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31295014915903.Pdf (3.446Mb) A SYSTEMATIC STUDY OF INDEHISCENT-FRUITED YUCCAS IN THE CHIHUAHUAN DESERT by R. LAURIE ROBBINS, B.S. in ED., M.A., M.A. A DISSERTATION IN BOTANY Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Approved December, 1983 d,^ ,3 ACKNOWLEDGMENTS I deeply appreciate the support of friends and family along the way. I am also indebted to Dr. David K. Northington for his direction on this project and to the members of my committee for their helpful criticism. ii TABLE OF CONTENTS ACKNOWLEDGMENTS ^^ ABSTRACT IV LIST OF TABLES VI LIST OF FIGURES Vll SECTIONS I. INTRODUCTION 1 II. MATERIALS AND METHODS 5 III. RESULTS 10 Yucca baccata and If. endlichiana 10 Yucca camerosana and Y. faxoniana 14 Yucca decipiens and Y. filifera 19 Yucca torreyi and Y_. treculeana 22 IV. DISCUSSION 28 V. TAXONOMIC CONCLUSIONS 31 LITERATURE CITED 33 APPENDICES 35 1. Key to species 35 2. Taxonomic treatment 37 3. Collecting localities by species, country, state, and county 49 4. Specimens examined from other herbaria 55 iii ABSTRACT Plants of the genus Yucca (Family Agavaceae) are strictly of New World distribution, occurring in the U.S., Mexico, and Central America. Because of the great size of many members of the genus, and because of previous inaccessibility of much of their ranges until fairly recently, little has been done to evaluate the variabi­ lity within the group. With the exception of Trelease's (1902) work on the genus, other studies have been done strictly within the con­ fines of political boundaries. This is the first to deal with the group within an ecological unit, namely, the Chihuahuan Desert. Eight taxa of indehiscent-fruited yuccas occur wholly or in part in this area. Of these, six appear as "pairs" that closely resemble each other and possess complimentary ranges (either north- south or east-west). Data and voucher material consisting of gross plant morphology (measurements), flower, fruit, and leaf measurements, and flavonoid patterns from greenhouse-grown seedlings were taken. Analyses of these data provided the basis for evaluating not only the amount of variability present in each of the taxa, but to reevaluate the taxonomic alignment of the taxa, as well. It was found that gross plant morphology and fruit morphology provide the most important characters for separating the taxa. Flowers are extremely conservative within the entire genus, but leaves are highly variable, even within a single taxon. The available data has lead to a realignment of the Y_. carnerosana-Y. faxoniana complex to more accurately reflect the probable species alignment, and a combination iv of two previously recognized species (Y_. torreyi and Y_. treculeana) into a single species (Y. treculeana), LIST OF TABLES 1. Classification of the Yucceae as per Trelease (1902) 53 2. Classification of the yuccas of the southwestern United States as per McKelvey (1938, 1947) 60 3. Classification of the yuccas of the southwestern United States as per Webber (1953) 62 4. Classification of the yuccas of Mexico as per Matuda and Pina L. (1980) 63 5. Measurements (in centimenters) and characters recorded on entire Yucca plants 65 6. Measurements (in millimeters) and determinations taken on each of 10 flowers per Yucca plant 66 7. Measurements (in millimeters) taken from each Yucca fruit 68 8. Measurements (in millimeters) and characters taken from Yucca leaves 69 9. Means, ranges, and standard deviations for 12 morphological characters taken on indehiscent-fruited species of yuccas in the Chihuahuan Desert 70 10. Means, ranges, and standard deviations for flower measurements for species of indehiscent-fruited yuccas in the Chihuahuan Desert 72 11. Means, ranges, and standard deviations for fruit measurements for species of indehiscent-fruited yuccas in the Chihuahuan Desert 75 12. Means, ranges, and standard deviations for leaf measurements for species of indehiscent-fruited yuccas in the Chihuahuan Desert 76 vi LIST OF FIGURES 1. Collection localities, specimens examined, and distribution of "torreyi" (x), "treculeana" (triangles), "Cuatro Cienegas" (closed circles), "Del Rio" (open circles), and "Post" (+) populations used in this study 78 2. Collecting localities, specimens examined, and distribu­ tion of Y. decipiens (circles) and Y. filifera (x) used in this study 80 3. Collecting localities, specimens examined, and distribu­ tion of Y. carnerosana (open circles) and Y. faxoniana (closed circles) used in this study 82 4. Collecting localities, specimens examined, and distribu­ tion of ][. endlichiana (triangle) and Y_. baccata (closed circles) used in this study 84 5. Plot of Factor III against Factor I for whole plant data for Y. baccata (open circles) and Y. endlichiana (closed circles) populations . 86 6. Plot of Factor II against Factor I for fruit data on Y. baccata (open circles) and _Y. endlichiana (closed circles) populations 88 7. Plot of Factor II against Factor I for leaf data of Y, baccata (open circles) and Y. endlichiana (closed circles) populations 90 8. Plot of Factor III against Factor I for whole plant data for populations of Y_. carnerosana (open squares) and Y. faxoniana (closed circles). 92 9. Plot of Factor II against Factor I for flower data on populations of ][. carnerosana (open squares) and Y. faxoniana (closed circles) 94 10. Plot of Factor III against Factor I on leaf data for populations of Y. carnerosana (open squares) and Y. faxoniana (closed circles) 96 11. Plot of Factor II against Factor I of total plant data from populations of Y. filifera (closed circles) and Y. decipiens (open squares) 98 12. Plot of Factor II against Factor I for flower data on populations of Y. filifera (closed circles) and Y. decipiens (open squares). 100 vii 13. Plot of Factor II against Factor I on fruit data from populations of Y. filifera (closed circles) and Y. decipiens (open squares). 102 14. Plot of Factor II against Factor I for leaf data on populations of _Y. filifera (closed circles) and Y. decipiens (open squares) 104 15. Plot of Factor III against Factor I for total plant data for populations of "torreyi" (open circles), "treculeana" (closed circles), "Cuatro Cienegas" (closed triangles), and "Del Rio" (open diamonds) plants 10^ 16. Plot of Factor II against Factor I for flower data of "torreyi" (closed triangles), "Post" (closed squares), "treculeana" (closed circles), "Del Rio" (open circles), and "Cuatro Cienegas" (open diamonds) populations 108 17. Plot of Factor II against Factor I on fruit data for "torreyi" (open circles), "treculeana" (closed circles), "Del Rio" (open diamonds), "Cuatro Cienegas" (closed triangles) populations of yuccas viii A SYSTEMATIC STUDY OF THE INDEHISCENT-FRUITED YUCCAS IN THE CHIHUAHUAN DESERT I. INTRODUCTION The genus Yucca is strictly of New World distribution, occurring throughout most of the United States, Mexico, and Central America. Although traditionally placed in the Liliaceae, Hutchinson (1934) included it in the Agavaceae, a treatment followed here. PiKa L. (1980) recognizes 47 species in the genus. Within the group, nomenclatural puzzles have arisen as a result of several confounding factors. Sargent (1895) summarized the situation rather succinctly in stating "Yuccas are difficult plants for the botanist to manage owing to their size and nature; and it is almost impossible to preserve in herbaria specimens that adequately illustrate the species. Individuals of the same species vary widely in the length and width of their leaves and in the size of their flowers and fruit, so that the seedlings of one plant scattered through the gardens of different countries, might be given different names by botanists familiar only with a few individuals. .... It is not surprising, therefore, that there is hardly a group of garden-plants about which there is so much confusion in books, and this confusion will never be cleared up until the different species are studied extensively in the field, and then gathered in some California or Mexican garden where their development under normal conditions can be patiently watched." Engelmann (1873) was the first to monograph the group. His treatment, consisting of two subgenera and three sections under one of the subgenera, is regarded as valid today. What he (Engelmann, 1871) initially termed sections Sarcocarpa, Clistocarpa, and Chaenocarpa under the subgenus Euyucca, became Sarcoyucca, Clistoyucca, and Chaenoyucca in his later publication (Engelmann, 1873). Within this classification scheme, he placed 14 naturally-occurring species. Baker (1892) maintained this alignment but included only 11 species. It was not until later, however, that Trelease (1902) provided the first major revision of the group and seriously addressed the problems that had arisen as a result of extensive plantings in Europe of materials originating in the New World. Descriptions of new species abound in European horticultural and botanical publications dating back to as early as 1597. Indeed, according to Trelease, one species of a Yucca from the South Carolina-Florida area had been in continuous cultivation in Europe since 1596 and, at the time of his writing, was represented by a considerable number of garden forms. Trelease (1902) took on the difficult task of clarifying the taxonomy to that time. At the same time, he attempted to resolve the taxonomy of what he regarded as "spontaneous" (or naturally-occurring) species in the United States, Mexico, and Central America. Available to him were herbarium specimens (including those obtained from the border surveys) and material from his own travels and collections. Travel in the arid, sparsely populated areas of northern Mexico and the southwestern United States at that time was difficult. When one views Trelease's accomplishments with the appropriate time frame in mind, one can appreciate the enormity of the task representing some 16 years of herbarium, field, and garden work that went into his 1902 treatise. Table 1 gives the classification as arranged by Trelease (1902) in that publication.
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