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ArticleAnnals of botany2026

The taphonomic puzzle of Notothylacites filiformis: reinterpretation of bryophyte fossils from the Late Cretaceous of Czechia reveals the co-occurrence of hornworts and liverworts.

Veronika Veselá, Marcela Svobodová, Veronika Kadlecová, Jiřina Dašková, Jiří Kvaček

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Article in Annals of botany, 2026. The graph could read no effect estimate from its abstract, so it casts no vote on the map. Not yet cited in PubMed.

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5 authors.

Veronika VeseláFaculty of Science, Institute of Geology and Palaeontology, Charles University in Prague, Praha 2, 120 00, Czech Republic.
Marcela SvobodováInstitute of Geology, Department of Palaeontology, The Czech Academy of Sciences, Praha-Lysolaje 165 00, Czech Republic.
Veronika KadlecováFaculty of Science, Institute of Geology and Palaeontology, Charles University in Prague, Praha 2, 120 00, Czech Republic.
Jiřina DaškováInstitute of Geology, Department of Palaeontology, The Czech Academy of Sciences, Praha-Lysolaje 165 00, Czech Republic.
Jiří KvačekNational Museum, Prague, Department of Palaeontology, Praha 1, 110 00, Czech Republic.

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6 · The paper itself

Abstract

BACKGROUND AND

aimsHornworts are rarely found in the fossil record, so each new find provides important insights regarding their evolution and diversity. Here we revisit a controversial genus, Notothylacites, described from the Late Cretaceous of Central Europe, which has liverwort morphology but bears hornwort spores.

methodsThe fossil material was originally studied in 1970 by Pacltová using pollen preparation techniques. Using a single grain method, we extracted spores from the preparation matrix, and observed them with scanning electron microscopy. For comparison, we studied spores of the extant hornwort genus Notothylas. KEY

resultsWe reinterpret the original Notothylacites filiformis holotype as a conglomerate of two fossils: a hornwort, Notothylacites, and a liverwort, which we designate as a new species Ricciopsis pacltovae sp. nov. Notothylacites filiformis was lectotypified, using material from seven slides and the original drawing, which emphasized a key character - a sporangium with apical longitudinal dehiscence slit containing psilate, trilete spores. Ricciopsis pacltovae, erected based on comparisons with seven described species, is a liverwort that closely resembles extant Riccia, with a prostrate circular thallus consisting of dichotomously branched, linear segments, and up to six dichotomies per branch, each branch with a pronounced narrow medial sulcus. These and other morphological traits distinguish it from previously described species in the genus.

conclusionsNotothylacites filiformis is reinterpreted as one of the rare examples of a fossil hornwort sporophyte. Gametophyte material previously assigned to Notothylacites filiformis is assigned to a new fossil liverwort species, Ricciopsis pacltovae. Hornworts frequently grow on wet soil together with Riccia liverworts. The co-occurrence of the two fossil species further highlights the presence of bryophyte diversity in the Late Cretaceous that mirrors extant ecological associations. This study corrects decades of taxonomic confusion, demonstrating how taphonomic mixing can mislead interpretations, and emphasizes the importance of re-examining collections with modern methods.

Indexed as

BryophytaFossilsHepatophytaBiological EvolutionCzech RepublicMicroscopy, Electron, ScanningSporesAnthocerotophytaCentral EuropeFossil bryophytesLate CretaceousMarchantiophyta

Identifiers

PMID40926531
PMCPMC13275004

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