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Water Course Vegetation on Granitic and Calcareous Substrates in the Eastern Mojave Desert, California

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Water Course Vegetation on Granitic and Calcareous Substrates in the Eastern Mojave Desert, California WATER COURSE VEGETATION ON GRANITIC AND CALCAREOUS SUBSTRATES IN THE EASTERN MOJAVE DESERT, CALIFORNIA by Julie M. Evens A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment of the Requirements for the Degree Master of Arts In Biology May, 2000 WATER COURSE VEGETATION ON GRANITIC AND CALCAREOUS SUBSTRATES IN THE EASTERN MOJAVE DESERT, CALIFORNIA by Julie M. Evens We certify that we have read this study and that it conforms to acceptable standards of scholarly presentation and is fully acceptable, in scope and quality, as a thesis for the degree of Master of Arts. Approved by the Master's Thesis Committee P. Dawn Goley Todd Keeler-Wolf Michael R. Mesler Milton J. Boyd, Graduate Coordinator Approved by the Dean of Graduate Studies Ronald A. Fritzsche ABSTRACT Desert water courses are topographically distinct features with periodic flooding, which are floristically rich and structurally complex. This study was undertaken to determine if plant species and vegetation types change along desert water courses as environmental gradients are crossed. In four granitic and four calcareous water courses, vegetation patterns were classified and described for 262 samples in the eastern Mojave Desert, California. In a broad altitudinal range from mountain slope to bajada, vegetation samples were systematically taken at 300 m intervals in a 9 km distance of each water course. A total of 15 alliances and 32 associations were delineated using Ward's method cluster analysis and Twinspan and were validated by Bray-Curtis ordination. The classification included alliances of one forest, two woodlands, two intermittently flooded shrublands, two temporarily flooded shrublands, and eight shrublands. Of these alliances, ten were at the canyon position, seven were at the arroyo position, and six were at wash position. Alliances dominated by Acacia greggii, Chilopsis linearis, and Prunus fasciculata were the most widely distributed, at more than one topographic position and on both granitic and calcareous substrates; however, these alliances had more than double the number of associations found specifically on limestone as compared to granite. Environmental gradients showed strong relationships to the overall vegetation patterns as expressed by multi-response permutations procedures, Indicator Species analyses, and Bray-Curtis ordinations. Vegetation types appear to be expressions of moisture and temperature gradients, nutrient availability, and regional locations as iii functions of aspect, elevation, geologic substrate, geographic position, surface rock cover, and topographic position. The influence of topography and geologic substrate was apparent across all watercourses. Each water course had differential species and at least one distinct alliance per topographic position, and the water courses had differential species and vegetation types depending on geologic substrate. iv ACKNOWLEDGMENTS First and foremost, I thank John Sawyer, my major professor, for his faithful support and guidance. I have greatly benefited from his organizational and editorial skills as well as his research expertise. I also thank Dawn Goley and Michael Mesler, two committee members, for their valuable insights and constructive comments. I extend further thanks to Todd Keeler-Wolf, a third committee member, for providing inspiration, data collection design, technical support, and insightful review of this manuscript. I am also thankful for the gracious support from the following entities. Peter Stine of the U.S. Geological Survey of Biological Resources Division and Humboldt State University of Biological Sciences contributed financial support and equipment. Additionally, the California Native Plant Society provided financial support. Jim Andre and Claudia Luke of the University of California at Granite Mountains Reserve contributed housing, lab facilities, research design, and field assistance. The Mojave National Preserve granted office support and equipment as well as access to study sites and collection of plants. Mitchell Caverns State Park granted access to the Gilroy Canyon study site and collection of plants. Andy Sanders of the University of California at Riverside Herbarium and Jim Andre assisted in plant identification. I am grateful to numerous other people who have given support and field assistance. My parents provided financial support during my graduate studies. Jeffrey Cox assisted with research design, reconnaissance, and data management. Alex Evens and Jane Melville assisted in field work. Lastly, I thank my family and friends for their enduring emotional support that allowed me to stay focused and determined. TABLE OF CONTENTS Page ABSTRACT iii ACKNOWLEDGMENTS LIST OF TABLES viii LIST OF FIGURES LIST OF APPENDICES xii INTRODUCTION 1 STUDY AREA 3 Location 3 Climate 5 Landforms and geology 13 METHODS 22 Sampling 22 Data analyses 27 RESULTS 35 Direct analyses of the complete datasets 35 Classification and ordination of the complete dataset 45 Classification of canyon dataset 47 Classification of arroyo/wash dataset 53 Bray-Curtis ordination of the canyon datasets 57 Bray-Curtis ordination of the arroyo/wash datasets 67 vi TABLE OF CONTENTS (CONTINUED) Page Relationship of vegetation types to existing classifications 71 Vegetation descriptions 75 DISCUSSION 99 Environmental gradients 99 Relationship to existing classifications 108 CONCLUSIONS 112 LITERATURE CITED 114 vii LIST OF TABLES Table Page 1 Elevation, mean temperature, and percent of mean annual precipitation in the summer and winter seasons at three weather stations in the eastern Mojave Desert, California 8 2 Calcareous rock units in the study area 18 3 Granitic rock units in the study area 19 4 Sedimentary deposits in the study area 21 5 Criteria to establish coverage class values for sampling 24 6 Ranked classes for aspect 26 7 Total and mean number (± 1 S.D.) of vascular species per 0.1 ha in the eight water courses on granite and limestone in the complete dataset .. 36 8 Mean species percent cover (± 1 S.D.) per 0.1 ha in the water courses on granite and limestone in the complete dataset 37 9 Elevation range and mean percent slope (± 1 S.D.) per 0.1 ha in the eight water courses on granite and limestone in the complete dataset 38 10 Mean percent cover of rock and sediment (± 1 S.D.) per 0.1 ha in the water courses on granite and limestone in complete dataset 39 11 Multi-response permutation procedures for topographic position in the complete dataset (p<0.00000 for each analysis) 41 12 Indicator species and values for geologic substrate in the complete dataset 42 13 Indicator species and values for topographic groups in the complete dataset 44 viii LIST OF TABLES (CONTINUED) Table Page 14 Coefficients of determination (r2) between distances in the ordination space and distances in the original n-dimensional space for the canyon dataset and the arroyo/wash dataset 60 15 Pearson and Kendall correlations of species with the ordination axes for the canyon dataset 61 16 Pearson and Kendall correlations of environmental variables with the ordination axes in the canyon dataset 63 17 Indicator species and values for geologic substrate in the canyon dataset 66 18 Pearson and Kendall correlations of species with the ordination axes in the arroyo/wash dataset 68 19 Pearson and Kendall correlations of environmental variables with the ordination axes in the arroyo/wash dataset 69 20 Indicator species and values for geologic substrate in the arroyo/wash dataset 72 21 Source and environmental characteristics of the alliances (with at least three replicate samples) 73-74 ix LIST OF FIGURES Figure Page 1 Location of the study area in the Mojave Desert, California 4 2 Monthly mean precipitation at three weather stations in the eastern Mojave Desert, California. 7 3 Mean annual precipitation at the Granite Cove weather station in the Granite Mountains, California, from 1986 to 1999 9 4 Monthly maximum/minimum mean temperature and mean precipitation at the Granite Cove weather station in the Granite Mountains, California, averaged from 1986 to 1999 10 5 Bray-Curtis diagram of 258 vegetation samples in the complete dataset based on topographic position 46 6 Ward's dendrogram for the 74 canyon samples 48 7 Ordination diagram of Ward's method groups at the four-cluster level in the canyon dataset 50 8 Ordination diagram of Twinspan groups at the four-cluster level in the canyon dataset 50 9 Bray-Curtis ordination diagram for alliances of 74 canyon plots of the eastern Mojave Desert, California 51 10 Bray-Curtis ordination diagram for associations of 74 canyon plots of the eastern Mojave Desert, California 52 11 Ward's dendrogram for the 181 arroyo and wash samples 54-55 12 Ordination diagram of Ward's method groups at the four-cluster level in the arroyo/wash dataset 56 13 Ordination diagram of Twinspan groups at the four-cluster level in the arroyo/wash dataset 56 LIST OF FIGURES (CONTINUED) Figure Page 14 Bray-Curtis ordination diagram for alliances of 181 arroyos and washes of the eastern Mojave Desert 58 15 Bray-Curtis ordination diagrams for associations of 181 arroyos and washes of the eastern Mojave Desert 59 16 Bray-Curtis ordination diagram of canyon alliances with overlay of environmental and floristic variables (r>0.5) 64 17 Bray-Curtis ordination diagram of arroyo and wash alliances with overlay of environmental and floristic variables (r>0.5) 70 xi LIST OF APPENDICES Appendix Page A Species list of plants
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