Checklist of fossil liverworts suitable for calibrating phylogenetic reconstructions

Abstract

The number of available liverwort fossils substantially increased within the past decade, which is mainly due to new findings from Cretaceous and Cenozoic amber deposits. Many of them, however, are fragmentary and not predestined for consideration in evolutionary analyses. Here, we list those liverwort fossils that we suggest as suitable for calibrating phylogenetic reconstructions, along with brief descriptions, justification of their use, and age information. Our recommendations are based on thorough microscopic investigation of available fossils from worldwide amber collections including recent findings. We recommend that the following 42 fossil taxa can be used as confident minimum age constraints in phylogenetic reconstructions: Acrolejeunea ucrainica (35 Ma), Anastrophyllum rovnoi (35 Ma), Bazzania polyodus (34 Ma), Blepharolejeunea obovata (15 Ma), the genus Bryopteris with B. bispinosa and B. succinea (15 Ma), Calypogeia stenzeliana (34 Ma), Cephaloziella nadezhdae (35 Ma), the genus Ceratolejeunea with C. antiqua, C. palaeomexicana, and C. sublaetefusca (15 Ma), Cheilolejeunea latiloba (34 Ma), Cheirorhiza brittae (158 Ma), Cololejeunea sp. (15 Ma), Cyclolejeunea archaica (15 Ma), Dibrachiella grollei (15 Ma), Diettertia montanensis (112 Ma), Drepanolejeunea eogena (15 Ma), the genus Frullania with F. baerlocheri, F. cretacea, and F. partita (99 Ma), Frullania subgen. Frullania with F. casparyi (34 Ma) and F. riclefgrollei (35 Ma), F. subgen. Trachycolea with F. rovnoi (35 Ma) and F. schumannii (34 Ma), Gackstroemia cretacea (99 Ma), Geocalyx heinrichsii (34 Ma), the genus Lejeunea with L. hamatiloba, L. miocenica, L. resinata, and L. urbanioides (15 Ma), Lopholejeunea subnigricans (15 Ma), Marchantites cyathodoides (228 Ma), Marchesinia brachiata (15 Ma), Metzgeriothallus sharonae (383 Ma), Microlejeunea nyiahae (52 Ma), Neurolejeunea macrostipula (15 Ma), Nipponolejeunea europaea (34 Ma), Notoscyphus balticus (34 Ma), Odontoschisma (sect. Iwatsukia) dimorpha (34 Ma), Plagiochila groehnii (34 Ma), Porella subgrandiloba (34 Ma), Protolophozia kutscheri (34 Ma), Radula (subgen. Odontoradula) cretacea (99 Ma), R. (subgen. Amentuloradula) heinrichsii (99 Ma), Scapania hoffeinsiana (34 Ma), Solenostoma berendtii (34 Ma), Spruceanthus polonicus (34 Ma), Stictolejeunea squamata (15 Ma), Tetralophozia groehnii (34 Ma), Thysananthus auriculatus (15 Ma), Thysananthus contortus (34 Ma). Furthermore, we transfer Lophozia kutscheri to Protolophozia, Archilejeunea grollei to Dibrachiella, Frullania ucrainica to Acrolejeunea, and Mastigolejeunea extincta to Spruceanthus, based on new fossil evidence and morphological revisions.

Miocene Ethiopian amber: A new source of fossil cryptogams

“…Within Jungermanniidae, there seems to be some preservation bias. The mainly epiphytic Porellales, and especially Lejeuneaceae and Frullaniaceae, are very diverse, whereas Jungermanniales are represented by significantly less species (Feldberg et al, 2014, 2018, 2021a, 2021b; Heinrichs et al, 2018a, 2018b; Katagiri, 2018; Mamontov et al, 2018, 2019, 2020; Li et al, 2020). The Porellales show several distinct characters, for example, the incubously inserted, complicate lobed lateral leaves with a large dorsal lobe and a smaller ventral lobule, which is generally folded on the ventral side of the dorsal lobe.…”

“…Thysananthus aethiopicus sp. nov. is the fourth extinct species assigned to this genus (Heinrichs et al, 2018a; Feldberg et al, 2021a) (Table 2). The fossil shows similarities to other Thysananthus fossils from Miocene Dominican and Eocene Baltic amber in the general habit but can easily be distinguished (Gradstein, 1993; Grolle & Meister 2004 as Mastigolejeunea ; Sukkharak & Gradstein, 2017; Yu et al, 2020).…”

“…Frullaniaceae are the most diverse group of liverworts found as amber inclusions (Feldberg et al, 2018, 2021a, 2021b; Heinrichs et al, 2018a; Mamontov et al, 2019, 2020; Li et al, 2020). The extant genus Frullania Raddi is represented by 19 fossil species dating from the mid‐Cretaceous to the Miocene.…”

“…Frullaniaceae are the most diverse group of liverworts found as amber inclusions (Feldberg et al, 2018(Feldberg et al, , 2021a(Feldberg et al, , 2021bHeinrichs et al, 2018a;Mamontov et al, 2019Mamontov et al, , 2020Li et al, 2020) Frullania is most diverse in Cenozoic ambers of Europe, with eight currently accepted species from Baltic and Bitterfeld amber (Grolle, 1985b;Grolle & Meister, 2004;Heinrichs et al, 2018a;Feldberg et al, 2018), six species from Rovno amber (Mamontov et al, 2015a(Mamontov et al, , 2017(Mamontov et al, , 2018(Mamontov et al, , 2019(Mamontov et al, , 2020Feldberg et al, 2021a), and one species occurring in all three deposits (Konstantinova et al, 2012) (Table 2). In addition, one representative of Frullania from Miocene Dominican amber was described at the subgeneric level and is probably also present in Rovno amber (Konstantinova et al, 2012) (Table 2).…”

Divergent evolutionary trajectories of bryophytes and tracheophytes from a complex common ancestor of land plants

“…Progress has also been made recently towards resolving the ambiguous phylogenetic relationships at the root of land plants (Cox et Su et al, 2021). Further, the bryophyte fossil record has recently undergone a radical reinterpretation such that there are now many more potential calibrations to constrain the timing of early land plant evolution (Tomescu et al, 2018;Feldberg et al, 2021;Flores et al, 2021). Even in the absence of new fossils, novel methods have been developed to provide calibrations by using horizontal gene transfers to provide relative time constraints (Szöllősi et al, 2020).…”