The Effect of Fire on Yellowstone Ecosystem Seed Banks by David
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The effect of fire on Yellowstone ecosystem seed banks by David Lee Clark A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biological Sciences Montana State University © Copyright by David Lee Clark (1991) Abstract: This paper describes the seed banks of fourteen habitat types which span the full spectrum of environments found in the Greater Yellowstone ecosystem, from dry grasslands, through forested zones, to alpine tundra. Seed bank characteristics are also described across two secondary seres in which lodgepole pine was dominant, and for two mid-seral stages of a sere in which whitebark pine was co-dominant. The effects of wildfire on the seed banks were measured. The effect of three specific temperatures (50,100, and 150°C) on germination of seed bank taxa was measured in the laboratory. Seed bank compositio was determined by identifying and enumerating germinants from soil cores collected at three distinct sites in each habitat type. The effect of fire was estimated by comparing samples from adjacent burned and unburned portions of each site. Three successive germination trial periods were used to insure germination of most seeds. Undisturbed seed bank densities averaged about 4000 seeds/m2, with fewer (less than 1000 seeds/m2) occurring in dry grasslands and dense coniferous forests, and more appearing in aspen groves (7000 seeds/nr) and seasonally wet meadows (14000 seeds/m2). Seed bank species richness averaged about 35 taxa/habitat type, with fewer species (10-20) in dry grasslands and dense forests. Fire reduced seed bank density and species richness in nonforested habitat types by approximately 20% and 15%, respectively. In forests, seed bank density and species richness were reduced by approximately 80% and 65%, respectively. The density and diversity of soil seed banks parallel the aboveground density and richness. They are low where resources are limited by climate or competition. Seed bank density declines with stand age in the seres investigated, while species richness and correspondence with the aboveground vegetation remain more nearly constant across each sere. The effect of fire on seed banks of nonforested plant communities is relatively minor. The seed banks of forests are more heavily impacted by fire, and especially in the dense coniferous forests, where undisturbed seed banks are initially depauperate. On such sites, the contribution of buried seeds to postfire revegetation may be limited. THE EFFECT QF FIRE ON YELLOWSTONE ECOSYSTEM SEED BANKS by David Lee Clark A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science i n Biological Sciences MONTANA STATE UNIVERSITY Bozeman, Montana December 1991 ii Z jV 7/f APPROVAL of a thesis submitted by David Lee Clark This thesis has been read by each member of the thesis committee and has been found to be satisfactory regarding content, English usage, format, citations, bibliographic style, and consistency, and is ready for submission to the College of Graduate Studies. Ic^lcW ^ T l Ti AN Date Chairperson, Graduate Committee Approved for the Major Department 2(o KbMeMber AAi Date Head, Major Department Approved for the College of Graduate Studies iii STATEMENT OF PERMISSION TO USE' In presenting this thesis in partial fulfillment of the requirements for a master's degree at Montana State University, I agree that the Library shall make it available to borrowers under rules of the Library. Brief quotations from this thesis are allowed without special permission, provided that accurate acknowledgement of source is made. Permission for extensive quotation from or reproduction of this thesis may be granted by my major professor, or in his absence, by the Dean of Libraries when, in the opinion of either, the proposed use of the material is for scholarly purposes. Any copying or use of the material in this thesis for financial gain shall not be allowed without my written permission. Date i v ACKNOWLEDGMENTS I would like to express my sincere appreciation to those who provided assistance during the course of this study. The encouragement and support given by my supervisor, Or. Don- Despain, Research Biologist, Yellowstone National Park, were essential to the success of the project. ■ The advice and guidance of my major professor, Dr.. Tad Weaver, Biology Department, Montana State University, were equally vital. Gai I Arundel I, Janet Edwards, Mary Emerick, Chris Lyness, Steve Stinnett, and Sarah Welch helped collect soil samples during the initial phase.of the study. Rob Ament assisted with sieving and oven heating of subsamples. Dr. Matt Lavin, Peter Stickney, and Jennifer Whipple helped with the identification of germinahts. Tom and Victoria Smith watered germination flats when I had to be away. My wife, Katharina, assisted with seedling inventories and the preparation of voucher specimens. I thank them all for their generous aid. Statistical analyses were accomplished with MSUSTAT, Version 4.12, developed by Dr. Richard E. Lund, Montana State University, Bozeman, MT 59717. This study was funded by the National Park Service Postfire Research Program. TABLE OF CONTENTS Page LIST OF TABLES....................................................■..............................................vii List of figures........................, ........................................................................ ix ABSTRACT................................................. x INTRODUCTION..........................................................................•..............................I Objectives.......................................................................:.............................2 Literature Review.................................................................. 3 Concepts....................................................................................................3 Seed Banks of Undisturbed Communities...............................................5 Effect of Fire on Seed Banks.......................................... 8 Seed Banks of Successional Series.................................................... 12 Methodology................................... 15 METHODS...............'...............:.................................................................................17 Study Area.......................................................... 17 Habitat Type Selection......................................•.......................................T9 Stipa comata/Bouteloua gracilis (Stco/Bogr)..................................21 Agropyron spicatum/Poa sandbergii-Stipa comata (Agsp/Posa-Stco)............................................................................21 Festuca idahoensis/Aqropyron spicatum (Feid/Agsp)'...........'..........21 Deschampsia caespitosa/Carex spp. (Deca/Carex)...........................23. Pseudotsuga menziesii/Calamaqrostis rubescens (Psme/Caru).................................................................. 23 Pseudotsuga menziesii/Symphoricarpos albus (Psme/Syal)..............23 Populus tremuloides/Calamaqrostis rubescens (Potr/Caru)............25 Artemisia tridentata/Festuca idahoensis (Artr/Feid)...................25 Abies Iasiocarpa/Calamaqrostis rubescens (Abla/Caru).................25 Festuca idahoensis/Aqropyron caninum (Feid/Agca)............ 27 Abies lasiocarpa/Vaccinium scopariurn (Abla/Vasc).........................27 Abies lasiocarpa/Vacciniurn globulare (Abla/Vagl)..................... ..27 Abies lasiocarpa/Vaccinium scoparium-Pinus albicaulis ^Abl a/Vasc-Pi al) ..... ...................... -.28 Festuca ovina/Poa alpina (Feov/Poal)...............................................28 Sampling Site Selection....................................................... 29 Soil Sample Collection................................................................................29 Subsample Treatment..................................................................................... 31 Greenhouse Procedures..................................................................................34 vi -TABLE OF CONTENTS--Contjnued Page RESULTS.................. '...................................-.......................'..................................38 Seed Bank Composition, Density, and Treatment Effects................... 40 Stipa comata/Bouteloua graci Iis.'......................................................40 Aqropyron spicatum/Poa sandberqii-Stipa comata............................ 40 Festuca idahoensis/Agropyron spicatum.............................................41 Desohampsia caespitosa/Carex spp...........■.........................................41 Pseudotsuga menziesii/Calamaqrostis rubescens..............................42 Pseudotsuga menziesii/Symphoricarpos albus....................................43 • Populus tremuloides/Calamaqrostis rubescens..................................44 Artemisia tridentata/Festuca idahoensis.........................................44 Abies Iasiocarpa/Calamaqrostis rubescens.................................. 45 Festuca idahoensis/Agropyron caninum...............................................47 Abies lasiocarpa/Vaccinium scopariurn...............................................48 Abies lasiocarpa/Vaccinium qlpbulare..............................................50 Abies lasiocarpa/Vaccinium scoparium-Pinus albicaulis................51 Festuca ovina/Poa alpina.....................................................................52 Comparison of Vegetation and Seed Bank Composition...........................