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Hymenophyllaceae (filmy ferns), with ca. 430 species, are the most species-rich family of early diverging leptosporangiate ferns but have a poor fossil record dating back to the Late Triassic period. Traditionally, Hymenophyllaceae comprise two species-rich genera or clades: Hymenophyllum (hymenophylloids) and Trichomanes sensu lato (s.l.) (trichomanoids). Unequivocal fossils of Hymenophyllum have been reported from the Early Cretaceous of central Mongolia and the early Eocene of Okanogan Highlands, Washington, USA. However, despite being a highly diversified lineage with an estimated 184 extant species, Trichomanes s.l. lack a definitive fossil record, which severely affects the reliability of the molecular dating of this group. Here, we report the first unequivocal fossil record of Trichomanes s.l. as T. angustum comb. nov. on the basis of fertile material with tubular involucres and long exserted receptacles from the mid-Cretaceous Kachin amber, Myanmar. This species was previously tentatively assigned to Hymenophyllites due to a lack of fertile evidence. Inferred to be an epiphytic fern, T. angustum further enriches the species diversity of the epiphytic palaeocommunities in the mid-Cretaceous Kachin amber, which are mainly composed of Porellalean leafy liverworts and Dicranalean and Hypnodendralean mosses. Fossil records indicate that Hymenophyllaceae probably originated in the tropical Pangea at the latest in the Triassic when all continents were coalesced into a single landmass and had already accumulated some notable diversity in low-middle latitude areas of Laurasia by the mid-Cretaceous period. [ABSTRACT FROM AUTHOR]

Hymenophyllaceae (filmy ferns), with ca. 430 species, are the most species-rich family of early diverging leptosporangiate ferns but have a poor fossil record dating back to the Late Triassic period. Traditionally, Hymenophyllaceae comprise two species-rich genera or clades: Hymenophyllum (hymenophylloids) and Trichomanes sensu lato (s.l.) (trichomanoids). Unequivocal fossils of Hymenophyllum have been reported from the Early Cretaceous of central Mongolia and the early Eocene of Okanogan Highlands, Washington, USA. However, despite being a highly diversified lineage with an estimated 184 extant species, Trichomanes s.l. lack a definitive fossil record, which severely affects the reliability of the molecular dating of this group. Here, we report the first unequivocal fossil record of Trichomanes s.l. as T. angustum comb. nov. on the basis of fertile material with tubular involucres and long exserted receptacles from the mid-Cretaceous Kachin amber, Myanmar. This species was previously tentatively assigned to Hymenophyllites due to a lack of fertile evidence. Inferred to be an epiphytic fern, T. angustum further enriches the species diversity of the epiphytic palaeocommunities in the mid-Cretaceous Kachin amber, which are mainly composed of Porellalean leafy liverworts and Dicranalean and Hypnodendralean mosses. Fossil records indicate that Hymenophyllaceae probably originated in the tropical Pangea at the latest in the Triassic when all continents were coalesced into a single landmass and had already accumulated some notable diversity in low-middle latitude areas of Laurasia by the mid-Cretaceous period.

Filmy ferns (Hymenophyllaceae) are the most diverse lineage of the early-diverging leptosporangiate ferns with ca. 430 species widely distributed around the world but with the highest diversity in the humid tropics. However, their fossil record is scarce because of the low preservation potential of the delicate, membranous laminae. So far, no Hymenophyllaceae fossils have been reported from tropical Asia. Here, we describe some fern remains and their syninclusions (spike-mosses) in four pieces of Kachin amber from the mid-Cretaceous of Hukawng Valley, Northern Myanmar, as Hymenophyllites angustus sp. nov., H. kachinensis sp. nov., H. setosus sp. nov. (Hymenophyllaceae) and Selaginella alata sp. nov. (Selaginellaceae), respectively. These fern remains are assigned to Hymenophyllaceae based on the filmy, one-cell thick, decompound pinnatifid laminae and dichotomous venation. They represent the first fossil record of Hymenophyllaceae in tropical Asia. The growth habits of these ferns and associated spike-mosses and their implication for paleoenvironment are discussed. Our study expands the diversity of the cryptogams in mid-Cretaceous Kachin amber. Together with other contemporaneous findings, the present fossils indicate that Hymenophyllaceae have already accumulated some notable diversity in the Cretaceous.

Filmy ferns (Hymenophyllaceae) are the most diverse lineage of the early-diverging leptosporangiate ferns with ca. 430 species widely distributed around the world but with the highest diversity in the humid tropics. However, their fossil record is scarce because of the low preservation potential of the delicate, membranous laminae. So far, no Hymenophyllaceae fossils have been reported from tropical Asia. Here, we describe some fern remains and their syninclusions (spike-mosses) in four pieces of Kachin amber from the mid-Cretaceous of Hukawng Valley, Northern Myanmar, as Hymenophyllites angustus sp. nov., H . kachinensis sp. nov., H . setosus sp. nov. (Hymenophyllaceae) and Selaginella alata sp. nov. (Selaginellaceae), respectively. These fern remains are assigned to Hymenophyllaceae based on the filmy, one-cell thick, decompound pinnatifid laminae and dichotomous venation. They represent the first fossil record of Hymenophyllaceae in tropical Asia. The growth habits of these ferns and associated spike-mosses and their implication for paleoenvironment are discussed. Our study expands the diversity of the cryptogams in mid-Cretaceous Kachin amber. Together with other contemporaneous findings, the present fossils indicate that Hymenophyllaceae have already accumulated some notable diversity in the Cretaceous.