A new conifer from the Upper Triassic of southern Poland linking the advanced voltzialean type of ovuliferous scale with Brachyphyllum-Pagiophyllum-like leaves

The Triassic is an exceptionally interesting period from the point of view of conifer evolution. The oldest representatives of some modern families appear in the fossil record in the Upper Triassic and all recent conifer families probably originated during this time. The stageswhereby the primitiveVoltziales transformed into the evolutionarily advanced families of modern conifers in the Triassic are yet to be fully documented. A newly discovered Upper Triassic (Norian) locality in Patoka (SW Poland, Upper Silesia) with a unique fossil record offers us a rare opportunity to fill this gap. In this paper, we describe the very well preserved remains of a new conifer from this locality.We have been able to reconstruct thewhole plant and propose a new taxon, Patokaea silesiaca gen. et sp. nov., on the basis of organic attachment and similar cuticular details on leaves, ovuliferous cones, seed scale-bract complexes, ovules and mature seeds as well as polliniferous cones containing pollen grains of the Enzonalasporites type in situ. This plant combines shoots bearing Brachyphyllum-Pagiophyllum-type leaves with a new type of seed scale-bract complex clearly derived from evolutionary advanced Voltziales and polliniferous cones somewhat similar to the Cheirolepidiaceae (Classostrobus) type. Based on this distinctive and hitherto unique combination of features, a new conifer family - Patokaeaceae - has been proposed. The ovuliferous cones of this conifer are lax and borne singly at the end of leafy twigs. Ovule-bearing scales are stalked and trilobate with two lateral oval lobes, each bearing one ovule, and one sterile reduced lobe between them, all in one plane. The bract is small and leaf-like. Polliniferous cones are simple with helically arranged microsporophylls. Five to seven pollen sacs are arranged around the microsporophyll stalk. Polliniferous cones are borne singly at the end of leafy twigs. This is also the first evidence of a relationship between Enzonalasporites pollen and the parent plant. The pollen occurs in the polliniferous cones, in the micropyle inside the ovule and in the micropylar region inside the seed of this new conifer. This plant expands our view of voltzialean conifer diversity at the roots of modern conifer families.