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Tricostate Mosses and Associated Bryophilous Fungi from The TRICOSTATE MOSSES AND ASSOCIATED BRYOPHILOUS FUNGI FROM THE EARLY CRETACEOUS OF VANCOUVER ISLAND (CANADA) By Glenn W.K. Shelton A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Science in Biology Committee Membership Dr. Alexandru M.F. Tomescu, Committee Chair Dr. Ruth A. Stockey, Committee Member Dr. Stephen C. Sillett, Committee Member Dr. Michael R. Mesler, Graduate Coordinator May 2015 ABSTRACT TRICOSTATE MOSSES AND ASSOCIATED BRYOPHILOUS FUNGI FROM THE EARLY CRETACEOUS OF VANCOUVER ISLAND (CANADA) Glenn W.K. Shelton Two novel fossil mosses (Tricosta plicata gen. et sp. nov. and Krassiloviella limbelloides gen. et sp. nov.) and several fungal morphotypes associated with one of these mosses are described here based on anatomically preserved material from the Early Cretaceous (Valanginian-136 Ma) Apple Bay locality of Vancouver Island (Canada). The mosses provide additions to the extremely short list of anatomically preserved bryophytes currently described from the pre-Cenozoic and have tricostate leaves (i.e., bearing three leaf veins or costae per leaf), a trait only known in a few other Mesozoic fossil mosses. Although the sporophytes of Tricosta and Krassiloviella are unknown, the high level of anatomical detail preserved in these fossils allows for whole-plant reconstructions of the gametophytes and the recognition of a new family of pleurocarpous mosses, Tricostaceae fam. nov. The bryophyte-inhabiting (bryophilous) fungi, known exclusively in association with the gametophytes of T. plicata, are represented by ii several distinct morphologies including: epiphyllous hyphae and appressoria, intracellular haustoria and cell aggregates, and endophytic perithecioid fruiting bodies embedded within shoot tips. This is the first occurrence of bryophilous fungi documented in the fossil record. Both the moss gametophytes and the bryophilous fungi are common throughout the Apple Bay material, yet inconspicuous. Their characterization holds promise for future studies of anatomically preserved material in terms of documenting past bryofloras and mycofloras, both of which are fascinating but little-studied components of paleo- ecosystems. iii ACKNOWLEDGEMENTS I would like to thank my committee for their perseverance in making it through this tome the with still a healthy mind for comments and discussion. A special thanks goes to my advisor, Dr. Alexandru “Mihai” Tomescu and all of the HSU Biology Department staff and faculty who have helped shape me academically throughout the past five years as an undergraduate and Master’s Candidate. It should also be said that, the inspirational environment and solid work ethic found in the “Lanphere-Reiss-Tomescu Lab” (in part, my office), were great motivators for me during my years there (2010- 2015). This work is supported in part by: the Natural Sciences and Engineering Research Council of Canada grant A-6908 to R. A. Stockey; the HSU Biology Department’s Master’s Student Grant; and the Gregory Mark Jennings Award (HSU Biology). iv TABLE OF CONTENTS ABSTRACT ........................................................................................................................ ii ACKNOWLEDGEMENTS ............................................................................................... iv LIST OF TABLES ........................................................................................................... viii LIST OF FIGURES ........................................................................................................... ix INTRODUCTION .............................................................................................................. 1 TRICOSTA PLICATA GEN. ET SP. NOV. ...................................................................... 4 Fossil Record of Pre-Cenozoic Mosses .......................................................................... 4 Materials and Methods .................................................................................................... 7 Systematics ..................................................................................................................... 9 Description .................................................................................................................... 12 Habit, branching, shoot architecture, and stem anatomy .......................................... 12 Leaf morphology and anatomy ................................................................................. 20 Specialized branches ................................................................................................. 29 Discussion ..................................................................................................................... 35 The tricostate condition ............................................................................................. 35 Tricostate analogues in extant mosses ...................................................................... 36 Tricostate mosses in the fossil record ....................................................................... 38 Taxonomic placement of Tricosta plicata gen. et sp. nov. ....................................... 40 Pleurocarpous mosses in the pre-Cenozoic fossil record .......................................... 48 Gametangia in the fossil record ................................................................................ 51 Outstanding questions ............................................................................................... 53 Conclusions ................................................................................................................... 56 v KRASSILOVIellA LIMBELLOIDES GEN. ET SP. NOV.............................................. 58 Introduction ................................................................................................................... 58 Materials and Methods .................................................................................................. 59 Systematics ................................................................................................................... 61 Description .................................................................................................................... 64 Habit, branching, shoot architecture, and stem anatomy .......................................... 64 Leaf morphology and anatomy ................................................................................. 72 Specialized branch .................................................................................................... 80 Discussion ..................................................................................................................... 84 The tricostate condition ............................................................................................. 84 Taxonomic placement of Krassiloviella limbelloides gen. et sp. nov. ..................... 84 Moss-animal interactions .......................................................................................... 88 Conclusions ................................................................................................................... 88 IMPLICATIONS OF TRICOSTA AND KRASSILOVIA .............................................. 90 BRYOSYMBIOTIC FUNGI ............................................................................................ 92 Introduction ................................................................................................................... 92 Materials and Methods .................................................................................................. 93 Description of Fungal Morphotypes ............................................................................. 95 Tubular septate hyphae ............................................................................................. 95 Moniliform hyphae ................................................................................................. 105 Intracellular cell aggregations ................................................................................. 109 Phragmospores and dictyospores ............................................................................ 114 Aperturate amerospores .......................................................................................... 118 vi Perithecioid fruiting bodies ..................................................................................... 118 Interfoliar stromata .................................................................................................. 121 Haustoria and other fungal types ............................................................................ 124 Discussion ................................................................................................................... 127 Ascomycete specialists ........................................................................................... 127 Biotrophs or decomposers? ..................................................................................... 128 Life history of the Tricosta plicata bryophilous fungi ............................................ 129 Systematics ............................................................................................................. 131 Conclusions ................................................................................................................. 135 REFERENCES ..............................................................................................................
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