Graduation Date

Spring 2024

Document Type



Master of Science degree with a major in Biology

Committee Chair Name

Dr. Alexandru Mihai Tomescu

Committee Chair Affiliation

HSU Faculty or Staff

Second Committee Member Name

Dr. Terry Henkel

Second Committee Member Affiliation

HSU Faculty or Staff

Third Committee Member Name

Dr. Kelly Matsunaga

Third Committee Member Affiliation

Community Member or Outside Professional

Fourth Committee Member Affiliation

Community Member or Outside Professional


Devonian, Wyoming, Beartooth Butte, Cottonwood Canyon, Plant fossil, Compression fossil, Zosterophyll

Subject Categories



The fragmentary state of plant fossils and the modular organization of plants make whole-plant reconstructions of fossil species desirable and feasible. Such reconstructions are key for integrating fossil species in systematic studies. The ca. 410 Ma Beartooth Butte Formation of Wyoming (USA) hosts the only rich Early Devonian plant assemblages in western North America, which fills a major gap in the phytogeography of this interval. I construct a whole-plant concept for a new zosterophyll from the Beartooth Butte Formation based on a detailed morphometric study. More than 600 fragments of the new zosterophyll were observed and 200 of those were measured. Epidermal features were observed in cuticular material recovered on cellulose acetate. Resulting data were used to characterize variability in axis thickness and taper, branching density, and sporangial orientation, based on descriptive statistics, correlations, and principal component analysis. The new zosterophyll, Nowenia wyomingense gen. et sp. nov., consisted of decumbent axes with long internodes that exhibited K-branching and bore more densely branched erect axes, as well as branches with delayed development (dormant lateral meristems and circinate protrusions). Axes had circinate apices and bore isolated bivalvate sporangia in small numbers, primarily on upright portions. Phylogenetic analyses recover Nowenia sister to Forania, from which it differs primarily in the absence of spinescent protrusions. Nowenia is the first zosterophyll for which an empirically-based whole-plant concept, making explicit and extensive use of quantitative data, is available. The approach used to reconstruct the Nowenia plant introduces a method for integrating morphometric data in constructing whole-plant concepts of early tracheophytes with simple body plans. Application of this and similar methods to additional fossil species could produce reconstructions at similar levels of detail and accuracy, crucial for reaching well-supported resolution of early vascular plant relationships.

Citation Style


ElAbdallahEtAl2024_SupplementalDatasets.xlsx (45 kB)
Spreadsheets including raw data



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