316 Wilhelm Bock 1969
Other scales illustrated came from additional specimens, of which sub-fig. 6 (fig. 544) suggests a scale placement of the rhomboidal apophysis. The units in fig. 545 are particularly interesting, as they represent two or more scales with some definite outline, being more roundish, or heart-shaped with a quite narrow stalk, which could only yield space for a small seed, if any. They possibly are sterile ones from the very tip of the cone.
The epidermal pattern of this species is rather simple and little differentiated. The surface cells of the foliage follow straight to wavy lines, being squarish to elongate, occasionally polygonal, which may change toward the leaf margin into long-rhomboidal shape. The cell pattern of the male pericone and its cone scales are more or less the same as in the foliage cell structure. The surface of the megasporophyll apophysis is typically wrinkled, or granulated in appearance (fig. 546).
Phylogenetic remarks. Morphogenetically, it is worth mentioning that the fused seed scales and bract, resp. sterile scales, are paramount in Primaraucarians, which are involved in a distinct apophysial structure with a cross ridge, radial rays and a distinct, small rhomboidal umbo resembling greatly those of living species, ushering this upper Triassic tree into a well advanced position. The microsporangial scales could be clearly considered to be derived from the pericone scale leaves and the latter from normal lamina. Obviously, the microsporophylls are directly attached to a morphologically slightly altered leaf scale. What influenced the presumable sex-foliate double leaf of some lamina, is still unclear, although it may have some bearing on the sterile seed scale development.
The most important parallel may be the close similarity of the seed scale unit, found in the Permian Walchiaraucaria. Both possessed a distinct apophysial armor with a small and pointed umbo. The imbedded individual seed was squeezed into bulging portions of the seed scale which were attached to each other below the base of the seed, but apparently were three-sectionaly patterned, lengthwise. The scale unit in Primaraucaria is already closely fused, while the bract scale in Walchiaraucaria may not have been yet completly joined, which should be considered phylogenetic advancement of the former. While the Permian araucarian type is an informative transition form toward the Triassic type, the latter represents an equal transitional position toward the still more advanced recent araucarians, separated, or partially separated bract scale conditions being only expressed in immature stages of the latter. As, therefore, the fused scale conditions are not yet completely met and are obviously in a form of incomplete transformation, Primaraucaria and Walchiaraucaria have not fully attained the standing of the Pinales and are retained in the Voltziales. It further suggests that in this phylogenetic sequence the stalked, more or less pendent seed scale, was on its way to be abandoned, and one of the last survivors appears to be the uppermost Triassic genus Hirmeriella and possibly the new species Glyptolepis delawarensis (516-517).