Oral Paper

         Paleobotany

Ovuliferous complexes of Wollemia nobilis and the dual bundle system of conifer seed cones

Presenting Author
Kelly Matsunaga
Description
The relationship between the anatomy of modern conifer seed cones and the organization of axillary ovuliferous shoots of fossil conifers was central to Rudolph Florin’s foundational model of seed cone homology. In particular, the presence of two series of vascular bundles within the ovuliferous (bract-scale) complexes of modern seed cones was of special significance. In cones with clear differentiation of bract and ovuliferous scale, one set of bundles supplies the bract and have adaxial-abaxial polarity consistent with leaf homology (adaxial xylem, abaxial phloem). The second set of bundles supplies the ovuliferous scale but is inverted relative to the bract bundles: polarity is reversed and phloem is adaxial. Importantly, this dual bundle system is also present in taxa without clear differentiation of bract and scale (e.g., Cupressaceae, Araucariaceae), an observation central to the hypothesis that ovuliferous complexes in modern conifers are homologous and evolved through modification of an axillary shoot and its subtending bract. However, it remains unclear what these bundles correspond to structurally and their evolutionary significance, and thus further study of the development and variation of the dual bundle system is essential for understanding the homology and evolution of conifer seed cones. Here we describe the anatomy and vascular architecture of ovuliferous complexes in Wollemia nobilis, for which these traits have never been formally documented. Individual ovuliferous complexes were fixed, embedded in paraffin, serially sectioned, and stained using a modified triple-stain protocol derived from Sam (1983, Ann. Bot. vol. 51). Ovuliferous complexes are broad and parenchymatous basally, becoming narrow and more sclerotic distally. In the specimen studied, they are vascularized by a single bundle, which branches continuously to form a row of ~9-10 bundles distally. The seed is vascularized by 4 inverted bundles that are adaxial to the main bundle system. Two of these bundles branch from lateral bundles near the base of the ovuliferous complex and the other two diverge from medial bundles. The inverted bundles initially diverge laterally but subsequently rotate to produce the inverted orientation, resulting in phloem that points towards the adaxial side of the ovuliferous complex. The vasculature entering the seed appears to contain only phloem. Vascular anatomy of Wollemia ovuliferous complexes is similar to that of Agathis and Araucaria and falls within the range of variation documented for these genera. The inverted bundle system is more complex than that of Agathis australis and is more similar to A. macrophylla and some species of Araucaria. Unlike most Araucaria, the upper bundle system does not persist beyond the seed. Study of additional specimens will be necessary to understand intraspecific variation of these traits in Wollemia nobilis, but the pattern is nevertheless consistent with the presence of an ovuliferous scale that is reduced, fully fused to the bract, and does not extend beyond the seed. This study fills a conspicuous gap in our knowledge of the anatomy of Araucariaceae, providing data necessary for understanding the evolution, homology, and fossil record of conifer seed cones.