100 Ma) amber from Myanmar. Heinrichsia cheilanthoides is characterized by a pinnate‐pinnatifid frond that bears apical, marginal sori protected by a pseudoindusium. Sporangia are of the polypod type and contain tetrahedral‐globose, trilete spores with a striate perine. This discovery provides a new calibration point to test and refine molecular clock‐based concepts of the evolutionary history of the Pteridaceae. Heinrichsia cheilanthoides further substantiates the suggestion that the Cretaceous forests of Myanmar were home to a rich fern flora.

    • Yuriy S. Mamontov, and Michael S. Ignatov
      2019, 57 (4): 339–360
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      Three new fossil bryophytes are described from Jurassic and Lower Cretaceous deposits of the Transbaikalia region of Russia. The complex thalloid hepatic Khasurtythallus monosolenioides gen. et sp. nov. belongs to the Marchantiidae, but its combination of characters precludes unequivocal placement in any of the five orders of this subclass, representing most likely an extinct lineage. Paleaethallus squarrosus gen. et sp. nov. is a thalloid plant with scales similar to those of complex thalloid hepatics, although their arrangement and the overall plant structure has little in common with any extant hepatics. Dispersed moss capsules, three of which have attached calyptrae, are described as a form genus Kulindobryum gen. nov. Despite incomplete preservation, some rare characters indicate possible relationship to the genus Tayloria of the Splachnaceae, an extant family of mosses adapted to grow on animal dung, dead bodies and bones. Notably, Kulindobryum co‐occurs with bones of the small feathered dinosaurs Kulindadromeus, which also supports an affiliation of Kulindobryum with the Splachnaceae. The most common and best known Mesozoic moss for the region, the genus Bryokhutuliinia, is appraised for its systematic position and probable affinities with the Dicranales. A scoring approach is introduced for the comparative method of taxonomic placement of fossils with partial suites of morphological characters at the family or order level.
    • Gaik Ee Lee, Julia Bechteler, Tamás Pócs, and Alfons Schäfer-Verwimp
      2019, 57 (4): 361–370
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      A new species from Fiji, Lejeunea heinrichsii is described and illustrated herein based on morphological characters and molecular phylogenetic analyses. The taxonomic descriptions of L. heinrichsii and the comparison with their allies are presented, together with the Maximum Likelihood (ML) and Bayesian phylogenetic analyses based on sequences of two chloroplast regions (trnL‐trnF, rbcL) and the nuclear ribosomal ITS region. This delicate and rather minute species is characterized by its autoicy, eplicate perianths, the oblong‐rectangular leaf lobes, the distantly arranged underleaves with lobes to 1/2‐2/3 underleaf length, and the leaf cells without well‐developed trigones and intermediate thickenings. In the molecular phylogeny, L. heinrichsii is placed sister to the Asian L. compressiuscula from Indonesia and forms an independent lineage including accessions of Neotropical L. adpressa from the Dominican Republic and the widespread Palaeotropical L. anisophylla from São Tomé and Príncipe. Morphologically, L. heinrichsii is very similar to L. capensis, however the result based on the molecular phylogenetic analyses revealed that the two species are very distinct.
    • Silvia Pressel, and Jeffrey G. Duckett
      2019, 57 (4): 371–381
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      To test the premise that fertilization distances in bryophytes are limited by spermatozoid motility, we measured the distances between over 80 000 male and female thalli of Marchantia polymorpha on heathland over 2 successive years after a major fire and recorded the incidence of sporophytes. The first comparative data on spermatozoid numbers per antheridium across bryophytes and pteridophytes were calculated from antheridial sizes and spermatid diameters and cross‐checked with the likely numbers produced by successive antheridial mitoses. Individual antheridia of Marchantia produce over 200 000 spermatozoids. Extrapolating from individual antheridia to the numbers of mature antheridia per antheridiophore and then to individual male plants, we calculated that a single flooding event on a male thallus with 10–12 antheridiophores will result in the release of over 50 million spermatozoids. Assuming radial dispersal in surface water films by lipids released from the dehiscing antheridia, spermatozoids can reach distances exceeding 20 m from the parent plants, in line with our finding of 100% fertilization in female plants of Marchantia up to 19 m from the nearest males, far beyond published fertilization distances in dioicous bryophytes. We attribute this to (i) much greater spermatozoid production and numbers in Marchantia than in other bryophytes and (ii) highly effective sperm transport both within the antheridiophores and archegoniophores via their overlapping scales and grooves in the stalks and between male and female thalli via surface water films. These features, coupled with the massive production of small spores, explain the success of Marchantia as the primary colonist of open habitats.

    • Ying Yu, Hong-Mei Liu, Jun-Bo Yang, Wen-Zhang Ma, Silvia Pressel, Yu-Huan Wu, and Harald Schneider
      2019, 57 (4): 382–394

      Sequencing the plastid genomes of land plants provides crucial improvements to our understanding of the plastome evolution of land plants. Although the number of available complete plastid genome sequences has rapidly increased in the recent years, only a few sequences have been yet released for the three bryophyte lineages, namely hornworts, liverworts, and mosses. Here, we explore the disparity of the plastome structure of liverworts by increasing the number of sequenced liverwort plastomes from five to 18. The expanded sampling included representatives of all major lineages of liverworts including the genus Haplomitrium. The disparity of the liverwort genomes was compared with other 2386 land plant plastomes with emphasis on genome size and GC‐content. We found evidence for structural conservatism of the plastid genomes in liverworts and a trend towards reduced plastome sequence length in liverworts and derived mosses compared to other land plants, including hornworts and basal lineages of mosses. Furthermore, Aneura and Haplomitrium were distinct from other liverworts by an increased GC content, with the one found in Haplomitrium only second to the lycophyte Selaginella. The results suggest the hypothesis that liverworts and other land plants inherited and conserved the plastome structure of their most recent algal ancestors.

    • Marta R. Pereira, Alice Ledent, Patrick Mardulyn, Charles E. Zartman, and Alain Vanderpoorten
      2019, 57 (4): 395–403

      Bryophytes are a group of land plants in which the role of hybridization has long been challenged. Using genotyping by sequencing to circumvent the lack of molecular variation at selected loci previously used for phylogeny and morphology, we determine the level of genetic and morphological divergence and reproductive isolation between the sibling Syrrhopodon annotinus and S. simmondsii (Calymperaceae, Bryopsida) that occur in sympatry but in different habitats in lowland Amazonian rainforests. A clear morphological differentiation and a low (0.06), but significant Fst derived from the analysis of 183 single nucleotide polymorphisms were observed between the two species. Conspecific pairs of individuals consistently exhibited higher average kinship coefficients along a gradient of geographic isolation than interspecific pairs. The weak, but significant genetic divergence observed is consistent with growing evidence that ecological specialization can lead to genetic differentiation among bryophyte species. Nevertheless, the spatial genetic structures of the two species were significantly correlated, as evidenced by the significant slope of the Mantel test based on kinship coefficients between pairs of interspecific individuals and the geographic distance separating them. Interspecific pairs of individuals are thus more closely related when they are geographically closer, suggesting that isolation‐by‐distance is stronger than the interspecific reproductive barrier and pointing to interspecific gene flow. We conclude that interspecific introgression, whose role has long been questioned in bryophytes, may take place even in species wherein sporophyte production is scarce due to dioicy, raising the question as to what mechanisms maintain differentiation despite weak reproductive isolation.

    • Rafael Medina, Matthew G. Johnson, Yang Liu, Norman J. Wickett, A. Jonathan Shaw, and Bernard Goffinet
      2019, 57 (4): 404–417

      Selection on spore dispersal mechanisms in mosses is thought to shape the transformation of the sporophyte. The majority of extant mosses develop a sporangium that dehisces through the loss of an operculum, and regulates spore release through the movement of articulate teeth, the peristome, lining the capsule mouth. Such complexity was acquired by the Mesozoic Era, but was lost in some groups during subsequent diversification events, challenging the resolution of the affinities for taxa with reduced architectures. The Funariaceae are a cosmopolitan and diverse lineage of mostly annual mosses, and exhibit variable sporophyte complexities, spanning from long, exerted, operculate capsules with two rings of well‐developed teeth, to capsules immersed among maternal leaves, lacking a differentiated line of dehiscence (i.e., inoperculate) and without peristomes. The family underwent a rapid diversification, and the relationships of taxa with reduced sporophytes remain ambiguous. Here, we infer the relationships of five taxa with highly reduced sporophytes based on 648 nuclear loci (exons complemented by their flanking regions), based on inferences from concatenated data and concordance analysis of single gene trees. Physcomitrellopsis is resolved as nested within one clade of Entosthodon. Physcomitrella s. l., is resolved as a polyphyletic assemblage and, along with its putative relative Aphanorrhegma, nested within Physcomitrium. We propose a new monophyletic delineation of Physcomitrium, which accommodates species of Physcomitrella and Aphanorrhegma. The monophyly of Physcomitrium s. l. is supported by a small plurality of exons, but a majority of trees inferred from exons and their adjacent non‐coding regions.

    • Hong‐Mei Liu, Libor Ekrt, Petr Koutecky, Jaume Pellicer, Oriane Hidalgo, Jeannine Marquardt, Fatima Pustahija, Atsushi Ebihara, Sonja Siljak‐Yakovlev, Mary Gibby, Ilia Leitch, and Harald Schneider
      2019, 57 (4): 418–430

      Recent studies investigating the evolution of genome size diversity in ferns have shown that they have a distinctive genome profile compared with other land plants. Ferns are typically characterized by possessing medium‐sized genomes, although a few lineages have evolved very large genomes. Ferns are different from other vascular plant lineages as they are the only group to show evidence for a correlation between genome size and chromosome number. In this study, we aim to explore whether the evolution of fern genome sizes is not only shaped by chromosome number changes arising from polyploidy but also by constraints on the average amount of DNA per chromosome. We selected the genus Asplenium L. as a model genus to study the question because of the unique combination of a highly conserved base chromosome number and a high frequency of polyploidy. New genome size data for Asplenium taxa were combined with existing data and analyzed within a phylogenetic framework. Genome size varied substantially between diploid species, resulting in overlapping genome sizes among diploid and tetraploid spleenworts. The observed additive pattern indicates the absence of genome downsizing following polyploidy. The genome size of diploids varied non‐randomly and we found evidence for clade‐specific trends towards larger or smaller genomes. The 578‐fold range of fern genome sizes have arisen not only from repeated cycles of polyploidy but also through clade‐specific constraints governing accumulation and/or elimination of DNA.

Song Ge
Jun Wen
Impact Factor
JCR 2019 IF ranking: 56/234 (Plant Sciences, top 23.72%, Q1 quartile)
Journal Abbreviation: J Syst Evol
ISSN: 1674-4918 (Print)
1759-6831 (Online)
CN: 11-5779/Q
Frequency: Bi-monthly




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