Palissya – absolutely incomprehensible or surprisingly interpretable: a new morphological model, affiliations and phylogenetic insights.

The morphology of the adaxial structures of cones belonging to Palissya Endlicher 1847 emend. nov. are reinterpreted based on exquisitely preserved permineralised material from the Lower Cretaceous of Queensland. Although the material was not found in situ, it likely derives from the Orallo Formation, which is Valanginian in age. The cones have dual vascular bundles in each bract/scale complex, and the different tissue types in the bract and ovule/scale complex support interpretation of the cone as a compound structure. Since the early twentieth century it has been widely accepted that each ovule is surrounded by a cup-shaped structure, but the detailed morphology of the “cup” has hitherto been unclear. These new three-dimensionally preserved specimens with in situ ovules are described as Palissya tillackiorum sp. nov. This study demonstrates that the “cup” is formed from a pair of thin scales that subtend but are not fused to each ovule; each pair of scales comprises a thicker outer and thinner inner scale. The organographic relationships among ovules and scales in Palissya show a high degree of synorganisation. The adaxial surface of the bract/scale complex has 2–6 pairs of erect (orthotropous) ovules. The ovule/scale units are arranged symmetrically in two parallel rows on either side of the midline of the bract/scale. Individual ovule/scale units are comparable to those seen in extant Podocarpaceae and Taxaceae. The ovules are thin-walled and are interpreted to have a single integument and a non-thickened (non-lignified) micropyle. These new insights allow reinterpretation of material previously referred to Palissya. A new species is described from Yorkshire, England, as P. harrisii C.R. Hill ex Pattemore & Rozefelds sp. nov. All species based on well preserved cones are reconsidered herein: P. sphenolepis (Braun 1843) Nathorst 1908 emend. Florin 1958, P. elegans Parris, Drinnan & Cantrill 1995 emend. nov., P. bartrumii Edwards 1934 emend. nov., P. antarctica Cantrill 2000 and P. hunanensis Wang 2012. Palissya ovalis Parris et al. 1995 differs structurally from Palissya and is transferred to Knezourocarpon Pattemore 2000 emend. nov. Representatives of this genus may superficially resemble those of Palissya in compressions and impressions, and their congeneric status has been previously suggested; hence its inclusion in this study. Knezourocarpon has adaxial processes that are positioned in two parallel rows but it lacks ovules and paired lateral scales that formed a cup-shape, and its processes attach directly to a central vascular trace. The improved understanding of Palissya’s morphology allows for definite separation of these genera, although the higher-order affiliation of Knezourocarpon remains unclear.