Rooting Portions of a Young Pseudosporochnalean from the Catskill Delta Complex of New York

Abstract

Premise of the Research: Pseudosporochnales (Cladoxylopsida) were conspicuous elements of the Earth's earliest forests. Recent evidence has done much to clarify basic aspects of the pseudosporochnalean architecture, but important questions remain about the developmental processes responsible for growth from juvenile individuals to trees of sometimes considerable size.Methodology: Presented here is combined compression/permineralization evidence of a young member of the group from a late Devonian (early Frasnian) locality also containing Eospermatopteris (Wattieza), currently the largest reconstructed pseudosporochnalean tree. Standard pyrite preparations were made and analyzed with reflected light.Pivotal Results: The anatomically preserved portion of the trunk with expanded base lacking a central vascular column shows abundant evidence of appendages with apparent rooting function supplied by traces comprised of primary and often secondary xylem. Traces arise within parenchyma near trunk center and follow lax courses with multiple divisions outward and downward to the surface, finally enveloping the plant base for some distance. In the upper portion of the specimen, likely near the transition between base bearing rooting appendages and aerial shoot, the traces form a vascular plexus toward the periphery of the stem, with the bulk of vascular tissues comprising secondary xylem. Similar but differently oriented vascularization also occurs near the base.Conclusions: Here we hypothesize a unique form of "bipolar" development in this specimen, and potentially all pseudosporochnaleans, by means of a trunk base bearing an appendicular system of positively geotropic rooting appendages. In addition, we hypothesize that diffuse meristematic activity of the base plus the vascular plexus may have a previously unrecognized role in development of pseudosporochnaleans from the small specimen observed here to large body size. We also suggest that this tissue also offers an explanation for the enigmatic genus Xenocladia known from tissue fragments of large size found in coeval marine sediments of New York State, USA. Given current incomplete understanding of development within the Pseudosprochnales, considering the rooting system as "sui generis" confers the advantage of adequate description of this organ, without necessarily specifying correspondence or homology with other groups.