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Sarcodon in the Neotropics: New Species from Belize, Colombia

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Sarcodon in the Neotropics: New Species from Belize, Colombia SARCODON IN THE NEOTROPICS: NEW SPECIES FROM BELIZE, COLOMBIA, GUYANA, AND PUERTO RICO By Arthur Charles Grupe II A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Science Biology Committee Membership Dr. Terry Henkel, Committee Chair Dr. Erik Jules, Committee Member Dr. Matthew Smith, Committee Member Dr. Michael Mesler, Committee Member, Graduate Coordinator July 2015 ABSTRACT SARCODON IN THE NEOTROPICS: NEW SPECIES FROM COLOMBIA, BELIZE, GUYANA, AND PUERTO RICO Arthur Charles Grupe II Eight species of the ectomycorrhizal (ECM) genus Sarcodon (Bankeraceae, Thelephorales, Basidiomycota) are described as new to science. Sarcodon quercophilus, and Sarcodon umbilicatus are described from Quercus (Fagaceae) cloud forests within the Maya Mountains of Belize. Sarcodon pakaraimensis is described from forests dominated by the ECM trees Pakaraimaea dipterocarpacea (Dipterocarpaceae) and Dicymbe jenmanii (Fabaceae subfam. Caesalpinioideae) in the Pakaraima Mountains of Guyana. Sarcodon portoricensis is described from lower montane wet forest within the El Yunque National Forest of Puerto Rico. Sarcodon colombiensis, Sarcodon rufogriseus, Sarcodon pallidogriseus, and Sarcodon bairdii are described from the Colombian Amazon in forests dominated by Pseudomonotes tropenbosii (Dipterocarpaceae) and “white sand” forests dominated by species of Dicymbe (Fabaceae subfam. Caesalpinioideae) and Aldina (Fabaceae subfam. Papilionoideae). The discovery of these eight new species is significant given that the majority of the approximately 87 previously described Sarcodon species are north temperate or boreal in distribution and frequently associate with coniferous host plants; these constitute the most recent records ii for Sarcodon from the greater Neotropics. Each of the new species is morphologically consistent with accepted diagnostic characters for Sarcodon: pileate-stipitate stature, a dentate hymenophore, determinate basidiomatal development, fleshy, non-zonate context, and brown, tuberculate basidiospores. DNA (ITS) sequence analysis corroborated the generic placement of S. pakaraimensis, S. portoricensis, S. quercophilus, S. umbilicatus, S. colombiensis, S. rufogriseus, S. pallidogriseus, and Sarcodon bairdii, and along with morphological differences, supported their recognition as distinct species. Macromorphological, micromorphological, habitat, and DNA sequence data from the nuc rDNA internal transcribed spacer region (ITS) are provided for each of the new species. iii ACKNOWLEDGEMENTS The author thanks the following for financially supporting this project: the George Verback Scholarship from the Humboldt Bay Mycological Association, Sonoma County Mycological Society Graduate Student Scholarship, CSUPERB Travel Scholarship, and the Greg Jennings Memorial Scholarship. A special thank you goes out to all the people over my time at Humboldt State University that contributed to my personal and professional development. Some of these people are: J. Uehling, D. Husbands, N. Najarian, I. Zacher, M. DeSiervo, A. Sahara, E. Wu, K. Ludwig, S. Brandt, S. McGuire, E. DeStigter, S. Faulkner, J. Jackson, E. Alvey, P. Lei, J. Mola, Y. Wenzel, L. McCrigler, M. Reed, A. Baker, L. Weaver, C. Satter, G. Shelton, K. Matsunaga, E. Grupe, S. Gowan, and M. Friedman. Field assistance in Guyana was provided by C. Andrew, P. Joseph, F. Edmond, and L. Edmond. Research permits that were granted by the Guyana Environmental Protection Agency, and Ministry of Amerindian Affairs. I would also like to acknowledge the contributions of my committee members. Dr. Erik Jules for giving me my first research opportunities, and letting me borrow his prized Grateful Dead tapes. Dr. Michael Mesler for his encouragement in my development as a teacher and researcher, and for informing me that I would not be alone in “Pollinator Hell”. Dr. Matthew Smith for always being willing to help me with molecular data and problem solving. Dr. Terry Henkel for always pushing me to better myself in my academic career, showing me what it takes to execute a successful iv international field expedition, how to spell “Pakaraimaea”, and those times we performed at the Black Alice show. Most of all I would like to thank my family: my sister Alison Cardenas, my parents Ann Weber, Dan Weber, and Arthur S. Grupe for their unwavering support and belief in me, no matter what. v TABLE OF CONTENTS ABSTRACT........................................................................................................................ ii ACKNOWLEDGEMENTS............................................................................................... iv LIST OF TABLES............................................................................................................ vii LIST OF FIGURES ......................................................................................................... viii INTRODUCTION .............................................................................................................. 1 CHAPTER 1 SARCODON IN THE NEOTROPICS I: NEW SPECIES FROM GUYANA, PUERTO RICO, AND BELIZE...................................................................... 3 Introduction..................................................................................................................... 4 Materials and Methods ................................................................................................... 6 Results............................................................................................................................. 9 Taxonomy ..................................................................................................................... 12 CHAPTER 2 Sarcodon in the Neotropics II. A NEW SPECIES AND DISTRIBUTION RECORD FROM COLOMBIA........................................................................................ 45 Introduction................................................................................................................... 46 Materials and Methods ................................................................................................. 48 Results........................................................................................................................... 49 Taxonomy ..................................................................................................................... 52 LITERATURE CITED ..................................................................................................... 75 vi LIST OF TABLES Table I. Diagnostic morphological characters of new Sarcodon species from Guyana, Puerto Rico, Belize, and Colombia………………………………………………………19 vii LIST OF FIGURES Figure 1. Most parsimonious phylogram (384 steps; mid-point rooted) based on internal transcribed spacer (ITS) ribosomal DNA sequences depicting phylogenetic relationships of Neotropical Sarcodon species. Support values above the nodes are maximum likelihood bootstrap support values whereas values below the nodes are maximum parsimony support values. Nodes with bootstrap support values less than 75 are not shown. Sequences for all Sarcodon species found outside of the Neotropics were obtained from GenBank with numbers shown next to species epithets. .......................... 11 Figure 2. Basidiomata of Sarcodon pakaraimensis (HOLOTYPE; Henkel 9513). A. Developmental series. B. Basidioma showing the black auto-oxidation and staining reaction on the pileus. C. Longitudinal section showing pink staining of the exposed pileus trama, and hollow stipe with internal squamules. Bar = 10 mm............................ 16 Figure 3. Basidia and basidiospores of Sarcodon pakaraimensis (HOLOTYPE; Henkel 9513). Bar = 10 µm........................................................................................................... 17 Figure 4. Scanning electron micrographs of basidiospores of new Neotropical Sarcodon species (×4000). A. Sarcodon pakaraimensis (HOLOTYPE; Henkel 9513). B. Sarcodon portoricensis (HOLOTYPE; Baroni 8776). C. Sarcodon quercophilus (HOLOTYPE; CFMR-BZ-3833). D. Sarcodon umbilicatus (HOLOTYPE; Baroni 10201). ×4000 .......................................................................................................................................... 21 Figure 5. Basidiomata of Sarcodon portoricensis (HOLOTYPE; Baroni 8776). Bar = 10 mm…………………………………………………………………………………….....25 Figure 6. Basidia and basidiospores of Sarcodon portoricensis (HOLOTYPE; Baroni 8776). Bar = 10 µm……................................................................................................... 26 Figure 7. Basidiomata of Sarcodon quercophilus (HOLOTYPE; CFMR-BZ-3833). A. Dorsal view. B. Ventral view, showing pale hymenophore. Bars = 10 mm..................... 32 Figure 8. Basidia, basidiospores, and stipitipellis terminal cells of Sarcodon quercophilus (HOLOTYPE; CFMR-BZ-3833). Bar = 10 µm................................................................ 33 Figure 9. Basidiomata of Sarcodon umbilicatus (HOLOTYPE; Baroni 10201). Bar = 10 mm. ................................................................................................................................... 39 viii Figure 10. Basidia and basidiospores of Sarcodon umbilicatus (HOLOTYPE; Baroni 10201). Bar = 10 µm......................................................................................................... 40 Figure 11. Most parsimonious phylogram (384 steps; mid-point rooted) based on internal transcribed
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