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Systematics of theSchizachyrium scoparium (Poaceae) complex in North America
Bruner, Joe Leonard, Ph.D.
The Ohio State University, 1987
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University Microfilms International SYSTEMATICS OF THE SCHIZACHYRIUM SCOPARIUM (POACEAE) COMPLEX IN NORTH AMERICA.
DISSERTATION
Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University
By
Joe L. Bruner, B.S., M.S.
% * * * *
The Ohio State University 1987
Reading Committee: Approved by: Dr. Daniel J. Crawford Dr. Gary L. Floyd Dr. Katherine L. Gross Aavisor Dr. Tod F. Stuessy Department of Botany to Tom - warm friend who introduced me to the mind's eye ACKNOWLEDGEMENTS
A number of people should be acknowledged for their contributions during the course of this study. I must thank Dr. Daniel J. Crawford, my major professor, for his patience and continued encouragement over the past 7 years. My committee, Drs. Daniel J. Crawford, Gary L. Floyd, Katherine L. Gross and Tod F. Stuessy provided valuable comments and criticisms on the dissertation. I thank the Department of Botany and Sigma Xi for their grants and financial aid that made much of this research possible. Special thanks go to Dr. Randall J. Bayer for his unyielding faith, encouragement and friendship. I must finally express my love and thanks to my parents, whose support has made this dissertation possible. VITA
September 5, 1952 Bom - Tulsa, Oklahoma. 1974 B.S., Oklahoma State University, Stillwater, Oklahoma. 1 9 7 4 -1 9 7 6 Graduate Teaching Assistant, Oklahoma State University. 1975 Graduate Teaching Assistant, University of Oklahoma, Norman, Oklahoma. 1976 M.S., Oklahoma State University, Stillwater, Oklahoma. 1 9 7 6 -1 9 7 7 Graduate Teaching Assistant, Colorado State University, Fort Collins, Colorado. 1 9 7 8 -1 9 8 5 Graduate Teaching Assistant, Instructor, The Ohio State University. 1981 Sigma Xi Resesarch Grant. 1 9 8 3 -1 9 8 7 Instructor, Columbus College of Art and Design, Columbus, Ohio. PUBLICATIONS
Bruner, J. L. 1977, Chromosome reports in IOPB chromosome number reports LVII. Taxon 26(5/6):557-565.
Bruner, J. L., R. J. Tyrl, et al. 1985. A biosystematic study of the relationship of Nama hispidum and Name stevensii (Hydrophyllaceae). The Southwestern Naturalist 29(4):367-375.
Bruner, J. L., Key and descriptions for the genus Sporobolus to be included in Grasses of Oklahoma, (edited by . J. R. Estes and R. J . Tyrl.)
Bruner, J. L., Keys and descriptions for the genus Schizachyrium to be included in The Florida of Florida, (edited by Helen Correll and Dick Wunderlin.)
FIELDS OF STUDY
Major Field: Botany
v TABLE OF CONTENTS
page DEDICATION...... ii ACKNOWLEDGEMENTS...... iii VITA...... iv LIST OF TABLES...... viii LIST OF FIGURES...... ix LIST OF PLATES...... xii INTRODUCTION...... 1 CHAPTER 1. SYSTEMATICS OF THE SCHIZACHYR1UM SCOPARIUM (POACEAE) COMPLEX IN NORTH AMERICA. I. PHENETIC ANALYSES...... 3
Introduction ...... 3 Materials and Methods ...... 4 R esults ...... 6 Discussion ...... 10 S um m ary...... 15 Literature Cited...... 18
2. SYSTEMATICS OF THE SCHIZACHYRIUM SCOPARIUM (POACEAE) COMPLEX IN NORTH AMERICA. II. TAXONOMY...... 46
Introduction ...... 46 Previous Taxonomy ...... 47 Growth Form...... 48 C aiyopses ...... 49 Leaf Anatomy ...... 50
vi page Chromosomes ...... 52 E cology ...... 53 Origin of the Complex ...... 55 T axonom y ...... 59 Literature Cited...... 93 SUMMARY...... 139
APPENDIX
A. Data matrix for all OTUs used in phenetic studies of Schizachyrium ...... 141
LITERATURE CITED ...... 164 LIST OF TABLES
TABLE CHAPTER I page
1. List of 29 characters and states used in phenetic studies of the Schizachyrium scoparium complex ...... 21
2. Partial list of varieties of Schizachyrium scoparium and their diagnostic characters ...... 22
3. List of character correlations for Schizachyrium s c o p a riu m...... 23
CHAPTER II
4. Major taxonomic treatments of members of the Schizachyrium scoparium complex ...... 97
5. Collections that provided chromosome counts ...... 98
v i i i LIST OF FIGURES
FIGURE CHAPTER I page
1. Principal components analysis composed of 60 OTUs for the major varieties of Schizachyrium scoparium...... 25
2. Correlation phenogram (UPGMA) com posed of 60 OTUS of the major varieties of Schizachyrium scoparium ...... 27
3. Discriminant analysis of the major varieties of Schizachyrium scoparium...... 2 9
4. Mean, range and standard deviation for five key varietal characters for Schizachyrium scoparium...... 31
5. Correlation phenogram (UPGMA) composed of 190 OTUs of North American species of Schizachyrium...... 33
6. Principal components analysis composed of 190 OTUs of North American species of Schizachyrium ...... 35
7. Discriminant analysis of North American species of Schizachyrium ...... 37
8. D iscrim inant analysis of: Schizachyrium scoparium, S. stoloniferum and S. rhizomatum...... 39
9. Mean, range and standard deviation of five key characters used to delimit species of Schizachyrium ...... 41
10. Principal components analysis composed of 242 OTUs of both North and South American species of Schizachyrium... 43
i x page 11. Discriminant analysis of North and South American species of Schizachyrium ...... 45
CHAPTER II
12. Distribution of the Schizachyrium scoparium complex in North America ...... 100
13. Different types of growth form in Schizachyrium...... 102
14. Caiyopsis of Schizachyrium scoparium. X 33...... 104
15. Caryopsis of Schizachyrium scoparium. X 75...... 104
16. Cross-section of leaf: typical Schizachyrium species ...... 106
17. Cross-section of leaf: filiform leaves of 106 Schizachyrium gracile and S. cubense...... 106
18. Distribution of the Schizachyrium scoparium complex and the S. condensatum complex in the Western Hemisphere.... 112
19. Distribution of Schizachyrium gracile ...... 114
20. Distribution of Schizachyrium maritimum ...... 114
21. Features of Schizachyrium gracile ...... 116
22. Features of Schizachyrium maritimum ...... 118
23. Features of Schizachyrium maritimum ...... 120
24. Distribution of Schizachyrium niveum...... 122
25. Features of Schizachyrium niveum...... 124
26. Distribution of Schizachyrium scoparium var. divergens 126
27. Distribution of Schizachyrium scoparium var. littordle 126
x page
28. Features of Schizachyrium scoparium var. divergens...... 128
29. Features of Schizachyrium scoparium var. littorale...... 130
30. Distribution of Schizachyrium scoparium var. polycladus 132
31. Distribution of Schizachyrium scoparium var. rhizomatum .. 132
32. Features of Schizachyrium scoparium var. polycladus 134
33. F eatu res of Schizachyrium scoparium var. rhizomatum 136
34. Distribution of Schizachyrium cubense...... 138
x i LIST OF PLATES
I. Chromosomes for the taxa studied: all 2n = 40, all at X450 except for la (X200) ...... 108
II. Chromosomes for the taxa studied; all at X450 ...... 110
x i i INTRODUCTION
Schizachyrium Nees (Andropogoneae) is a genus of grasses well represented in tropical and subtropical regions of the world by more than 70 species. While at one time included as a section within the genus Andropogon, the majority of modern workers maintain it as a separate genus. In North America there are two distinct groups of species in Schizachyrium. One group, the Schizachyrium cirratum-S. sanguineum complex (Hatch, 1975), is composed of a number of very similar species that express a 10-based polyploid series from 2n = 2 to 2n = 100. Another group, which is the subject of this study, consists of nine species that can be found (with one notable exception) primarily in the southeastern United States and the Caribbean. One species of this group, S. scoparium, occurs in a variety of habitats, throughout North America east of the Great Basin, from Canada, south to Mexico. The remaining species are restricted to rather narrow ecological regimes and distributions. Schizachyrium scoparium is widespread and exhibits considerable variability, and this species and others of the group have generated diverse taxonomic appraisals during this century. The number of species in North America has been judged to range from as few as one to as many as eleven (Hitchcock & Chase, 1950, Roberty, 1960, Gould, 1967). Many of the taxa described during this time were based upon few morphological characters.
1 2 Schizachyrium scoparium is a tetraploid of some economic importance, and its origin has been the subject of some speculation. Stebbins (1975) suggested that it might be the mesic allotetraploid derivative of two diploids from opposite ecological extremes: arid and high precipitation habitats. There was a diploid candidate from the Southwest ( Schizachyrium cirratum), and it was proposed that the other diploid might be found among those unstudied species of the southeastern United States. The diversity of taxonomic opinion of this group and the intriguing speculation about the origin of S. scoparium exposed a need for a study that would develop an accurate understanding of the variation within the S. scoparium complex. In the first chapter, various phenetic analyses are discussed in an attempt to characterize the morphological variation that has traditionally been used to make taxonomic decisions. Under closer scrutiny, many of the taxa maintain their taxonomic integrity while some do not. In the second chapter, chromosome numbers, and morphological, ecological and geographic information are brought together to produce a taxonomy. In addition, a new hypothesis is generated concerning the origin of Schizachyrium scoparium in the light of the current knowledge about variation within the S. scoparium complex. CHAPTER I Systematics of theSchizachyrium scoparium (Poaceae) Complex in North America. I. Phenetic Analyses.
Introduction The Schizachyrium scoparium complex is a group of closely related perennial grasses distributed throughout the eastern two- thirds of North America and the West Indies. The majority of the species demonstrate ecological and distributional specificity in Florida or along the Gulf Coast while Schizachyrium scoparium (Michx.) Nash ranges widely east of the Rocky Mountains and can be found in many habitats. Because knowledge about species ranges and morphological variation within this group has been incomplete, limits of taxa have been the subject of considerable disagreement. Hitchcock (1917) and Hitchcock and Chase (1950) recognized eleven species while Gould (1967) acknowledged only seven of Hitchcock's nine continental United States species. Roberty (1960) in an assessment of worldwide Andropogoneae recognized only a single, variable S. scoparium. The morphological variation within S. scoparium, even at its most conservative circumscription, has been responsible for the description of eleven varieties. Such differences in interpretation of species level taxonomy and the proliferation of varieties in S. scoparium have prompted an effort to consider all members of this complex and define distributions, delimit taxa and determine relationships. This paper represents the results of 3 morphological analyses and examines variation within S. scoparium and species limits within the S. scoparium complex. In addition the morphological relationship of this complex to other South American species is discussed. The South American species of Schizachyrium which are morphologically similar to those of the S. scoparium complex remain largely unstudied. Many of the species have been included in a number of floras (Nees, 1829; Hackel. 1885; Herter, 1943) and one revision (Turpe, 1984), but little accurate information exists about the geographical, ecological and morphological variation of the species.
Methods and Materials Representative specimens of all taxa involved in this study were gathered from living collections made in the course of field work from 1980 to 1984 and various herbaria (BH, FLAS, FSU, FTG, GH, NEBC, NELU, NY, PAC, RM, SDU, SMU, US, USF, TAES, TUC). Living material of S. cubense was unavailable and, due to the inadequate material at hand (three poor specimens), was not included in the detailed numerical analyses. Two hundred and thirty three individuals, selected from over 2000 specimens to represent the full range of variation within taxa, were evaluated fcr 29 morphological characters (Table 1). Cluster analyses (UPGMA) and Principal Components Analyses (PCA) were accomplished with the NTSYS program of Rohlf, Kisbaugh and Kirk (1974). The Biomedical Statistical Program (BMDP, Dixon et al„ 1981) provided univariate statistics and a stepwise discriminate analysis. Three-dimensional 5 illustrations of the results of the PCA were obtained with the use of MACSPIN Graphical Data Analysis Software (Donoho, et al, 1985) for the APPLE MACINTOSH personal computer. Four matrices were used to investigate different levels of phenetic relationships: 1) a 233 OTU data matrix of all North and South American species (Appendix A), 2) a subset (191 OTUs) of this matrix of all North American species, 3) a second subset (60 OTUs) of representatives from within the traditional limits of S. scoparium and 4) a third subset (65 OTUs) representing S. stoloniferum, S. rhizom atum and S. scoparium. Sixty specimens were chosen to represent the abundant variation present in Schizachyrium scoparium. Type specimens were included for all but var. divergens (presumed lost with the Berlin herbarium in WW II) while vars. villosissimum and ducis were represented only by their types (due to identification difficulty). These specimens were then subjected to cluster, principal components and discriminant analyses. Latitude and longitude were coded for each specimen in order to determine correlations with the characters and test for agreement with McMillan's (1964, 1965) discovery of clinal trends in Schizachyrium scoparium. For the actual morphological comparisons of taxa in the analysis these two traits were deleted from the computations. Discriminant analysis can be used to determine the taxonomic identity of unidentified specimens or to visualize the amount of overlap of established taxonomic categories. It is used here with the latter intent. While the OTUs (and groupings of OTUs) are referred to by their traditional 6 taxonomic identities, the interpretations of morphological analyses and final taxonomic decisions were not determined by these previously conceived taxonomic identities. Following the analysis of variation of S. scoparium the eight species of this group that occur in North America were then subjected to various numerical analyses. Specimens of S. condensatum were included in the analyses of North American species since it reaches into central Mexico, and material from living collections was available. South and Central American specimens identified as S. gracilipes, S. lactijlorum, and S. plum igerum (from US) along with those of S. condensatum were chosen as the distinctive representatives of the South American taxa and included in the PCA and DA in this study.
Results Schizachyrium scoparium. Examination of the correlation of various characters provides an indication of how some morphological characters vaiy in relation to one another. Significant character correlations (at .01 level) are presented in Table 3. In general, vegetative size characters (1,3,4,5.6) tend to be positively correlated with one another, with spathe length (27) and with pedicelled spikelet length (25). Leaf vestiture characters (7-10) are highly positively correlated with each other. Since they are coded discontinuously it would be misleading to give much weight to correlations of these characters with the majority of the other 7 continuously varying characters. Spikelet number (13) is positively correlated with vegetative size (1,3,6) and raceme length (12) and is negatively correlated with sessile spikelet length (20) and pedicel length (24). Spathe length (14) correlates positively with longer peduncles (28). Rachis length (15) is positively correlated with raceme length (12), spathe length (14) and fertile lemma length (21). Rachis pubescence length (17) is positively correlated with rachis pubescence amount (18) and callus pubescence length (19). The remainder of the inflorescence characters (20-26) tend to correlate positively with one another, while sessile spikelet length (20) and pedicel length (25) are negatively correlated with spikelet number (13). Latitude (30) is negatively correlated with vegetative size (1,3,4,6) as well as raceme length (12), spikelet number (13) and pedicelled spikelet size (25). Leaf vestiture (7-10) tends to be negatively correlated with latitude. Longitude (31) is negatively correlated with fertile lemma awn length (23) and pedicel length (24). Latitude-character correlations demonstrate that vegetative robustness as represented by greater height and leaf size characters is characteristic of plants as one proceeds south. Large racemes (characteristic of southern plants) are covered by longer spathes but are composed of more, smaller reproductive units (sessile spikelets). Pedicelled spikelets are generally larger in southern plants. Most key varietal characters tend to vary independently from one another and from latitude and longitude. Only leaf pubescence is restricted to the south-central Gulf area. 8 In a PCA (Fig. 1) the first three principal components axes account for 42.47 % of the variation. Characters that load Factor 1 are primarily vegetative characters (3-10) while Factor 2 is loaded by inflorescence size characters ( 13,17-20, 23,24). Factor 3 is loaded by both vegetative and inflorescence characters. These results demonstrate that the varieties are not all morphologically discrete. OTUs of variety littorale demonstrate a rather distinct group, while OTU's of vars. virile, divergens and villosissimum group together. The remaining OTU's form a well-mixed, morphological group. C luster Analysis (Fig. 2), w hich is m ost useful for dem onstrating the relationship of individual OTUs rather than overall groupings, reveals two large clusters and demonstrates morphological groupings similar to the PCA. In the upper cluster, OTU's recognizable as var. littorale tend to cluster together, while OTU's primarily of vars. divergens and virile occur in mixed clusters. OTU's of var. littorale do not cluster as to geographical area. This upper, major cluster is largely composed of plants from the central Gulf Coast area. The lower, major cluster, composed of the remaining OTU's, shows little clustering of plants as to taxonomic category, and suggests a lack of varietal integrity . A discrim inant analysis (Fig. 3) produces results sim ilar to those of the PCA and Cluster analysis. Sheath vestiture (8), pedicelled spikelet length (24) and rachis pubescence length (17) (all important varietal key characters) best separate scoparium into at most three groups: littorale, divergens-virile-villosissimum and one that includes the remaining OTU's. 9 North American Species. Once a basic understanding of variation in S. scoparium was obtained, attention was directed to the remaining North American species.' Cluster analysis (Fig. 5) demonstrates the discreteness of S. niveum, S..condensatum, S. gracile and S. sericatum. OTU’s of Schizachyrium maritimum cluster with three S. scoparium OTUs (of var. littorale ,a taxon of habitats similar to those of S. maritimum) but is otherwise a cohesive group. Schizachyrium rhizomatum OTUs tend to cluster with those of both S. scoparium and S. stoloniferum. These latter three species cluster together, indicating much overlap in morphological variation. The resu lts of the PCA (Fig. 6) coincide w ith those of the cluster analysis. The first three factors generated account for 46% of the variation. Eigenvalues for each of these factors are greater than 1 indicating that most characters are not highly correlated. Spikelet size characters and vegetative pubescence load the first factor while no major character groups are obvious for the second and third. In the top diagram in Figure 6, OTUs of S. maritimum, S. gracile, S. sericatum and S. condensatum cluster distinctly and are well separated from the other taxa by the first two factors. In the bottom diagram in Figure 6, the third factor effectively defines S. niveum The remaining unresolved taxa are : S. scoparium, S. stoloniferum and S. rhizomatum. The first two canonical variables produced by the discriminant analysis are shown in Figure 7. Inflorescence and spikelet characters (21,20,18,17,26 in order of importance to loading) best separate taxa. Schizachyrium maritimum, S. niveum, S. condensatum, S. 1 0 gracile and S. sericatum are discrete while S. scoparium, S. stoloniferum, and S. rhizomatum are not. A discrim ination (Fig. 8) was performed involving only these three taxa. These groups do have some resolution when emphasis is placed upon other characters (1,4,27.23,24). Related South American Species. A PCA produces three factors that account for only 45% of the variation. The eigenvalues for these factors are all much greater than 1 attesting to the lack of great character correlation. The vector diagram (Fig. 10) produces two major results First, three separate point clouds are evident: OTUs of S. maritimum, the North American species and the South American species (including S. condensatum) cluster discretely. Second, OTUs classified as S. plum igerum cluster with those of S. niveum. The discrim inant analysis (Fig. 11) resu lts agree w ith those of the PCA but emphasize the morphological similarities of S. niveum and the South American species. Schizachyrium niveum clusters with the South American species when spikelet characters (17, 19, 21, 22, 29) are employed to discriminate among taxa.
DISCUSSION A number of varietal taxa have been named within the traditional limits of Schizachyrium scoparium (Table 2) based upon various combinations of such characters as inflorescence pubescence, size of pedicelled spikelet and vegetative pubescence. These varieties have seldom been of practical use and assigning a varietal name to a specimen has always been an onerous task. Morphological analyses 11 help to reveal the reasons for such difficulty and point to a better understanding of the variation within this species. One can understand the difficulty in determining an appropriate varietal name for given specimens. Named varieties are based upon combinations of a few, key characters that Eire not mutually exclusive. For example, sessile spikelet size may be used for recognizing one variety while rachis pubescence is emphasized for recognizing another variety. These key characters represent only single states of a character that varies continuously from one taxon to another. Consideration of some simple statistics for four of these taxonomically important characters clearly demonstrates the degree of overlap among taxa (Fig. 4). These four are representative of the vast majority of the quantitative characters. In none of these characters is one variety completely discontinuous from another. Evaluation of multiple morphological characters along with consideration of geographical or ecological criteria suggests that there are three distinct groups of plants within the traditional limits of the variation of S. scoparium. There is one group plants (traditionally identified as var. littorale) that are restricted to the sand dunes of certain coastal areas. This consists of populations from three quite disjunct areas - southern Texas, the north Atlantic Coast and Lake Michigan. Some ecotypic variation was apparent in var. littorale during growth in the greenhouse (southern plants were more robust). A second small group is composed of plants from the south-central Gulf Coast area while the majority of the remaining plants of North America comprise a third group. One can find 1 2 distinctive plants of S. scoparium throughout North America but they are likely to be a part of a large repetitive morphological mosaic that stretches across North America. McMillan's (1959, 1965) ecological studies of several North American grasses described the clinal nature of the variation of culm height, pubescence and phenology. Flowering times are delayed in southern populations and plants have extra time for increased vegetative growth. Culms and leaves tend to be larger. The results of this study indicate that while the inflorescence of southern plants may be longer than those of northern populations, the individual spikelets (and presumably the caryopses) are smaller. Morphological and ecological variability in S. scoparium is comparable to that found in Andropogon virginicus, a related species that occurs in similar habitats across the United States (Chapman & Jones, 1975; Campbell, 1983). These results maintain the morphological integrity of some N orth A m erican species (S. maritimum, S. niveum) and detect overlaps in the variation of S. gracile with S. sericatum, and S. scoparium with both S. stoloniferum and S, rhizomatum. Again, as with the varietal taxa discussed earlier, some simple statistics alone do much to reveal the nature of the considerable variation present in many of the species. Four characters are presented (Fig. 9) with their means, standard deviations and ranges for all of the species. These four characters accurately reflect the situation for all twenty of the quantitative characters. Schizachyrium gracile and S. sericatum are very similar. It appears that S. sericatum represents a slightly more 1 3 robust population of S. gracile. Similar data also help to explain the overlaps in phenetic variation of S. scoparium, S. stoloniferum and S. rhizomatum. Schizachyrium rhizomatum is generally less variable than the other two species but, for most characters variation is highly coincident. The remaining qualitative characters that often typify certain taxa (i.e.the rhizomes of S. stoloniferum and S. rhizomatum) do not weigh heavily in the clustering and PCA and those that demonstrate zero variance within groups do not enter into the discriminant analyses. The only substantial difference between these three traditionally recognized species is the presence or absence of rhizomes and it is questionable if such a trait is important enough to delimit species. S. stoloniferum and S. rhizom atum are ecologically distinctive from S. scoparium but not morphologically so . Schizachyrium condensatum belongs to a group of species that are closely related to the S. scoparium complex. Turpe (1984) maintains that most of the South American taxa must be recognized as varieties of S. condensatum based upon current information. In his revision, Turpe describes Schizachyrium condensatum as a polymorphic species that includes a minimum of 15 previously named varieties or species. These variants differ most noticeably in their habit of branching, with some being extremely fastigiate. This species very much resembles S. scoparium morphologically and it, too, occurs in many different habitats, across Central and South America. Turpe carried out a PCA analysis involving five characters (awn length, glume length, spathe length, leaf length and leaf width). He found low character correlation and no distinct groupings within a 1 4 large S. condensatum and concluded that these characters cannot be used to subdivide this species. Information on morphology confirms, for the most part, the discreteness of the North American taxa from other, related species of Central and South America. There are some morphological groupings apparent among the OTUs that represent S. condensatum when a large number of characters are considered. Chromosome numbers suggest that S. condensatum is composed of two chromosomal races. Diploid (2n = 20)populations are reported from Brazil and Uruguay (Gould, 1956) and El Salvador (Davidse & Pohl, 1974) while all of the Mexican populations of this species sampled by this author proved to be tetraploid. There are no reports of tetraploids in South America. This suggests that our understanding of variation in S. condensatum would benefit from further study. The great morphological diversity of S. condensatum and related taxa in South America together with reports of diploids suggest that the origin of S. condensatum might be found in southern South America. But wherever the origin, over time tetraploids must have moved northward into southern Mexico. One of the original intents of this research was to locate in the southeastern United States the diploid ancestors of the tetraploid S. scoparium however, no diploid relatives were discovered. The results of this study raise the possibility that S. condensatum might be the progenitor of S. scoparium. This is a distinct possibility since S. scoparium is also a tetraploid. While the phenetic analyses are used to find distinct patterns or clusters of morphological variation S. scoparium is very 1 5 similar morphologically to the S. condensatum that is found in Mexico. The morphological similarity of S. niveum to the South American species raises additional questions about its origins. Characters of this plant (branching habit, spikelet characters and overall coloration) suggest a closer relationship to S. condensatum than to any North American taxa. Is it a result of long distance dispersal from South America( it particularly resembles three Colombian collections of S. condensatum from US) or Central America? Or is S. niveum a relict that remained behind when the glacial periods restricted a once more widely distributed S. condensatum?
Summary The phenetic results provide a clearer picture of the morphological variation of the Schizachyrium scoparium complex in North America. They suggest that S. maritimum, S. niveum, S. gracile and S. scoparium are morphologically quite distinct and can be recognized at the species level. Schizachyrium scoparium proves to be an extremely variable taxon that includes the following recognizable morphological varieties: var. divergens, var. littorale, var. stoloniferum and var. rhizomatum. The characters typically employed to identify species and varieties have been demonstrated to be unreliable. This should come as little surprise to those who work with various grass taxa. Whether the genus is Poa (Kellogg, 1985), Andropogon (Campbell, 1983) or 1 6 Schizachyrium, traditional use of one or a few characters for taxonomic delimitations has been shown to be unreliable under the scrutiny of more comprehensive morphological analyses. Grass taxa that have a wide geographic range are likely to develop distinctive ecological and morphological variants but individual characters are just as likely to vary independently from one another. Official recognition of distinctive character states or combinations of character states without an awareness of the total morphologic variation proves invariably to be an unwise taxonomic strategy. Upon completion of an examination of morphological variation wathin the Schizachyrium scoparium complex, interest naturally turns to the original sources of variation and how the variation is maintained. The patterns of variation most likely have resulted from factors that include the interaction of pre-glacial variation and the control of flowering by climate. McMillan proposed the existence of three centers of variation in pre-glacial S. scoparium : a western- montane area, a southern area (Texas, Oklahoma and Louisiana) and the southeastern United States. McMillan's idea is consistent wath the results of this investigation and supported by his evaluation of flowering behavior (confirmed by my greenhouse observations). The northern populations are characterized by both earlier flowering and minimal growth. Similar trends exist in S. scoparium from higher to lower latitudes. Today, northern parts of North America are populated by genotypes that probably originated in the northern- montane areas, the eastern part has been populated by genotypes from the southeastern United States and populations in the central 1 7 part of the continent are derived from mixtures of genotypes that arose from all three original areas. While outcrossing and wind pollinated, flowering times restrict gene flow and allow for the localization of many genotypes and phenotypes across North America. Comparison of the S. scoparium complex with what may be called the S. condensatum complex introduces many interesting questions. What are the origins of S. scoparium and S. niveum, and how are the widely separated yet very similar species of North and South America related to one another? The South American species appear to have diversified by increasing the degree of branching, a trend also recognizable in the Andropogon virginicus complex (Campbell, 1983). Is there any significance to this observation? Any accurate investigation into these questions must be based upon detailed study of the morphology, ecology and cytology of the South American species. 1 8
LITERATURE CITED
Allred, K. W. 1984, Studies in the genus A ristida (Gramineae) of the southeastern United States. I. Spikelet variation in A. purpurescens, A. tenuispica and A. virgata. Rhodora 86:73-77. Campbell, C. S. 1983. Systematics of the Andropogon virginicus complex (Gramineae) Joum. Arnold Arbor. 64:171-254. Chapman, R. H. and S. B. Jones. 1985. Ecotypic differentiation of Andropogon virginicus (Gramineae). Bull. Torrey Bot. Club 102(4):166-171. Church, G. L. 1929. Meiotic phenomena in certain Gramineae. II. Paniceae and Andropogoneae. Bot. Gaz. 87: 608-629. Dixon, W. J . £t ah 1981. Biomedical Data Program. 700 pp. University, of California Press, Berkeley. Donoho, A. W., D. L. Donoho and M. Gasko. 1985. Macspin graphical data analysis software. D2 Software, Inc. Austin, Texas. Gould, F. W. 1967. The grass genus Andropogon in the United States Brittonia 17(l):70-76. Hackel, E. 1885. Andropogoneae Novae. Flora 7:115-128. Herter, J. 1943 Plantae Uruguayenses novae vel criticae. Revista Sudamer. Bot. 7:(6/8): 193-194. Hitchcock, A. S. 1909. Grasses of Cuba. Contr. U.S. Natl. Herb.
12(6 ): 183-258. 1 9 ______. 1936. Manual of the grasses of the West Indies USDA. Misc. Publ. 243. 385-402 pp. ______1950. Manual of the Grasses of the United States (Agnes Chase, rev. ed.). Dover Publications, Inc. New York. 1051 pp. Kellogg, E. A. 1985. A biosystematic Study of the Poa secunda complex. Journ. Arnold Arbor. 66:201-242. McMillan, C. 1959. The role of ecotypic variation in the distribution of the central grassland of North America. Ecol. Monogr. 29:285-308. ______. 1964. Ecotypic differentiation within four North American prairie grasses. I. Morphological variation within transplanted community fractions. Amer. J. Bot. 51:1119- 1128. ______. 1965. Ecotypic differentiation within four North American prairie grasses. II. Behavioral variation within transplanted community fractions. Amer. J. Bot. 52:55-65. Metcalfe, C. R. 1960. Anatomy of the Monocotyledons. I. Gramineae. Oxford, Clarendon Press. Nees von Esenbeck, C. G. 1829. "Agrostologia Brasiliensis " in M artius' Flora Brasiliensis. ( 2:208) 1829. Ostergren, G. and W. Heneen. 1962. A squash technique for chromosome morphological studies. Hereditas 48:332-341. Riggins, R. 1977. A biosystematic study of the Sporobolus asper complex (Gramineae). Iowa State J. of Res.. 51(3):287-321. 20 Roberty, G. 1970. Monographique systematique des Andropogoneae du globe. Boissiera 9:216-239. Rohlf, F. J., J. Kishpaugh and D. Kirk. 1971. NT-SYS. Numerical taxonomy system of multivariate statistical programs. Tech. Rep. St. Univ. of New York at Stony Brook. New York. Scribner, F. L. and C. R. Ball. 1898. Studies on American Grasses III. Miscellaneous notes and descriptions of new species. U.S.D.A. Bull. 24: 40-41. Turpe, A. M. 1974. Revision of the South American species of Schizachyrium (Gramineae). Kew Bull. 39:169-178. 21 Table 1. List of 29 characters and states used in the phenetic studies of the Schizachyrium scoparium complex. Discontinuous character states coded are: *- glabrous (0)/ pubescent (1)/ glaucous (2); enclosed/half exerted/ totally exerted. Latitude (30) and longitude (31) were included in initial analyses to provide geographical correlations but were deleted during morphological analyses.
1. Height of Flowering Culms 17. Rachis Pubescence Length
2. Rhizomes (0/1) 18. Rachis Pubescence % ofRachis
3. Leaf Blade Length 19. Callus Pubescence Length
4. Leaf Blade Width 20. Sessile Spikelet Length
5. Leaf Sheath Length 21. Fertile Lemma Length
6. Ligule Length 22. Relative Length of Lemma Cleft
7. Blade Surface Covering (0/1/2)* 23. Fertile Lemma Awn Length
8. Sheath Surface Covering (0/1/2) * 24. Pedicel Length
9. Throat Pubescence (0/1) * 25. Pedicelled Spikelet Length
10. Auricle Pubescence (0/1) * 26. Pedicelled Spikelet Awn Length
11. Raceme Exertion (0/1/2) ** 27. Spathe Width
12. Raceme Length 28. Peduncle Length
13. Spikelet Number 29. Callus Length
14. Spathe Length 30. Latitude
15. Length ofRachis 31. Longitude 22 Table 2. Characters used to delimit common varieties of Schizachyrium scoparium.
VARIETY DIAGNOSTIC CHARACTERS
divergens (Hack.) Gould larger spikelets, fewer spikelets per inflorescence, less inflorescence pubescence ducis (Fern.) Gould more branched, rames not very flexuous, culms not very compressed
freq u en s(Hubb.) Gould smaller sessile spikelet, vestigial pedicelled spikelet, little rachis pubescence
littorale (Nash) Gould large spikelets, glaucous foliage, compressed culm
neomexicanum (Nash) profuse rachis pubescence, Gould straightened raceme
septentrionalis (Fem.& plants less robust, more rachis Grisc.) Gould pubescence, fewer spikelets
villossissimum (Kearn.) sheath much pubescent Gould
virile (Shinners) Gould larger pedicelled spikelets, less inflorescence pubescence 23 Table 3. Character correlations for Schizachyrium scoparium. Underlined characters express a negative correlation.
CHARACTER SIGNIFICANT CORRELATIONS
1. Height 3,4,12, 13, 25, 30 2. Rhizomes 3. Blade Length 1.5.6.7.8.25.27.30 4. Blade Width 1.5.6.7.8.9.10.27.30 5. Sheath Length 3.4.5.8.20.25.27 6. Ligule Length 3.4.5.7.8.9.10.13.30 7. Blade Pubescence 3.4.6.8.9.10.30 8. Sheath Pubescence 3.4.5.6.7.9.10.30 9. Throat Pubescence 4.6.7.8.10.30 10. Auricle Pubescence 4.6.7.8.9.30 11. Raceme Exertion 12. Raceme Length 1.13.14.15.28.30 13. Spikelet N um ber 1.3.6.12.20.24.30 14. S pathe Length 12,15,21,28 15. R achis Length 12,14,21 16. R achis Tip W idth 17. Rachis Pub. Length 18.19.23.26 18. Rach. Pub. % ofRachis 17.22 19. Callus Pub. Length 17.26 20. Sess. Spklt. Length 5.13.21.23.24.25.27 21. Fertile Lemma Length 14.15.20.25 22. Lemma Cleft Length 18.23 23. Fert. Lem. Awn Length 17.20.22.24.26.31 24. Pedicel Length 1 3 .2 3 .3 1 25. Ped. Spklt. Length I.3.5.20.21.30 26. Ped. Spklt. Awn Length 17.18.19.23 27. Spathe Width 3 ,4 ,5 ,8 ,2 0 28. Peduncle Length II.1 2 ,1 4 . 29. Callus Length 30. Latitude 1.3.4.6.7.8.9.10.12.13.25 31. Longitude ______2 3 .2 4 ______F ig u re 1. PCA composed of 60 OTUs representing the spectrum of variation in Schizachyrium scoparium. Basic group outlines indicated by solid lines. TyPe specimens indicated by underlined symbols. -fc divergens, ^ ducis, • frequens, <>- littorale, □ neomexicanum, 0 septentrionalis, ☆ villosissimum, # virile, ★ type specimen of S. scoparium.
24 1 25
•<> 0 I—
ririle
divergent
o — F ig u re 2. Correlation phenogram (UPGMA) composed of 6 0 OTUs of Schizachyrium scoparium. Traditional variety designations are indicated by the first three letters of the varietal epithet.
26 2 27
LIT I VIR 2 DIV 2 DIV J VI* 3 DIV I DIV 3 VII.LTYP DIV 3 DIV 6 VI* 7 SEP 10 LIT 3 LIT 5 LIT 6 LIT 9 VI* I VIR 9 VIR 6 VIR 11 VI* s FRE I LIT 2 LIT 3 LIT 8 VI* 3 VIR IO VI* 3 FRE 3 * FRE 6 I FRE 9 SEPTYP FRE IO i SCOTYP NEO 7 FRE 3 SEP 9 r*E7 FRE 8 NEO 3 LIT 7 FRE 9 LIT IO SEP 7 DUCTYP NEO I ■ NEO 5 FRE 2 SEP 2 •SEP 6 SEP 8 NEO 2 NEO IO • NEO 8 SEP 3 NEO 9 ■SEP 3 ■ NEO 6 -NEO 3 -SEP 3 T ~ ~T~ i 0.2 0.5 0.9 correlation Figure 3. Results of a discriminant analysis of Schizachyrium scoparium varieties. Type specimens are represented by underlined symbols, 'b ducis, -fc divergens, • frequens, <> littorale, □ neomexicanum, O septentrionalis, ☆ villosissimum, virile, ★ type specimen of S. scopariurrL
28 virile
divergens
□□ littorale F ig u re 4. Mean, range and standard deviation for five key varietal characters used to delimit five major varieties of Schizachyrium scoparium. Taxa are indicated by the first three letters of the varietal epith ets.
30 31
SPIKELET HEIGHT (cm) NUMBER
DIV VIR
LIT L I J I I 30 65 100 135 170 200 6 7 8 9 10 11 12 13
RACHIS PUBE5CENCE 5ESS5ILE SPIKELET LENGTH (mm) LENGTH (mm)
SEP
NEO FRO
DIV VIR
LIT J l _ l I______I___ L 0 1 2 6 7 8 10 11 F ig u re 5. Correlation phenogram (UPGMA) composed of 190 OTUs of the North American species of Schizachyrium. Taxa indicated by the first three letters of their specific epithets. Cophenetic correlation coefficient is 0.774.
32 33
CORRELATION -j -.j j .1 .» j .» u
SCO
Sto
-.1 -.1 .1 .1 .9 .1 F ig u re 6. PCA com posed of 190 OTUs of North American species of Schizachyrium. 4 S. condensatum, 4 S. gracile, 0 S. maritimum, * S. niveum, • S. rhizomatum, S. scoparium, 'tx S. sericatum, + S. stoloniferum.
34 35
gracile v v > v>» *
sericatum
/n iv eu m* 1 v *
oooooo * o — >0
marilimum ♦♦ ♦ oo ♦ ♦ conuensatum
> ♦ i V#i ♦♦ ♦ ♦ ♦ • v *♦♦• ♦ ♦ ♦ ♦ OOO * »0 4♦000*0 • *ooo o ooooo ♦♦
condensatum F ig u re 7. Results of a discriminant analysis of the North American species of Schizachyrium. ★ S. condensation, • S. gracile, + S. maritimum, <> S. niveum, □ S. rhizomalum, S. scoparium, O S. sericatum, S. stoloniferum.
36 7
<► sco par iu m / rh izo matu m/slo lo a ife ru m ❖ * # o * * % * V J* * & # * * *%
t o * □rf?a * * v > *
* * * * * * * *
maritimum condensaUim
O sericatum # o o o 2 gracile l Figure 8. Results of a discriminant analysis of, Schizachyrium scoparium, ■#- S. stoloniferum and □ S. rhizomatum.
38 rhizoouUum
stolonifenim
# *
scoparium Figure 9. Mean, range and standard deviation of five key characters used in the delimitation of species of Schizachyrium. Taxa are indicated by the first three letters of their specific epithets.
40 41
HEIGHT (cm BLADEWIDTH (mm)
SCO-
—H
200 1.0 5.0 11.0
SESSILE SPIKELET PEDI CELLED LENGTH (mm) SPIKELET LENGTH (mm) 5C0
5T0
CON
MAR
NIV
RHI
GRA
5ER
4 0 7.5 11.0 0.0 5.0 10.0 Figure 10. PCA composed of 233 OTXJs of both North and South American species of Schizachyrium. O North American species (except S. maritimum), X S. maritimurru + South American species (including S. condensatum), □ S. niveum. A two dimensional plot of a three-dimensional representation from M acSpin.
42 43 10
•o
• ©
o ■ ■
South American Taxa
u> Factor 1 Figure 11. Results of a discriminate analysis of the North and South American species of Schizachyrium. North American species, • South American species (including S. condensatum). S.. niveum OTU's are enclosed in a circle.
44 i 4
£
* * v <* #<* * * * V * <* # # # #
Complex in North America,n . Taxonomy.
INTRODUCTION
The Schizachyrium scoparium complex is a group of nine closely related species that occur in open habitats throughout much of North America, from Canada to Mexico and the West Indies. The species demonstrate numerous morphological and ecological differences. Schizachyrium scoparium (Michx.) Nash, the most widespread and variable species, ranges east of the intermontane regions of the Rocky Mountains. The remaining taxa are restricted to Atlantic or Gulf coastal areas of the southeastern United States [Schizachyrium maritimum (Chapm.) Nash, Schizachyrium sericatum (Swallen) Gould, Schizachyrium littorale[Nash) Nash, Schizachyrium stoloniferum Nash, Schizachyrium rhizomatum (Swallen) Gould, and Schizachyrium niveum (Swallen) Gould], Cuba [Schizachyrium cubense (Hack.) Nash ] or the West Indies [Schizachyrium gracile (Spreng.) Gould)]. Prior to this study, investigations of members of this complex had focused primarily on the physiological and morphological variation of S. scoparium from an ecological point of view (Cornelius (1947), Larsen(1947), McMillan, (1959, 1964, 1965)]. Little was
46 47 known of the type or extent of morphological, cytological and ecological differences in this group and the taxonomy of these species had been confused because of their extreme variability. An evaluation of morphology (including character phenetics, rhizome production, leaf anatomy and caryopsis surface morphology), chromosome number and ecological information has been used to produce a taxonomy for this group.
PREVIOUS TAXONOMY
Various taxonomic treatments demonstrate some disagreement in delimiting species (Table 4). Hitchcock(1936, 1950), Gould(1967), and Roberty(1960) do not agree on a classification. Hitchcock recognized 11 species (as Andropogon) in this complex. He differentiated them on the basis of spikelet size, pubescence, possession of rhizomes and habitat preference. Gould believed that similarities required the concept of a more variable Schizachyrium scoparium and included two of Hitchcock’s species in that taxon. In his treatment of the Andropogoneae, Roberty reduced all taxa to varietal status under S. scoparium. His work did not include three recognized North American taxa: S. littorale, S. niveum and S. sericatum . Clayton (1964) has suggested that S. gracile, and S. cubense are not closely related to this group and belong to the genus Andropogon. These different treatments indicated a need for work which would characterize variation within the complex and establish taxonomic limits. The results of previous phenetic analyses (Chapter 48 I) suggest that S. maritimum, S. niveum, S. gracile/S. sericatum and S. scoparium form morphologically distinct groups, while S. scoparium proves to be a variable species composed of four distinguishable, morphological types.
GROWTH FORM
The character of rhizome production has been used to delimit taxa in this group but conflicting descriptions left it unclear as to which taxa actually produce rhizomes. Various accounts have identified Schizachyrium rhizomatum, S. stoloniferum, S. maritimum, S. littorale and S. cubense as having rhizomes. Gould(1967) believed that rhizome presence in S. littorale was of less importance than the inflorescence characters. At the same time he maintained the specific status of S. stoloniferum and S. rhizom atum based upon their rhizomatous growth in spite of close inflorescence similarity w ith S. scoparium. Herbarium sheets proved equivocal and only an examination of living plants reveals which taxa are truly rhizomatous. Schizachyrium niveum, S. scoparium, S. gracile and S. sericatum are caespitose (Figure 13). In this growth form, innovations produce an outward and slightly upward direction of growth each year while the deposition of litter and subsequent soil formation presumably keep the crown of each plant at, or just above, soil level. Observation of S. maritimum and S. littorale in the field and greenhouse demonstrated that these taxa do not produce true rhizom es. Schizachyrium maritimum is decumbent, rooting 49 infrequently at the nodes. Its stems are often buried by sand. Schizachyrium littorale is primarily caespitose, producing basal internodes that grow faster and become more elongated than those of the typical caespitose forms. These elongated internodes apparently help the plant cope with the near constant deposition of sand and prevent complete burial. The buried leaves of both taxa decay over time and stems superficially come to resemble rhizomes. Schizachyrium rhizomatum and S. stoloniferum produce true rhizomes (Figure 13). The rhizomes of S. stoloniferum are thin but n o t hindered in their growth by the sandy soils of Florida and Southern Alabama and Georgia. Intemodes tend to be long and not completely covered by the scale-leaves. Rhizome production becomes more aggressive in populations that occur in the southern part of its range and the plants often produce vigorous stands along roadsides. Schizachyrium rhizomatum possesses much shorter, tougher rhizomes and the intemodes are well protected by scale leaves. This taxon inhabits the shallow, marl soils of southern Florida and its rhizomes enable it to grow over and conform to the surface of this rock. Rhizome production is common in grasses of porous, sandy soils, such as Andropogon hallii, Sporobolus asper var. rnacrus and Sporobolus clandestinus (Riggins, 1977).
CARYOPSES
Since caryopses were known to vaiy somewhat in size in this group, scanning electron microscopy was used to examine the 50 surfaces of caryopses for differences in externa] morphology. Results (Figures 14, 15) suggest that there are no significant differences among the taxa under study. Specimens were mounted on double stick tape on 1 inch aluminum mounts, gold-coated and examined on a Jeol JSM-35 scanning electron microscope at lOKv at magnifications of X22 and X60. Photos were taken with use of a 55 P/N polaroid filter.
LEAF ANATOMY
In a survey of the leaf epidermises of species of Andropogon that included Schizachyrium gracile and other species of Schizachyrium, Hilu (1984) found variation but no patterns of taxonomic value.. There are two distinctive types of leaf shapes characteristic of the species in this complex: the leaves of S, gracile, S. cubense, and S. sericatum are terete while the remaining species possess typical, flattened blades. This suggested the possibility of anatomical characters that might be useful in determining relationships. While a terete versus flattened blade presents a striking morphological difference, their leaf anatomy do not differ radically . All blades have a typical Kranz anatomy cross-section, as described by Metcalfe (1960) for the Andropogoneae, with little variation (Figure 16). One large median vascular bundle is accompanied by smaller vascular bundles to either side. Chlorenchyma is radiate and the bundle sheaths are, for the most part, complete. Bulliform cells occur in the epidermis in irregular groups on the adaxial surface of 51 the blade. The basic organization of the leaf is the same for Schizachyrium sericatum, S. gracile and S. cubense (Figure 17) but the cross-section is terete. The upper epidermis appears to have proliferated and fused to become the center of the leaf while the rest of the blade appears curled around this central tissue. The adaxial surface is reduced to a small groove that is quite prominent at the base of the leaf, becoming less so toward the apex. The adaptive function of the gracile-type of leaf blade is unknown. Burduja & Toma (1971) report that the distribution of sclerenchyma associated with the vascular bundles can be correlated with ecological factors. Grasses from arid areas are characterized by well-developed sclerenchyma tissue while tropical grasses have a high proportion of smaller bundles not accompanied by sclerenchyma. This appears to be the case here. The species with terete leaves are the tropical ones and they do demonstrate a reduction in sclerenchyma development. The presence of I-beam vein construction (vascular bundles with groups of sclerenchyma cells extending to both upper and lower epidermises) is evident in all of the species studied. This type of arrangement of veins and sclerenchyma is reported to enable the strap-shaped leaves of the grasses to better withstand the physical stresses they experience (Ellis. 1976). That the terete leaves also possess the I-beam construction could indicate a recent origin of these types of leaves in the species. The reduced surface-to-volume ratio of the terete leaves might help reduce water loss for species that occur in open areas of high light intensity and heat. Examination of the leaf anatomy 52 entailed looking at a cross-section of the leaves. The cross-sections were obtained by placing leaves between two halves of a cork and shaving the end of the cork with a razor blade. The resultant cross-sections were crudely acquired but sufficiently thin and intact to view competently with the aid of a Zeiss compound microscope.
CHROMOSOMES
Original speculation on the origin of Schizachyrium scoparium (Stebbins, 1975) sought an ancestral diploid among the unstudied taxa of this complex in the southeastern United States. A polyploid series (of 2n = 20, 40, 60, 100), reported for the closely related S. cirratum - S. sanguineum complex (Hatch 1975), suggested the possibility of different chromosome numbers in the S. scoparium complex. But as in the Andropogon virginicus complex, whose members have distributions similar to the taxa of this complex (Campbell, 1983), these taxa reveal no variation in chromosome number . Examination of chromosome numbers, did not reveal the hoped-for diploids. Counts of 2n = 40 were made for all of the species in the complex (Table 5). First reports were obtained for S. sericatum, S. stoloniferum, S. rhizomatum and S maritimum. C ounts for the remaining taxa are confirmations of previously published reports. The counts for the taxa studied here are shown in Plates I and II. Schizachyrium condensatum from Central and South America, a species morphologically very similar to S. scoparium is included in these counts. Previous reports for Schizachyrium 53 condensation have been diploid (2n = 20; Pohl & Davidse, 1974. Gould, 1956). Church (1929) reported 2n = 56 for a meiotic count in S. scoparium but this is apparently aberrant. The North American taxa have diversified ecologically and to some extent morphologically, but there has been no diversification in chromosome number. For the cytological studies root tips were collected from the greenhouse and placed in a 15% solution (by weight) of colchicine at 4°C for approximately 8 hours. They were then fixed overnight (8-10 hrs) in Ostergren and Heneen's fluid (Ostergren and Heneen, 1962). Staining involved the Fuelgen reaction (Fuelgin and Ross. 1924) according to the procedures of Ostergren and Heneen (1962). Root tips were hydrolyzed in 1 N HC1 at 60°C for 8 minutes and washed briefly in distilled water. They were then transferred to Schiffs reagent (leuco-basic-fuschin) for 2 hours and finally a 5% pectinase solution for 1 hour. Roots were stored at -20°C in 45% acetic acid. Root tips were macerated in 45% acetic acid and pressure was gently, but firmly, applied to the coverslip prior to observation.
ECOLOGY
The majority of the taxa prove to be geographically and ecologically isolated from one another. Schizachyrium scoparium occurs in a wide range of habitats across North America (Figure 12). Most of the taxa traditionally recognized as varieties of S. scoparium are based upon one or few characters and these characters vary clinally across the continent (McMillan, 1965; Bruner 1974). Few characters are correlated with any geographical areas or ecological 54 regimes (Chapter I). Consequently, specimens identified as a single variety may be found in many places in North America and different varieties may be found within a single population (Gould, 1975). Maintenance of the majority of the varieties seems taxonomically inadvisable. Phenetic analyses note two exceptions: pubescent sheathed plants with larger pedicelled spikelets (typically referred to as var. divergens or var. villosissimum of the south-central pinelands of the United States (Figure 27)), and var. littorale, a sand dune inhabitant (with a much pubescent inflorescence) that has a striking disjunction (Figure 27). Schizachyrium scoparium var. littorale is one of a number of species that once occurred in suitable habitats along the Atlantic and Gulf coasts, during the last period of glaciation (Peattie, 1922). Var. littorale followed the margins of the glacier as it receded and today persists on present-day sand dunes in more seasonal climates: those along the Texas coast, the Atlantic Coast and areas around the Great Lakes. An alternative hypothesis, involving long distance dispersal by birds, has been proposed to explain this type of disjunction pattern for some species (Pierce, 1974) but this is cannot be applied to var. littorale. This taxon is neither attractive to birds nor is it likely to be inadvertently carried by birds for long distances. Long distance dispersal is also unlikely against the prevailing westerly winds. The hypothesis of Peattie best accounts for the current distribution of Schizachyrium scoparium var. littorale. With one exception the remaining taxa are geographically and/or ecologically isolated from one another. Schizachyrium 55 scoparium is not sympatric with any of the other taxa. Schizachyrium stoloniferum is the taxon with the widest distribution (Figure 30) following that of S. scoparium. It inhabits the sands of southern Georgia and Alabama, extending into Florida as far south as the Everglades. Schizachyrium rhizomatum occurs only on the oolite or the Everglades Keys and the Florida Keys (Figure 31). Schizachyrium niveum is rare and encountered only in the fragile Ceratiola-Pine-Oak woodlands of central Florida (Figure 24). Most of the suitable habitats for this species have been destroyed and used for orange groves and housing developments. Schizachyrium m aritim um is restricted to the ocean's edge along the Gulf Coast (Figure 20). Schizachyrium gracile can be found on oolite or sand in southern Florida and the West Indies (Figure 19). Schizachyrium cu.bense is known only from Cuba (Figure 34). At one site in Franklin county, Florida, one clone of S. stoloniferum was found growing within 100 feet of a population of S. maritimum but none of the others are sympatric. Schizachyrium niveum is surrounded by S. stoloniferum but is not sympatric with it.
ORIGIN OF THE COMPLEX
The results of this study support a taxonomy for the Schizachyrium scoparium complex based upon morphology and geographical/ecological considerations. On the basis of morphology there are four major groups that may be best recognized at the species level: Schizachyrium niveum, S. maritimum, S. gracile, and S. scoparium. The remaining taxa examined are all considered as a part 56 of the variation of S. scoparium. W ithin S. scoparium, the phenetic analyses support retention of variety divergens, a pubescent taxon of the south-central United States pinelands and variety littorale, a taxon with very pubescent racemes that inhabits coastal sand dunes. Schizachyrium rhizomatum and S. stoloniferum, traditionally recognized as species, are virtually indistinguishable (morphologically) from typical S. scoparium but because they are rhizomatous, ecologically restricted to specific habitats in the southeastern United States and spatially isolated from the rest of S. scoparium it. seems best to recognize them as varieties of S. scoparium. An original hypothesis concerning the origin of Schizachyrium scoparium provided much of the impetus for this study. Stebbins (1975) speculated that this species might be the allotetrapoid derivative of two diploid species: one from the wet, southeastern United States and one from the arid southwest. Such a hybrid (ecologically as well as morphologically intermediate) would have been well suited to colonize the large mesic expanse of most of North America that has existed during and since the onset of glaciation on the continent. At the time this hypothesis was put forth, diploid Schizachyrium cirratum was known from the Southwest and the most likely prospects for diploid progenitors in the southeast were to be found among the unstudied taxa of the southeastern United States. The results of this study find no related diploids in the southeast and dictate proposal of a new hypothesis. Current evidence suggests that the S. scoparium complex may have originated in 57 Central America and dispersed into North America from the south. A species closely related to the Schizachyrium scoparium complex occurs widely throughout South and Central America. Schizachyrium condensatum extends from the grasslands of Brazil and Argentina, along the Andes into northern South America and continues across Central America into central Mexico (Figure 18). This species appears to be at least as morphologically and ecologically diverse as S. scoparium. Schizachyrium condensatum is conspicuously more branched than S. scoparium but otherwise is very similar morphologically. The greater amount of morphological diversity of S. condensatum is in South America. Schizachyrium condensatum. unlike S, scoparium, is composed of both diploids and tetraploids. Diploid populations have been reported from Brazil and Uruguay (Gould 1956) and El Salvador (Davidse & Pohl, 1974), while several Mexican populations studied by this author (see Table 5) were tetraploid. The great morphological diversity and the presence of diploids in southern South America suggest that the origins of S. condensatum are to be found there. Dispersal northward into Central America would have been possible only since the Pleistocene (Raven & Axlerod, 1975), a time during which areas in the tropics became more arid and the grassland and savannah habitats that S. condensatum prefers became more extensive. Any earlier than the Pleistocene and Central America would have found a Central America that was too tropical with few, if any, arid areas suitable for colonization by S. condensatum. Allred (1983) has proposed a similar scenario for the dispersal of New World 58 Bothriochloa from South America into North America. The wind- borne propagules of S. condensatum, identical to those of Bothriochloa, could easily account for such dispersal. Once the more arid areas became available, diploid S. condensatum could have dispersed into Central America. One cam hypothesize either an autoploid or alloploid origin for the tetraploid S. condensatum If an alloploid it would have resulted from contact with another diploid relative. Schizachyrium cirratum ranges throughout the Western United States and Central America (Hatch, 1975) and can still be considered a possible diploid candidate. The resultant tetraploid spread into North America before or during the interglacial periods. Subsequent glacial advances compressed the habitat zones in North America, shifting plant distributions southward. Mexican populations would have become isolated from populations along the American Gulf Coast and Florida during the periods of glacial extension. Such isolation would result in the morphological distinctions that cause us to recognize the Mexican S. condensatum as distinct from the members of the S. scoparium complex. Subsequent expansion of S. scoparium from continental margins after the last glaciation (McMillan, 1965) would have produced the current distribution of morphologically similar taxa in this group of species. 59
Taxonomy
Schizachyrium Necs Annual or perennial grasses, typically caespitose but some species rhizomatous; culms round or strongly flattened, producing leaves with flattened, folded blades (rarely terete); flowering culms with few to many branches. Inflorescence a single raceme, straight and erect or strongly flexuous and divergent at maturity; rachis flattened and clavate, the apex cup-shaped and typically erose or lobed. Disarticulation below sessile spikelets with paired spikelets falling as a unit; sessile spikelet fertile, two-flowered, lower floret vestigial; glumes large; lemmas thin, hyaline; lemma of fertile floret bifid (sometimes extremely so) and awned; pedicelled spikelet awnless or short-awned, in some species vestigial and sterile, consisting of one or two small glumes, in others nearly as large and developed as the sessile spikelet and occasionally staminate. A genus of an estimated 70 species; worldwide, primarily tropical and subtropical.
Key to species of the Schizachyrium scoparium complex. 1. Leaf blades terete, no wider than 1 mm. 2. Plants caespitose 1. S. gracile 2. Plants rhizomatous ...... 2. S. cubense 1. Leaf blades flattened, occasionally folded, wider than 1 mm. 60 3. Base of fertile lemma indurate, white; lemma split 3/4 to 7/8 its length; callus pubescence to 1 mm long ...... 3. S. niveum 3. Base of fertile lemma membranous, not indurate, same color as the rest of lemma; lemma split only to 1/2 its length; callus pubescence to 0.5 mm long. 4. Pedicelled spikelet staminate, 4.5-8.5 mm long; rachis densely villous its entire length ......
...... 4. SL maritimum 4. Pedicelled spikelet sterile, 1-4.5 mm long; rachis sparsely villous for up to 2/3 its length ...... 5. S. scoparium 5. Plants rhizomatous. 6. Leaf blades less than or equal to 3 mm wide var.5a. rhizom atum 6. Leaf blades greater than 3 mm wide var. 5b polycladus 5. Plants caespitose, not with true rhizomes. 7. Rachis densely villous one-half to three-quarters its length from the tip, hairs 3.5-6 mm long; leaves glaucous var. 5c littorale 7. Rachis not densely villous its entire length, or if so rachis hairs 1.5-3.5 mm long; leaves usually not glaucous 8. Sheaths mostly pubescent . . 61 ...... var. 5d divergens 8. Sheaths mostly glabrous ...... var. 5e scoparium
1. Schizachyrium gracile (Spreng.) Nash in Small. FI. S.E. U.S. 60. 1903. Andropogon gracilis Spreng. in Linne, Syst. Veget. 1:284. 1825. Type: Hispaniola; type not seen. Andropogon sericatus Swallen, J. Wash. Acad. Sci. 31:355. 1941. Type: Florida: Monroe Co., Ramrod Key: 17 O ctober 1940, W. A. Silveus 6633, 1940 (Holotype, US!). Schizachyrium sericatum (Swallen) Gould, Brittonia 19:73. 1967.
Caespitose perennials; flowering culms slender, wiry, little branched, 30-80 cm long, 1.5-3.5 mm in diameter; peduncle 4.5-9.2 cm long, pubescent at apex; raceme flexuous at maturity; rachis 2.5-5 mm long, long villous (to 6.5 mm long) its entire length; sessile spikelet 4-7 mm long; lemma of fertile floret 2.5-5 mm long, basally indurate, split one-fifth to one-half of its length; awn twisted, 10.5- 19 mm long, inserted at the bottom of the cleft; callus pubescence to 1.5 mm long. Pedicelled spikelet 0.5-3 mm long, sterile; awn absent or up to 2.5 mm long; pedicel 3-4.5 mm long, villous its entire length. 2n = 40. (Figure 21). In openings and margins of pine woodlands on oolite. Flowering from June to August. Frequent, though overlooked when 62 not in bloom.
Distribution: Occurs (Figure 19) from Miami southward through the Florida Keys. Widespread throughout the West Indies.
Schizachyrium gracile has been considered a species of Schizachyrium primarily because it possesses only single rather than paired racemes. Clayton (1974) and Campbell (1983) maintain that S. gracile is more properly placed in the genus Andropogon because the species lacks a cupuliform rachis intemode and rounded glumes of the sessile spikelets, traits considered requisite for inclusion in the genus Schizachyrium. Since the focus of this study was not the generic traits of Andropogon and Schizachyrium, the data do not help resolve the disagreement. The morphological studies do indicate that S. gracile is quite distinct from all other taxa in the complex and observations call into question the crucial generic characters used to separate Schizachyrium from Andropogon. Robust specimens of S. gracile appear to approach the cupuliform rachis intemodes and the glumes appear rounded. Estes. Tyrl and Brunken (1982) point out the need for work on determining the limits of these two genera. Greenhouse observations confirmed the cleistogamous habit of this species however, late in the year, under a photoperiod much shorter than any to be expected in Florida, the last flowering spikelets exerted their anthers and released fertile pollen. No specimens of this species have been found to possess double racemes 63
Representative specimens: FLORIDA: Dade Co., 3 July 1977, Biem acki s.n. (FTG); SW 197 and SW 241 Junction, 1 Sep 1982, Bruner Fla-27. (OS); Deering Estate. Cutler, 26 Aug 1978, Correll & Popenoe 50086, (FTG); everglades, old Durham Hwy, 3 Dec 1962, Craighead s.n. (FTG); everglades, Old Durham Hwy, 9 Sep 1962, Craighead s.n. (FTG); Everglades National Park, 20 Oct 1963, Craighead s.n., (FTG); 29 Oct 1941, Davis s.n., (FLAS); Miami, Ju n e 1877, Garber 236. (FLAS); Krome Ave. Hialeah roadside, 13 Nov 1968, Gillis 7164. (FTG); South Miami, Spring 1942, McFarlin 12503. (FLAS); Miami, Oct 30-31, Thom pson 65. (FLAS); C hapm an Field, Coconut Grove, 2 Nov 1925, Weatherwax s.n. (FLAS); M onroe Co., Ju n e 1966, Bellinger 471. (FTG); Ramrod Key, 4 Nov 1980, Bruner Fla-12. (OS); on Sugarloaf Key, 25 Sep 1973, Correll 40117. (FTG); Big Pine Key, 8 Nov 1951, D ickson s.n. (FTG); No Name Key, 17 July 1942, Oneill s.n., (FLAS); BAHAMAS: 2 mi. E of McQueens, 15 July 1968, Byrne 522. (FTG); Grand Bahamas, NE of Freeport, 5 Nov 1973, Correll 40526. (FTG); Grand Bahamas, beach at Pelican Point, , 7 Nov 1973, Correll 40625. (FTG); N Eleuthera, between ferry slip and airstrip, 11 Jan 1974, Correll 41140. (FTG); Little Abaco, 1 mi. E of Cedar H arbour, 9 Ju ly 1974, Correll 42964. (FTG); Caicos Island, Pine Cay, 25 Aug 1974, Correll 43113. (FTG); Great Abaco, Green Turtle Cay, 26 Nov 1976, Correll 47665. (FTG); New Providence near airport, 18 1977, Correll 48403, (FTG); N Andros, S of airstrip, 8 May 1973, Correll 49713. (FTG); Andros, Mangrove Cay, 23 July 1978, Correll 50053, (FTG); New Providence, S side of Lake 64 Cunningham. 19 Sep 1978, Correll 50216. (FTG); New Providence, near airport. 7 Jun 1979, Correll 50731. (FTG); Andros, S of Red Bays, 13 May 1978, Fehling 51. (FTG); New Providence Island, west end, 2 Jan 1969, Gillis 7493. (FTG); Andros, 3 mi. S of Autec, 7 Jun 1975, Hill 3165. (FTG); Andros, Mangrove Cay, waterworks, 8 May 1973, Popenoe 255. (FTG); 2.5 mi. SE of Red Bays, 26 may 1977, Proctor 36924. (FTG); Eleuthera, N of Tarpin Bay, 2 Jun 1979, Sauleda 2645. (FTG); University of Miami, 24 May 1947, Williams s.n. (FTG); PUERTO RICO: Ftincon, Feb 1961, Gonzales 1716, (FLAS); HAITI: 28 Sep 1943, Holdridge 1725. (FLAS);
4. Schizachyrium maritimum (Chapman) Nash in Small, FI. S.E. U.S. 1903. Andropogon maritimus C hapm an FI. S. U.S. 668. 1883. Type: Florida: A. W. Chapman, 1883, "Florida." (Holotype US!) A, scoparius Michx. ssp. maritimus (Chapman) Hack. var. genuinis Hack, in DC.Phanerogamarum 6:385. 1889. Nom, illegit.
Plants decumbent, rooting at the nodes; flowering culms 37-80 cm long, glabrous with glaucous nodes; leaf blades folded, 3.5-5.5 mm wide, 6-18 cm long; sheaths keeled, 2.8-9 cm long, shorter that the intemodes; collar constricted and elongated; racemes not fully exerted, 2.5-6.3 cm long; peduncle 3.5-5.6 cm long; spathe 3.2-6.6 cm long, 3-6.5 mm wide; raceme flexuous at maturity; rachis stiff, 4- 5.5 mm long, conspicuously villous its entire length, hairs as long as 65 6 mm; sessile spikelet 9-11 mm long; lemma of fertile floret 6-8 mm long, split one-half to three-quarters of its length, awn 8-13 mm long, inserted at base of cleft; callus pubescence to 1 mm long; pedicelled spikelet 4,5-8.5 mm long, staminate, awn to 3.5 mm long; pedicel 5-7 mm long, as conspicuously villous as the rachis. 2n=40.
Characteristically at ocean waterline but also along the roads of low dune areas. Culms of this species are often covered with sand, rooting at the nodes, superficially resembling rhizomes. (Figures 22, 23).
This species can be found growing at the waterline and is often inundated with sea water. It must be the most salt-tolerant species in the group. It very effectively binds sands in the low areas along roads around Gulfport, Fla. This species also has the most extremely keeled stems and leaves of any of the taxa. Throats or collars of the leaves are very constricted, allowing the blades to twist or sway freely when subjected to winds and waves.
Distribution: Occurs (Figure 20) along the Gulf Coast from Piney Island (Wakulla Co., Fla.), westward to Mississippi.
Representative specimens: ALABAMA: Baldwin Co., 2 mi. E. of G ulfport on 180 West, 12 Sep 1982, B runer Ala-1. (OS); Alabam a Beach, 29 Sep 1949, Harper 4159. (US);11 mi. S of Foley, 25 Oct 1960, Shinners 25,888. (FSU); Dauphin Island, 18. Oct 1973, Taylor 66 15343. (NELU): Mobile Co.,Dauphin Island. 8 Oct 1967. Krai 29755. 1967, (FLAS); J. and C. Taylor. 1973. 15343. (NELU); FLORIDA: Bay Co., Holiday Lodge Marina. 11 Sep 1980, Bruner Fl-53. (OS); Mexico Beach, 15 Oct 1955, Godfrey & Krai 54204. (FSU); Beacon Hill, 6 Nov 1948, Kurz 220. (US); Dec 1970, T hom pson 139. (FLAS); Escambia Co., on Santa Rosa Island, 10 Dec 1941, Blair s.n. (FLAS); at Gulf Beach N of highway, 26 Oct 1972, Burkhalter 381. (FLAS); Pensacola Beach, Santa Rosa Island, 18 Oct 18, 1978, Godfrey, 76814. (FSU); 5 mi. E of Perdido Pass, Gulf shores, 15 Oct 1973, B. Hauso 2328. (FSU); Santa Rosa Island, 7 Nov 1964, McDaniel 5473. (FSU); Franklin Co., at entrance to St. Joe St. Park, 11 Sep 1980, Bruner Fl-50, (OS); St. Vincent Island, 27 Mar 1976, Barnes P-6-3- 27-10. (FSU); St. George Island, 9 Nov 1974, Clewell 4722. (FSU); Carrabelle Beach, 29, Nov 1954, Godfrey 52658. (FSU); Dog Island, 18 Oct 1969, Godfrey 69074, (FSU); St. George Island, 9 Nov 1971, Godfrey 71108. (FSU); St. Vincent Island, 14 Nov 1971, Godfrey 71164. (FSU); Dog Island. 30 Nov 1969, Godfrey 698278. (FSU); Cochran's Beach, 1 Nov 1941, Kurz s.n. (FLAS); St. Vincent Island, 30 Oct 1910, McAtee 1725A. (US); Gulf Co., Port St. Joe city limits (West), 11 Sep 1980, B runer Fl-51, Fl-52. (OS); MISSISSIPPI: Jackson Co., Deer Island, 1 Oct 1953, D. Demaree 34422. (US); Horn Island, 1 Oct 1895, Tracy 3798. (US); Horn Island, 6 Oct 1897, Tracy s.n. (US); H orn Island, Nov 1897, Tracy s.n. (US); 6 Oct 1897, Tracy 3786. (US); Harrison Co.,Ship Island, 27 May, 1951, D. Demaree 30583. (US); W end of Ship Island, Lenaire 904, 4 Dec 1959. (FSU); Gulfport. 25 Nov 1931, Vinall 52. (US). 67
3. Schizachyrium niveum (Swallen) Gould, Brittonia 19:73. 1967. Andropogon niveus Swallen, J. Wash, Acad. Sci. 31:354-355. 1941. Type: Florida, Oceola Co., sandy land about 15 mi. S of Kissimmee, Fla., 21 Oct 1940, W. A. Silveus 6684, (holotype, US!).
Plants caespitose; flowering culms 49-77 cm tall, glabrous; blades flattened, 2-4 mm wide, 6-10 cm long; sheaths keeled, 3.4- 5.7 cm long; racemes more or less exerted, only moderately flexuous, 3-4.5 cm long; peduncle to 4.6 cm long; spathe 2.5-4 cm long, 1.5- 3.5 mm wide; rachis 5-7 mm long, densely villous, the silvery white hairs up to 2.5 mm long contrasting sharply with the reddish-brown spikelets; sessile spikelet narrow, 5.6-6.5 mm long; lemma of fertile floret 3.5-5 mm long, split nine-tenths of its length, awn 10.5-12.5 mm long, inserted at bottom of cleft above the slightly indurate lemma base; callus conspicuously sharpened, to 1 mm long; callus pubescence up to 1 mm long; pedicelled spikelet 1-2 mm long, sterile, awn 1-2 mm long; pedicel 5-6.5 mm long and densely villous. 2n = 40. (Figure 25). In openings and sandhills of fragile Ceratiola-Pine-Oak w oodlands.
Distribution: Central-peninsular Florida (Figure 24), extremely rare. It is reported to occur in south-central Georgia but no evidence of such an occurrence is to be found. 68
This species is extremely rare and should undoubtedly be on the endangered species list. There are only two recent, extant localities recorded for S. niveum, at St. Josephine Creek and at the Archbold Wildlife Station, and only the former population has been located. Its habitat has been a favorite of real estate developers and orange growers. Perhaps it will benefit from the recent decline in the orange groves and increase in numbers but this is unlikely. Morphological analyses demonstrate the close affinity of Schizachyrium niveum, of central Florida, to S. condensatum. Schizachyrium niveum may either represent the remnant of a once widespread S. condensatum -like ancestor that survived in the uninundated Lake Wells Ridge of Central Florida (Ward, 1982) or it might simply be the result of long distance dispersal from Central America (or South America if tetraploids exist there).
Representative specimens: FLORIDA: Alachua Co., E of Gainesville, 16 Nov 1938, Swallen 5639. (US): Hernando Co.,2 mi. E of W eekiwachee Springs, 16 Oct 1959, Ray 9537. (US); H ighlands Co., Archbold Biological Station, 5 Nov 1945, Brass 15648. (US); St. Josephine Creek Bridge on US 27, 5 Nov 1980, Bruner Fl-15. (OS); Josephine Creek Bridge on US 27, 24 Oct 1964, Lakella 27805. (FLAS, FSU, FTG); Hillsborough Co.,USF Campus, 1960, Lakella 23701. (US, USF); Indian River Co.,Vero. Fla., 1 Nov 1920, Thom pson 158. (FTG, FLAS); Oceola Co.,Kissimmee, 21 Oct 1940, Silveus 6684. (US); Orange Co. Merrimack, 31 Oct 1897, Baker 34. 69 (US); Putnam Co., 1940, Laessle s.n. (FLAS).
5. Schizachyrium scoparium (Michx.) Nash in Small, FI. S.E. U.S. 59. 1903.
This species can be best represented by the recognition of five varieties:
5e. S. sco p a riu m (Michx.)Nash var. scoparium Andropogon scoparius Michx. FI. Bor. Amer. 1:57. 1803. Type: "Carolinas." (Holotype at Paris not seen; Photo of type and fragm ents at US!). Andropogon neomexicanus Nash, Bull. Torr. Bot. Club. 25:83. 1898. Type: New Mexico: Dona Ana, Wooten s.n., 1897, (holotype (SMU) not seen; isotype US!). S. scoparium (Michx.) Nash var. neomexicanum (Nash) Gould, Brittonia. 19:73. 1967. Schizachyrium acuminatum Nash, in Small, FI. SE. U.S. 59. 1903. Type: Mississippi: Oktibbeha, "Starkville, Miss., 16/11, 1890.” (Holotype NY!). Andropogon scoparius Michx. var. Jrequens Hubb., Rhodora 19:103. 1907. Type: Rhode Island: Newport, Femald, Long & Torrey 8476, 1913. (Holotype GH!, isotypes NEBC, PAS). Schizachyrium scoparium (Michx.) Nash var. Jrequens (Hubb.) Gould, Brittonia 19:73. 1967. Andropogon scoparius Michx. var. genuinis Fern. & Grisc., 70 Rhodora 37:143. 1935. Andropogon scoparius Michx. var. septentrionalis Fern. & Grisc.. Rhodora 37:143. 1935. Type: Canada: Ottawa. Rolland 19,199, 1925. (Holotype, GH!). Schizachyrium scoparium (Michx.) Nash var. septentrionalis (Fern.) Gould, Brittonia 19:73. 1967. Andropogon scoparius Michx.var ducis Fern. & Grisc., R hodora 37:145. 1935. Type: Massachusetts: Dukes Co., Fogg 2940, 1927. (Holotype, GH!) Andropogon scoparius Michx. var. genuinis forma calvescens Fern. & Grisc. Type: Virginia, Brunswick Co.,1 Oct 1942. Femald & Lewis 14,474. (Holotype GH!, isotype (not seen) at PAS).
Plants caespitose or rhizomatous; flowering culms 30-210 cm tall; blades flattened, 1.5-9 mm wide, 9-45 cm long, sheaths keeled: racemes exerted, 2-8 cm long; spathe 1.5-11 cm long, 0.5-4 mm wide; peduncle to 10 cm long; lemma of fertile floret 3-8 mm long, split from the tip to three-quarters its length; awn 2.5-17 mm long; pedicelled spikelet 1-9.5 mm long, neuter or staminate, awn to 4 mm long; pedicel 3-7.5 mm long, pubescent; callus 0.5 mm long, pubescence to 2.5 mm long. 2n = 40. An extremely variable species that occurs on a variety of soils in open habitats. An important component of the prairies. 71 Distribution: Throughout North America (Figure 26). east of the Continental Divide as well as into New Mexico. Arizona, Utah and Idaho, as far north as Alberta to New Brunswick, as far south as northern Mexico.
To say that typical S. scoparium is extremely variable is an understatement. Many of its morphologically features vary independently and continuously across North America coming together in distinctive combinations which in the past has prompted the description of several subspecific taxa. Most of these varieties have been proven unworkable, as one can often find two different "varieties" in the same population.
Representative specimens: ARIZONA: Apache Co., Gould & Robinson 5033, 1948, (TAES); Coconino Co.,Flagstaff, 29 Aug 1922, Hanson 285. (BH); Flagstaff, 13 Aug 1884, Jones 4070. (RM); 5 Sep 1943, Schallert s.n. (RM); Flagstaff, 10 Sep 1894, Tourney s.n. (RM); COLORADO: El Paso Co.,Williams Canyon, 10 Aug 1900, Harper s.n. (BH); Larimer Co., dry creek, 31 Aug 1901, Godding 576. (BH); Letterman Co., 1886, s.n. (BH); CONNETICUT: Hartford Co.,1901, Bessell s.n. (NEBC); Litchfield Co., Winsted, jet of US44 and CR8, 28 Aug 1974, Ahles 79524. (NELU); New Haven Co..19 Aug 1902, Harger 3508. (NEBC); 20 Aug 1879, Harger s.n (NEBC); GEORGIA: Oglethorpe Co., 9.3 mi. E of Lexington, 4 Oct 1964, Duncan 22388. (NELU); INDIANA: Henry Co.,Mt. Pleasant, 16 Aug 1880, Atwood s.n. (BH); Johnson Co.,Sioux City, 29 Aug 1896, Pammel 107. (BH); 72 Kossuth Co.,28 Aug 1897, Pammel 1022. (BH); IDAHO: Elmore Co.. Fall Creek, 1910, MacBride 745. (RM); ILLINOIS: Jack son Co.,S26, NW 1/4, NW 1/4, 19 Aug 1976, Heineke 1258. (NELU); Johnson Co., 4 mi. E of G rand Tower, 5 Oct 1974, Ebinger 015175. (NELU); KANSAS: Ellis Co., near Hays, 25 Sep 37, Cook s.n. (BH); Lyon Co.,15 mi. W, 2 mi. S of Emporia, s.n., 11 A ugust 1959, (NELU); R iley Co., 18 Sep 1895, Norton 582. (RM); Sedgewick Co., Wichita, 9 Sep 1890, Sm ythe 247. (BH); LOUISIANA: LaSalle Co., Lal26, 3.5 mi. S of La 124, 7 Sep 1976, Thom as 50131. (NELU); Rapides Co., S of G ardner, 11 Sep 1977, Thom as, et. al., 55181. (NELU); MASSACHUSETTES: Barnstable Co., 8 Aug 1914, Collins 2843. (NEBC); Dennis, 16 Sep 1916, Hubbard 811. (NEBC); Dukes Co., 19 Aug 1895, Harris s.n. (NEBC); Martha's Vineyard, 21 Sep 1916, Seym our 1493. (NEBC); Middlesex Co.,9 Sep 1932, Eaton s.n. (NEBC); Nantucket Co., 18 Aug 1878, Faxon s.n. (NEBC); W orcester Co., 12 Oct 1933, Seymour 4064. (NEBC); Aroostak Co.,26 Ju ly 1941, Pease & Bean 28,989. (NEBC); Hancock Co.,Mt. Desert Island, 31 July 1928, G. L. Stebbins 356. (NEBC); Knox Co.,30 Aug 1921, Long 1024. (NEBC); Oxford Co.,Lake Howell, 30 Aug 1918, Pease 17,294. (NEBC); Penobscot Co.,14 Sep 1898, M. L. Femald 2815. (NEBC); Piscataquis Cc., 3 Sep 1894, M. L. Femald s.n. (NEBC); Somerset Co.,21 Aug 1894, M. L. Fem ald s.n. (NEBC); Sagadaho Co., 4 Aug 1930, F asset 10458. (NEBC); MEXICO: Coahuila, 1963, F.W. Gould 10615. (TAES); San Luis Potosi,4 mi. E of Ciudad de Maiz, 20 Sep 1963, F. W. Gould 10800. (TAES); MISSOURI: Barton Co.,3 mi. NE of Milford, 22 Sep 1951, Palm er 53157. (NELU); n ea r M inden 73 Mines, 10 Oct 1958, Palm er 68093. (NELU); Greene Co., near Springfield, 20 Aug 1912, Standley 9084. (BH); Rollo Co.,near Oakwood, 11 Dec 1915, Davis s.n. (BH); NORTH DAKOTA: Benson Co., Leeds, 26 Aug 1907, Osterhout s.n. (RM); Kidder Co., William's Lake. 15 Aug 1917, Metcalf 284. (BH); Statsman Co., Jamestown, 24 July 1898, Schmidt s.n. (RM); Ward Co., Minot, 31 July 1903, Waldron 1803. (RM); NEBRASKA: Cherry Co.,3.5 mi. S of H12, 9 Sep 1972, C hurchill 421. (NELU); Hooker Co.,head of the Dismal River, 9 Aug 1893, Rydberg 1665. (BH); Howard Co.,S of Loup River on US 281, 16 Aug 1967, Krai 28866. (NELU); Kearney Co..Minden, 10 Aug 1891, Hapeman s.n. (RM); Lancaster Co., 1.1 mi. W of Em erald on 6, 21 Aug 1972, Churchill 365. (NELU); Sheridan Co.. 13 mi. N of Hay Spring. 18 July 1964, Nixon 172. (RM); NEW HAMPSHIRE: C hesire Co., Hinsdale, 23 Aug 1919, Batchelder s.n. (NEBC); Walpole, 31 July 1899, M. L. Femald 269. (NEBC); Troy, 11 Sep, 1897, Robinson 372. (NEBC); Coos Co..8 Sep 1914, Pease 16,315. (NEBC); Grafton Co., West Lebanon, 23 Aug 1907, Kennedy 1. (GH); Hillsborough Co.,7 Sep 1931, B atchelder s.n. (NEBC); NEW MEXICO: Clayton Co., Clayton, 19 Aug 1910, Wooten s.n. (BH); Eddy Co., 1968, Bostick 2536. (AZ); Otero Co., 1950, Norris s.n. (TAES); NEW YORK: Albany Co., W of Albany, 7 Sep 1932, Whitney 3124. (BH); Bronx Co.,in garden, 24 Aug 1918, Gershoy s.n., (BH); Nassau Co., Massapequa, 26 Aug 1918, Gershoy s.n. (BH); Richmond Co.,northwest side of Staten Island, 14 Sep 1917, Gershoy 758. (BH); Saratoga Co.,S bank of the Hudson River, 18 Sep 1899, Burnham s.n. (BH); Schenectady Co.,S of Schenectady, 11 Oct 1916, Metcalf & Wiegand 5492. (BH); Suffolk Co., EastHampton, 18 Aug 1938, Muenscher & Curtis 7833. (BH); Westchester Co.. Kensico, 21 Aug 1918, Gershoy s.n. (BH); Sullivan Co., N of High View, 9 Sep 1935, Muenscher & Curtis 5143. (BH); NORTH CAROLINA: Brunswick Co., 1 mi. S of supply on Hwy 17, 26, Oct 1980, Bruner NC-4, (OS); OHIO: Ottawa Co., Bay Point, 21 Aug 1914, J. Eames 280. (BH); OKLAHOMA: Cleveland Co..12 mi. E of Lexington, 13 Aug 1970, J & C Taylor 6880. (NELU); Comanche Co., 1 mi. W of Comanche High School, 21 Sep 1978, Rich 1325. (NELU); RHODE ISLAND: Providence Co.,Cumberland, 13 Sep 1903, Greenman 1802. (GH); SOUTH CAROLINA: Beaufort Co.,15 mi. S of Shallotte, North Carolina, 26, Oct 1980, Bruner SC-1, (OS); South of Charleston at the jet of 162 and 17 on 162, Bruner Sc-2, (OS); SOUTH DAKOTA: Brookings Co.,Brookings, 7 Sep 1898, Hefner s.n. (RM); Custer Co, Victoria Creek, 17 Aug 1909, Murdoch 3579.(RM); Grant Co., Big Stone, 1 Sep 1914, Moyer 566. (RM); Harding Co., Mt. Cave Hills, 1 Aug 1910, Vishner 36. (RM); TENNESSEE: Sevier Co.,S slope of East End, Bluff Mt., 18 Aug 1964, Thomas, et. al., s.n. (NELU); TEXAS: Bee Co., below Skidmore on Sinton Hwy, 3 Dec 1948, Tharp, et. al., 48-126. (TAES); Bell Co., near Salado, 15 Sep 1930, Walff 2 558.(TAES); Brazos Co.,.5 mi. S of Easterwood airport, 3 Oct 1975, Hill 3716. (TAES); Brewster Co., Big Bend, 13 Oct 1972, S. L. Hatch 1151a. (TAES); Gage Ranch, 7 July 1941, Wamack W582. (TAES); Briscoe Co., 11 mi. E of Silverton, 24 Sep 1953, Shinners 16349. (TAES); Brooks Co.,6 mi. S of Falfurrias, 26 Oct 1973, Lonard 4011. (NELU); Brown Co., 2 mi. NW of May. 19 Oct 1965, M cCart & Davis 32. (TAES); Chisos Co., 1932, Miller 46554. (TAES); 75 Collingsworth Co., 17 mi. N of Wellington. 25 Sep 1957, F. W. Gould & Thom as 7737. (TAES); Erath Co.. 7 mi. W of Dublin, 17 Oct 1964, J. L. H anoch 64-77. (TAES); Fayette Co.,W of Weimer, 10 Oct 1939, W. A. Silveus 5410, (TAES); Gillespie Co., 2 mi. S of Fredericksburg, 18 Sep 1942, Cory 40199. (TAES); Gregg Co., Sabine R. at 120, 10 Sep 1971, Thom as 25007. (NELU); Guadalupe Co., 10 mi. S of Seguin, 26 Oct 1973, Lonard 4003. (NELU); H ays Co.,Sabino Ranch of RR 12. Wimberly, 3 Oct 1981, P. R. H ubert 35. (TAES); Kerr Co.. 1 mi. W of Mt. Hume, 1 Oct 1961, F. W. Gould 9752. (TAES); 25 mi. NW of Kerrville, 27 Oct 1958, F. W. Gould 8458. (TAES); LaMar Co., 6.6 mi. NW of H eam e, 14 Oct 1952, Nash s.n. (TAES); McCullough Co., 3 mi. SW of Brady, 29 Sep 1967, F. W. Gould, 12426. (TAES); Mills Co.. 1 mi. W of Goldthwaite, 23 Sep 1957, F. W. Gould 7704. (TAES); 10 mi. E of Goldthwaite on 84, 11 Oct 1964, Knox 31. (TAES); Palo Pinto Co.,near Mineral Wells, 24 Sep 1953, F. W. Gould 356, Real Co., 16.25 mi. N of Leakey, 13 Sep 1937, Cory & Parks 24,349. (TAES); Robertson Co., 10 mi. SE of New Baden, 22 Oct 1956, F. W. Gould 7266. (TAES); Sutton Co.,24 mi. SE of Sonora, 17 Oct 1938, S tuckens 30775. (TAES); UTAH: Em ery Co., 10 mi. S of San Rafael River bridge, 1 Sep 1970, W elsh 10819. (NELU); VIRGINIA: Surrey Co., VEPCO Power Station, 25 Sep 1974, Ware 6045. (NELU); VERMONT: Chittenden Co., Burlington, 10 Aug 1883, Brainerd s.n. (GH); Essex Co.. 11 Sep 1940, Pease 28,440. (NEBC); WEST VIRGINIA: Hardy Co..3 mi. W of Old Fields, 21 Aug 1961, Wratchford s.n., (NELU); WYOMING: Albany Co.,15 mi. S of Laramie, 21 Aug 1901, Merrill & Wilcox 444. (BH); Park Co., Wind River, 1892, 76 Nelson 5075. (BH); CANADA: New Brunswick, York Co., 1922, M. L. Fem ald & Pease 24,832, (GH); Grondine, comte de Portneuf, 4 Aug 1935, Victorin 44, 066. (GH); lie Bizard, 15 Sep 1935, Victorin 43,556. (GH); Norway Bay.Norway Bay. 30 Aug 1935, Victorin 43,240. (GH); W altham , 14 Aug 1933, Victorin 43,911. (GH); Woodstock,1931, Victorin 44,639. (GH); Ottawa, Pontiac, 1 Aug 1932, Victorin 45,766. (GH); Q uebec, Richmond, Bromptonville, 13 Sep 1975, F orest 1298. (NELU).
5d. S. sc o p a riu m (Michx.) Nash var. d iverg en s (Hack.) Gould, Brittonia 19:73. 1967. Andropogon scoparius Michx. ssp. m aritim us var divergens Hack, in DC Phanaerogamarum 6:385. 1889. Type not seen. Andropogon divergens (Hack.) Anderss. ex H itchc., J.Wash. Acad. 23:456. 1933. A scoparius Michx. var. villosissimus Keam. in Scribn. & Ball, Bull. U.S. Dept. Agr. Agrost. 24:40. 1901. Type: Mississippi, along railroad at Waynesboro. 2 October 1896; T. J. Kearney 136. (Holotype, OS!). Schizachyrium villosissimum Nash in Small, FI. S.E. U.S. 59. 1903. A. scoparius Michx. var. virilis Shinners, Rhodora 56:36. 1954. Type: Texas: Upshur Co., 3.3 mi. S. or New Diana, 15 Sep 1953. Shinners 16,009. (Holotype SMU not seen. 77 isotype US!). Schizachyrium scoparium (Michx.) Nash var. virile (Shinners) Gould, Brittonia 19:73. 1967.
Culms 70-180 cm tall; leaves sparsely to densely pubescent, blades 3.5-5 mm wide, 12-30 cm long, sheaths often densely villous, throat pilose; racemes 3-5 cm long, of 7-12 spikelets; rachis 3.5-6 mm long, pubescence 1.5-3 mm long, often sparse; sessile spikelet 6-10 mm long, fertile lemma split from its tip less than half of its length, awn 9-15 mm long; pedicel 3.5-6.5 mm long; pedicelled spikelet 2-9 mm long, rarely staminate, awn to 2 mm long. (Figure 28). Distribution: This variety is common throughout the south- central pinelands of the United States (Figure 26) and varies widely as to degree of leaf pubescence. Leaf vestiture becomes confined to the sheaths as the variety extends toward the southeastern part of its distribution.
This variety is confined to the south-central United States but is not as ecologically defined as the other varieties. Morphologically it merges with typical S. scoparium and it could be argued that maintenance of this variety is not necessary. It does serve however, to mark the presence of a trend of vegetative pubescence. Foliage is long villous in the western part of its range with pubescence becoming shorter, more dense and largely confined to the sheaths toward the East into Mississippi. A. S. Hitchcock located the type specimen in the Berlin Herbarium and this type was presumably 78 destroyed during the destruction of the herbarium in World War II.
Representative specimens: LOUISIANA: Baton Rouge Co.,near Amite River and US 190, 20 Oct 1955, B arrett 36. (NELU); B ien ville Co., 1 /4 mi. N of Natchitoches, 16 Sep 1975, Parish. (NELU); B ossier Co., 4.5 mi. S of Plain Dealing at Collinsburg Creek, 14 Oct 1975, Thom as & Leggett 47742. (NELU); Caddo Co.,S of jet of La 2 and Lai on La 1, 16 Oct 1973, Thom as & Overby 37372. (NELU); Caldwell Co., S23, T14N, R3E, 21 Oct 1972, Max & Raym ond 224. (NELU); 8 mi. N of jet . of La 846 and La 4, 16 Sep 1973, Max & Wells 1591. (NELU); 2 mi. SE of Copenhagen, 21 Sep 1967, T hom as & Brode 4562. (NELU); .0.5 mi. S of O uachita Parish, 28 Sep 1973, Thom as & Max 37086. (NELU); Catahoula Co.,S of Suty, 21 Oct 1975, Thomas 47997. (NELU); Claiborne Co., 1.5 mi. S of Arkansas line on Us 79, 2 Aug 1978, Thom as, et. al., 59689. (NELU); DeSoto Co.,0.3 mi. N of La 346, 28 Sep 1978, Thom as & Dixon 61405. (NELU); Evangeline Co., N of T urkey Creek, 25, Sep 1977, Thom as, et. al., 55501. (NELU); Grant Co., SW of Georgetown, 14 Oct 1972, Thomas 33330. (NELU); Jefferson Davis Co.,US 165 N of Fenton, 18 Oct 1975, Thomas 47921. (NELU); LaSalle, 5 mi. N of Summervile, 23 Sep 1967, Thom as 4639. (NELU); Morehouse Co.,N of Sterlington, 27 Oct 1977, Thomas & Pias 56276. (NELU); Ouachita Co., SE of Luna. 12 Sep 1973, Thomas & Scurrier 36866. (NELU); 0.5 mi. N of Monroe, 11 Oct 1968, Thomas 12730. (NELU); 2 mi. S of Luna, 1 Oct 1970, T hom as 21441. (NELU); S of La 838, 16 Sep 1971, Thom as 25485. (NELU); Rapides Co., 5 mi. NW of Glenmora, 6 Oct 1978, Schutz 79 1709. (NELU); Flatwoods, 23 Sep 1971, Thom as 25735. (NELU); S of G ardner, 11 Sep 1977, Thom as et. al., 55181. (NELU); Red River Co., 4 mi. N of Coushatta, 3 Oct 1967, Thomas 5687. (NELU); St. H elena Co., 0.5 mi. W of Tangipahoa Parish, 28 Oct 1978, Thomas 62061. (NELU); St. Mary Co.,beside La 182, Berwick High School, 31 Oct 1971, Thom as et. al., 27122. (NELU); Vernon Co., 1 mi. N of the jet of 465 and Hwy 117, North of Kurthwood, 12 Nov 1980, Bruner La-1. (OS); US 171, N of Lessville, 22 Sep 1971, Thom as 25768. (NELU); TEXAS: Angeline Co., 2 mi. E of Zavalla, 29 Sep 1967, F. W. Gould 12442. (TAES); Brazos Co., 1.5 mi. inside College Range, 13 Oct 1966, Beason 12. (TAES); 2 mi. S of College Station, 20 Oct 1951, F. W. Gould 6159. (TAES); Sep 1938, Malone s.n. (TAES); W of College Station, 24 Sep 1966, Whitson 7. (TAES); Cass Co., 1.5 mi. NW of the Tx/La state line on Tx 77, 13 Nov 1980, Bruner Tex-7, (OS); Chambers Co., 3 mi. S of Winnie, 5 Oct 1967, Gould 12452. (TAES); Fayette Co.,W of Weiner, 10 Oct 1939, W. A. Silveus 5517. (TAES); Grimes Co., 14 mi. E of Navasota, 2 Oct 1956, Gould 7258. (TAES); 8 mi. E of Navasota, 23 Sep 1953, Gould s.n. (TAES); Hardin Co., 6 mi. SW of Kourtze, 24 Sep 1966, Gould 12007. (TAES); Henderson Co.. 1.6 mi. NE of Chandler, 2 Oct 1951, S hinners 19163. (TAES); LaMar Co., 1953, S hinners 16349. (TAES); Montgomery Co., 10 mi. NW of H untsville on 30, 12 Oct 1975, S. L. H atch 2116. (TAES); N ew ton Co., 1967, Krai 29132. (NELU); 12.4 mi. NE of Ja sp e r on 63. 11 Nov 1980, Bruner Tex-6, (OS); 3 mi. E of Kirbyville on 363, 30 Aug 1967, Krai 29132, (NELU); 16 mi. S of Newton, 4 Oct 1934, Parks & Cory 10857. (TAES); Orange Co., 10 mi. N of Orange, 1 Nov 1940, W. A. 80 Silveus 6764. (TAES); San Augustine Co., E of Zavalle, 19 Sep 1952, F W. Gould 6537. (TAES); Tyler Co., 1981, Pecottel 54. (TAES); Walker Co., 8 mi. E of Huntsville, 24 Oct 1956, F. W. Gould 7323. (TAES); 9.5 mi. N of Huntsville, 29 Sep 1934, Parks & Cory 10357. (TAES).
5c. S. sco p a riu m (Michx.) Nash var. litto ra le (Nash) Gould, Brittonia 19:73. 1967. Andropogon littoralis Nash in Britton, Manual Flor. North. St. Can. 1901. Type: New York, Richmond Co, on Staten Island 16 Oct 1894, Kearney. (Holotype NY not seen; isotype US!).
Culms 39-160 cm tall, glaucous, extremely compressed with proliferating basal innovations; leaf blades 3.5-6.5 mm wide, auricles yellowed, flexible; ligule 1.5-2 mm long; raceme of 8-11 spikelets, flexuous at maturity; rachis densely covered three-fourths of its length from the tip with long (3-6 mm), silvery hairs; sessile spikelet 8-10 mm long, awn of fertile lemma 10-16 mm long; pedicelled spikelet 2.5-4 mm long. (Figure 29).
Distribution: This variety grows on the shifting, coastal sand dunes of the Texas coast (from Port Aransas southward to South Padre Island), the Mid-Atlantic coast (from southern New York to northern North Carolina) and the east side of Lake Michigan (Figure 27). 81
This variety is a colonizer of unstable sand dunes on the margins of oceans and lakes. It makes an excellent dune binder. It has a distribution that identifies it as a member of a group of species first identified by Peattie (1922) [and much later by Pierce (1974)] that have similar disjunctions. Its distribution is a result of post Pleistocene dispersal. The variety has apparently been extirpated from Lake Erie (Ohio) due to elimination of its sand dune habitat but it still occurs at the Indiana Dunes National Lakeshore near Michigan City, Indiana and on the sand dunes of northern Michigan. Plants of the Atlantic coast (tall) and Texas coast (short) are ecotypes. Occasionally other plants of Schizachyrium scoparium can be found growing on sand dunes and may vegetatively resemble var. littorale but inflorescence characters will not be correct. Femald named var. ducis based upon such a specimen. Var. littorale is often confused w ith S. maritimum, they occur in somewhat similar habitats, but the large, pedicelled (often staminate) spikelets of the latter separate the two taxa.
Representative specimens. DELAWARE: Sussex Co.,Bethany Beach, 12 Oct 1940, C hase 12625. (US); INDLANA: Porter Co.. Indiana Dunes National Lakeshore, 3 May 1983, Bruner In-1. (OS); Pine, 1 Sep 1897, Chase 580. (US); MICHIGAN: M ackinack Co., 1947, Ball 3060. (US); NEW YORK: Richm ond Co.,Kearney s.n., 1894, (US); NEW JERSEY: Cape May Co.,Sandy Hook, 1 Oct 1885, Britton s.n., 1885, (US); Atlantic Co., Cape May, 4 Oct 1883, Bunk 82 s.n. (US); Atlantic City, 7 Sep 1884, L.F.W. s.n. (US); Atlantic City, Sep 1884, Vasey. (US); NORTH CAROLINA: Brunswick Co., Long Beach, 28 Oct 1950, Boyce & Godfrey, (US); Carteret Co., Near Fort Mason, 29 Oct 1937, Blomquist 10,200. (FLAS); Atlantic Beach at end of Hwy 58, 25 Oct 1980, Bruner Nc-2. (OS); Emerald Isle dunes, 25 Oct 1980, B runer, Nc-1. (OS); Dare Co., 6 mi. N. of Whalebone junction, 19 Sep 1974, McCaffrey s.n. (FLAS); New Hanover Co.,Wrightsville Beach, 26 Oct 1980, Bruner Nc-3, (OS); S of Wrightsville Beach, 27 Oct 1973, Hosier 4289. (FLAS); Onslow Co.,S of New River Inlet, 14 Sep 1950, Boyce 1497. (FLAS); Bear Island. 21 Oct 1961, Wilbur & Beal 6622. (FSU); Bear Island. 22 Oct 1961. Wilbur & Beal 6666. (FSU); OHIO: Erie Co., Cedar Point. 3 Aug 1902, Morriss A143. (US); TEXAS: Cameron Co., 18 mi. E of Brownsville on Hwy 4, 17 Oct, 1970. R. Lonard 2815 (TAES); 15 mi. N of Pt Isabel, 11 J u n 1968. F. W. Gould 12532. (TAES); Boca Chica Beach, 10 Dec 1937. Lynch & Hotchkiss 1118. (US); Del Mar Beach, 20 Nov 1940, H. B. Parks 2394, (TAES); Kennedy Co.,6.3 mi. S of Sasita, 15 Oct 1946, Lundell 14692. (US); 1949, Swallen 10594. (US); Kleburg Co., 2 mi. N of Riviera, 15 Oct 1946, Lundell 14685. (US); Nueces Co., along CR-358 into Padre Island, 11 Nov 1980, Bruner Tx-2.(OS); Padre Island, 21 Jan 1939, V. L. Cory s.n. (TAES); Refugio Co., Port Aransas ferry crossing, 11 Nov 1980, Bruner Tx-4. (OS); 5.5 mi. N. or Fulton city lim its, 11 Nov 1980, B runer Tx-5. (OS); San Patricio Co.,Padre Island dunes, 11 Nov 1980, Bruner Tx-3. (OS); Corpus Christi, 12 Nov 1939, Silveus 5475. (US); Aransas Pass, 22 Dec 1948, Thorpe & Brown 48-207. (US); Corpus Christi, 3 Dec 1948, Thorpe et al 48-1. 83 (US); VIRGINIA: Princess Anne Co.. Cape Henry, 23 Sep 1933, Femald & Griscom 2768. (FSU); 2 mi. N of Virginia Beach, 3 Sep, 1959, Krai 9485. (FSU).
5b. S. sc o p a riu m (Michx.) Nash var. p o ly c la d u s (Scribner and Ball) Bruner comb. nov. A. scoparius Michx. var. polycladus Scribner and Ball. USDA Bull. No. 24. 40-41. 1901. Type: Florida: Manatee Co. Braidentown, 3 Oct 1898. R. Combs 1298, 1898. (Holotype US!) Schizachyrium stoloniferum Nash, Flor. S.E. U.S. 59. 1903. (Type: Florida: "Herb. Chapman., Florida, Southern Flora!," holotype, NY!). Andropogon stolonifer (Nash) Hitchc., Am. J. Bot. 2:299. 1915. S. triaristatum Nash, Flor. S.E. U.S. 60. 1903. Type: "Florida" Ex Herb. Geo Thurber Leg. Chapman," GH! S. stoloniferum Nash var. wolfei DeSelm, Sida 6(2): 114. 1975.
Culms 58-210 cm tall, producing long, scaly rhizomes; leaves usually pubescent near the collar on both sheath and blade, blades 3.5-9 mm wide, 10-39 cm long; rachis with villous hairs up to 4.5 mm long; sessile spikelet 5-10 mm long; fertile lemma awn 6-14 mm long; callus pubescence to 2.5 mm long. (Figure 32) This variety inhabits the sandy soils of woodland openings and roadsides. Blooms from September through October. 84 Distribution: Throughout Florida north of the Everglades oolite reaching into southern Georgia and Alabama (Figure 30).
This is a variety that has one of the wider distributions. There are two types of plants to be found in its range: northern populations are made up of few, widely spaced, weak culms on rather bare sand, while southern populations are vigorous, dense stands composed of taller, robust culms. These southern populations appear to be spread along roadsides by grading equipment. Natural seed set is very low, and it is speculated that this variety's flowering and seed production may be extremely sensitive to changes in environmental conditions. Some clones, especially those of the south may be largely sterile. One collection (by A. S. Hitchcock at US) has been made along the coast of Vera Cruz, Mexico. This specimen probably represents an errant propagule taken across the Gulf of Mexico by man or hurricane. Collection in Vera Cruz has found no present-day populations.
Representative specimens: ALABAMA: Baldwin Co., outside of Barnwell city limits on Hwy 98, 9 Nov 1980, Bruner Ala-2. (OS); GEORGIA: Carlton Co.,east of Enigma on 82, 28 Aug 1982, Bruner Ga-2. (OS); Cypert 243, 1959, (UG); Duncan 5750, 1943, (UG); Decatur Co., Godfrey 74098, 1974, (FSU); McIntosh Co.,D uncan 20652, 1956, (UG); Grady Co..Faircloth 622, 1963, (UG); OciUa Co.. 5 mi. southeast of Waycross on Hwy 23, 29, Aug 1982, Bruner Ga-1. (OS); Toombs Co.,Duncan 14524, 1952, (UG); FLORIDA: Alachua Co., 85 Gainesville, 1 Oct 1907, Chase 4246. (US); Gainseville, 1 Oct 1907, C hase s.n. (US); Gainesville, 2 Dec 1933, Reed 18. (US); 18 Sep, 1940, Young s.n. (FLAS); Baker, E of McClenny, 23 Oct 1975, Godfrey 74666. (FSU); Olustee Nat. Forest. 28 Nov 1955, Krai 1862. (FSU); Bay, 3 mi. e of Lynn Haven, 3 Nov 1961, Godfrey 61606, 1961, (US, FSU); Brevard. 4 mi. W of Cocoa. 20 Ju l. 1957, Krai 5281. (FSU); on Merritt Island. 27 Aug 1957, Krai 5557. (FSU); Calhoun, 4.5 mi. NW of Clarkesville, 18 Oct 1957, Ford 5622. (FLAS); 2 mi. W of Clarkesville, 15 Nov 1958, Godfrey 57915. (FSU); .5 mi. N of Youngstown, 2 Nov 1961, Godfrey 61602. (US, FSU); Charlotte, on the approach to P unta Gorda, 6 Nov 1980, B runer, FI-17, (OS); 8 Sep 1946, Frye C-12. (FLAS); Citrus Co., across Hwy 41 from the Ivemess post office, 8 Nov 1980, B runer Fl-46, (OS); Clay Co.,9.2 mi. S of Green Cove Springs on Hwy 17, 27 Oct 1980, B runer Fl-1. (OS); 3.5 mi. W. of Carrabele Beach on Hwy 98, 8 Nov 1980, Bruner Fl-25. (OS); on SR 218 at Carter Spencer Road, Middleburg, 29 Aug 1982, Bruner Fl-29. (OS); 9.2 mi. S of Green Cove Springs, 6 Sep 1937, Survey s.n. (FLAS); Collier Co.,3 mi. south of Imolakee on 29, 5 Nov 1980, B runer FI-13. (OS); w est of Imolakee, 2 mi. W. of Hwy 29 on 82, 11 Nov 1980, Bruner Fl-14, (OS); Naples, 5 mi. E of 561 junction with 41S on 41S, 1 Sep 1982, Bruner Fl-28. (OS); Cooley 9061, 1962, (USF); Lakela 31585, 1968, (USF); Dade Co.. 1.2 mi. E of jet. of 94 and 27 on 94, west of Kendall, 1 Sep 1981, Bruner Fl-41, 1982, (OS); DeSoto Co.,Arcadia, Oct 1898, Com bs 1281. (US); Oct 1920, Thompson 201. (FLAS); west of Arcadia, 6 Nov 1980, Bruner Fl-16 (OS); Destin Co., 2 mi. E. of Destin on Hwy 98, 11 Nov 1980, 86 B runer Fl-54, (OS); Dixie, north of Cross City at the junction of CR 358, 8 Nov 1980, B runer Fl-47. (OS); 11 mi. NW of Cross City, 8 Nov 1969, Godfrey 69732. (FSU); Duval, SW of Jacksonville, 10 Oct 1944, Neal 2187. (FLAS); E of Jacksonville, 5 Oct 1939, Silveus 5330. (US): Franklin Co., 3.5 mi. west of Carrabelle Beach, 8 Nov 1980, Bruner Fl- 4 9 .(OS); SW of S um atra, 12 Nov 1969, Godfrey 69227. (FSU); Gadsen Co., on Ochlockonee Road at Iron Bridge, 8 Oct 1969, Godfrey 68951. (FSU); 26 mi. W of Tallahassee, 12 Nov 1955, Krai 1793A. (FSU); Hendry Co.,Davis, s.n., 1947, (USF); Hernando Co.,3.5 mi. W of Brooksville on Hwy 50, 7 Nov 1980, Bruner Fl-45. (OS); Brooksville, 18 Nov 1938, J. R. Swallen 5647. (US); Highland Co., H ighlands Hamm ock St. Park, 6 Nov 1980, B ru n er FI-18. (OS); Sebring city limits on 98, 31 Aug 1981, Bruner Fl-39. (OS); at junction of 70 and 27 on 27, 31 Aug 1981, Bruner Fl-40. (OS); Sebring, Fla., 24 Sep 1934, M cFarlin 9578. (FLAS); Sebring, Fla. 24 Sep 1934, McFarlin 9587. (FLAS); Hillsborough Co.,1/4 mi. W of McDonalds at Plant City, Bruner Fl-22, 1981, (OS); Tampa, 6 Oct 1898, Combs 1348.(US); Industrial park W of 4th St., 4 Oct 1963, Lakella 26521. (FLAS); Indian River Co.,1.1 mi. E of Yeehaw on 60E to Vero Beach, 29 Oct 1980, Bruner Fl-8. (OS); Jackson Co.,S of Sneads, 24 Oct 1977, Godfrey 76092. (FSU, FLAS); Mitchell 936, 1960, (FSU); Jefferson Co.,6 mi. NNE of Chaires, 5 Oct 1961, Godfrey 61532. (FSU); 2.5 mi. E of Wacissa. 20 Sep 1969, Godfrey 68836. (FSU); Lake Co., south of Clermont on C-651, north of the intersection with 33, 30 Aug 1981, B runer Fl-36. (OS); Leon Co.,on Welaunee Plantation, 7 Oct 1974, Clewell s.n. (FSU); W of 87 Tallahassee, 30 Sep 1964, Godfrey 64681. (FSU); near Larkins Bluff along Ochlockonee, 9 Oct 1969, Godfrey 68961. (FSU); near Larkins Bluff along Ochlockonee, 9 Oct 1969, Godfrey 68972. (FSU); along Forest Road 307, 18 Oct 1975, Godfrey 74596. (FSU. FLAS); on Ochlockonee River, 18 Oct 1955, Krai 1698. (FSU); 26 mi. W of Tallahassee, 18 Nov 1955, Krai 1792. (FSU); 26 mi. W of Tallahassee, 12 Nov 1955, Krai 1793. (FSU); 6 mi. NNE of Chaires, 12 Oct, 1956, Krai 3680. (FSU); Leon Co., 14 mi. E of Tallahassee, 22 Sep 1957, Krai 5886. (FSU); Levy Co.,1 mi. W of Janney, 30 Oct 1964, Godfrey 64757. (FSU); 3 mi. SW of Otter Creek, 13 Sep 1954, Perdue 1736. (FSU); Bronson, 7 Sep 1936, Survey s.n. (FLAS); Liberty Co.,E of Hosford, north side of the road, 2 Sep 1981, Bruner FI-19. (OS); 4 mi. S of Wilma, 16 Oct 1975, Clewell 4811. (FSU); Torreya St. Park, 12 Nov 1973, Godfrey 73145. (FSU); Madison Co.,5 mi. S of Lee. 20 Oct 1975, Godfrey 74615. (FSU); 8 mi. SSW of Ellaville, 11 Oct 1957, Krai 6187. (FSU); Manatee Co., Palmetta, 22 Aug 1895, Nash 2443. (US); Braidentow n, 1 Oct 1900, Sim pson 7092, (US); Nassau Co., Fernandina, 24 Oct 1975, Godfrey 74683. (FSU); Oceola Co.,2 mi. north of Holopaw at the junction with 441, 29 Oct 1980, Bruner Fl-6. (OS); at the junction of 441 and 92 13.6 mi. S of Holopaw, 29 Oct 1980, Bruner Fl-7. (OS); Sep 1947, Davis s.n. (FLAS); Okaloosa Co.,7 mi. NW of Baker, 27 Oct 1965, McDaniel 7149. (FSU); O kechobee Co., north of the Okeechobee city limits on 441, 30 Oct 1980, Bruner Fl-11. (OS); 2.1 mi N of 630A and 27 ju n ctio n on 27, 31 Aug 1981, B runer Fl-38. (OS); Orange Co., Grasmere E., 23 Sep 1898, Baker 1129. (US); Grasmere, 21 Sep 1898, Combs & Baker 1096. (US); 88 Palm Beach Co.. 1957, Krai 568. (USF); Polk Co..south of the junction of 27 and 60 on 27, west side of road, 1 Aug 1981, Bruner Fl-37. (OS); 4 mi. S of Dundee on 27, 7 Nov 1980, B runer Fl-44. (OS), 1 mi. N of Bartow on 98, 11 Dec 1969, Oswalt s.n. (FLAS); Putnam Co., 0.75 mi. N of the junction of 309 and the road to Pomona Park, 27 Oct 1980, Bruner Fl-2. (OS), at the San Mateo city limits on 207S, 29 Aug 1981, Bruner Fl-31. (OS); 9 mi. S or Melrose, 6 Nov 1969. Godfrey 69519. (FSU); Welaka, 28 Sep 1940, Laessle s.n., (FLAS); Santa Rosa Co.,5 mi. W of Munson, 14 Sep 1957, Ford&Wet 5592. (FLAS); Sarasota Co.,13 Oct 1940, McFarlin, £t ai. 12064. (US); Seminole Co., 7 mi. S of Geneva at Lake Mills Road, 28 Oct 1980, Bruner Fl-5. (OS), 2.5 mi. W of Sanford on 46, 30 Aug 1981, Bruner Fl-34. (OS), 2.5 mi. E of Sanford on 46, 30 Aug 1981, Bruner Fl-35 (OS); St. Johns Co.,at the jet. of 13 and 16 on 13, near the Ward's Creek Community Church, 29 Aug 1981, Bruner Fl-30. (OS); St. Lucie Co., 0.3 mi. W of Savannah County Park entrance, 29 Oct 1980, Bruner Fl-9. (OS); at the jet. of c-712 and State Hwy 70 on 70, 30 Oct 1980, B runer FI-10. (OS); Sumter Co., south of Bushnell on east side of road, 2 Sep 1981, Bruner Fl-23. (OS); Taylor Co.,2 mi. SE of Perry on Hwy 98 near the junction with C-361, 2 Sep 1981, Bruner Fl-20. (OS); 1.2 mi. SE of Salem, 7 Oct 1964, Godfrey 64719. (FSU); V olusia Co., at the gate to Valentine Park outside of Orange City on the road to Blue Springs St. Park, 28, )ct. 1980, Bruner Fl-3. (OS), S of Sanford north of the intersection of US 17/92 and 1/2 mi W of Geneva city limits, 28 Oct 1980, Bruner Fl-4. (OS), 1/2 mi. N of DeLeon Springs city limits on 17N, 30 Aug 1981, Bruner Fl-32. (OS); 89 just W of the jet of C439 and 44E on 44E, 30 Aug 1981, Bruner Fl- 33. (OS); Orange City, 4 Oct 1940, Hood 71877. (FL.\S); Wakulla Co., Newport city limits on Hwy 98, west edge of city, 8 Nov 1980, Bruner Fl-48. (OS); St. Marks Wildlife Refuge, 2 Oct 1961, Godfrey 61518. (FSU); 7 mi. N of Smith Creek, 15 Oct 1972, Godfrey 72245. (FSU); 5 mi. E of Wakulla Station on 98, 27, Oct 1970, Lazor 5080. (FSU); 5 mi. E of Wakulla Station of 319, 27 Oct 1970, Lazor 5082. (FSU); 2 mi. E of Wakulla Station on 319, 27 Oct 1970, Lazor 5125. (FSU); W alton, S30, TIN, R21W,. 16 Nov 1966, Chapman 0223. (FLAS); at Mossy Head, 3 Oct 1969, Godfrey 68892. (FSU); Seagrove Beach, 6 Nov 1976, Godfrey 75764. (FSU); 5.5 mi. E of Argyle, 6 Nov 1977, Godfrey 76216. (FSU); 4 Oct 1940, Hum e s.n. (FLAS); S5, T1S, R21W, 2 Oct 1950, Tyson 445. (FLAS).
5a. S. scoparium (Michx.) Nash var. rhizomatum (Swallen) Bruner comb, et stat. nov. Andropogon rhizomatus Swallen, J. Wash. Acad. Sci. 31:352- 353. 1941. Type: Florida, Dade, 16 Oct 1940, W. A. Silveus 6614, (Holotype US not found, paratype, Royal Palm State Park, W. A. Silveus 6606, US!. Schizachyrium rhizom atum (Swallen) Gould, Brittonia 19:73. 1967.
Culms 60-90 cm tall, usually glabrous; producing short scaly rhizomes; blades 1.5-3 mm wide, typically folded, 9.5-19 cm long; sessile spikelet 5-7.5 mm long; fertile lemma awn 2.5-10 mm long; 90 callus pubescence sparse, to 1.5 mm long. (Figure 33). Found in open glades and along the margins of open areas of pine woodlands, on oolite. Blooms from August to early September.
Distribution: Throughout the Everglades Keys from Miami southward into the Florida Keys to Big Pine Key (Figure 31).
Schizachyrium rhizomatum is confined to the marl outcrops of extreme southern Florida.where the thin soil is often saturated with water. These plants are much smaller than those of var. polycladus , the leaves are no wider than 3 mm and folded. It grows in sparse stands occasionally with S. gracile.
Representative specimens: FLORIDA: Dade Co., W of Kendall, 16 Sep 1960, Atwater s.n. (FLAS); Old Cutler Rd and SW 62 Avenue, 29 Aug 1965, Avery 464A. (FTG); Kendall, Florida, 14 Sep 1968, Avery 466. (FTG); E of Tam iam i airport, 14 Aug 1977, Avery 1765. (FTG); Sisal Hammock, 8 Oct 1966, Bellinger 556. (USF); Everglades National Park, 22 Oct 1966, Bellinger s.n. (FTG); 1/4 mi. W of the intersection of Redland Road and SW216 St. outside of Homestead, 1 Sep 1981, Bruner Fl-42. (OS); Palm Drive, pineland, 16 Sep 1962, Craighead s.n. (FTG); Mahogany Hammock Rd., 23 Aug 1963, Craighead s.n. (FTG); between Coral Gables and Black Creek, 26 Aug 1968, Lakela 31585. (USF); SW of Homestead. Fla., 15 Oct 1940, W. A. Silveus 6606. (FSU); Murden Hammock, 22 Jun 1915, Small & M osier s.n. (FTG); Monroe Co.,Big Pine Key on Wilder Rd., 29 Aug 91 1965, Avery s.n. (FTG); Big Pine Key, 31 Aug 1965, Avery s.n. (FLAS); Big Pine Key, 31 Aug, 1981, B runer Fl-56. (OS); Big Pine Key, 31 Aug 1981, Bruner Fl-57. (OS); Big Pine Key, NE of Watson Hammock, 1965, Craighead s.n. (FTG); Big Pine Key, 26 Oct 1950, Killip 40604. (US); Big Pine Key, 21 Nov 1950, Killip 40785. (US); Big Pine Key, 5 Dec 1950, Killip 40818. (US).
2. Schizachyrium cubense (Hack.) Nash, in North American Flora 17:109. 1912. Andropogon cubensis Hack., Flora 68:121. 1885. Type not seen: Wright 3898. (Not at US). Sorghum cubense (Hack.) Kuntze, Rev. Gen. 791. 1891.
Culms to 71 cm tall, producing short, scaly rhizomes; leaf blades lightly pubescent, 1 mm wide, from 6-12 cm long; raceme 4-5 cm long, containing up to 6 spikelets per raceme; sessile spikelet large, 5-6 mm long; fertile lemma to 3.5 mm long, awn to 8 mm long; pedicelled spikelet 2-3 mm long, awn to 1.5 mm long.
This species is not well known. It apparently grows on sand. With only three, poor specimens at US little can be said of the nature of its variation or to what species it is closely related. In aspects of its spikelets, and rhizome production it seems quite close to S. rhizom atum but the filiform leaves (if not a phenotypic response to the tropical, maritime habitat) suggest some affinity to S. gracile. 92 Distribution: Cuba, at Pinar del Rio. and the Island of Pines (Figure 34).
Representative specimens: CUBA: Damuji. 31 May. 1920, E. L. Ekman 11049. (US); Santa Barbara, 1920, E.L. Ekman 11993. (US); Isle of Pines, Feb-Mar 1916, Britton & Wilson 14291. (US). 93
Literature Cited
Allred, K. W. 1981. Cousins to the South: Amphitropical disjunctions in southwestern grasses. Desert Plants (3)2: 98-106. Burduja, C. and C. Toma. 1971. Pilot data on the experimental and ecological anatomy and histology of some grasses. 7. Leaves of Deschampsia Jlexuosa (L) Trin. Herb. Abs. 41:3030. Campbell, C. S. 1983. Systematics of the Andropogon virginicus complex (Gramineae) Journ.. Arnold Arbor. 64:171-254. Chapman, R. H. and S. B. Jones. 1985. Ecotypic differentiation of Andropogon virginicus (Gramineae). Bull. Torrey Bot. Club 102(4):166-171. Church, G. L. 1929. Meiotic phenomena in certain Gramineae. II. Paniceae and Andropogoneae. Bot. Gaz. 87: 608-629. Cornelius, D. R. 1947. The effect of source of Little Bluestem grass seed on growth, adaptation and use in revegetation seedlings. Jour. Agr. Res. 74:133-143. Davidse, G. and R. W. Pohl. 1974. Chromosome numbers, meiotic behavior and notes on tropical American grasses (Gramineae). Canadian Jour. Bot. 52(2):317-328. Ellis, R. P. 1976. A procedure for standardizing comparative leaf anatomy in the Poaceae. I. The leaf-blade as viewed in transverse section. Bothalia 12(1)65-109. Fuelgen, R. and H. Rossenbeck. 1924. Mikroskopish chemisher nachweis einer nucleusaure vom typus der thymonucleusaure 94 Fuelgen, R. and H. Rossenbeck. 1924. Mikroskopish chemisher nachweis einer nucleusaure vom typus der thymonucleusaure und die darauf beruheude elektive farbung von zellkemur in mikroskopischen praparatin. Hoppe-Seyler's Z. Physiol. Chem. 135:203-252. Gould, F. W. 1956. Cytotaxonomy of the Andropogoneae. Amer. J. Bot. 43:395-404. ______. 1967. The grass genus A ndropogon in the United States. Brittonia 17(l):70-76. Hackel, E. 1885. Andropogoneae Novae. Flora 7:115-128. Hatch, S. L. 1975. A biosystematic study of the Schizachyrium cirratum - S. sanguineum complex (PhD thesis, Texas A & M University, 1975). Hilu, K. W. 1984. Leaf epidermises of Andropogon sect. Leptopogon (Poaceae) in North America. Syst. Bot. 9(2):247-257. Hitchcock, A. S. 1909. Grasses of Cuba. Contr. U.S. Natl. Herb. 12(6): 183-258. Hitchcock, A. S. 1936. Manual of the grasses of the West Indies USDA. Misc. Publ. 243. 385-402 pp. ______1950. Manual of the Grasses of the United States (Agnes Chase, rev. ed.). Dover Publications, Inc. New York. 1051 pp. Kellogg, E. A. 1985. A biosystematic Study of the Poa secunda complex. Joum. Arnold Arbor. 66:201-242. Larsen, E. C. 1947. Photoperiodic responses of geographical strains of Andropogon scoparius. Bot. Gaz. 109:132-149. 95 McMillan, C. 1959. The role of ecotypic variation in the distribution of the central grassland of North America. Ecol. Monogr. 29:285-308. McMillan, C. 1964. Ecotypic differentiation within four North American prairie grasses. I. Morphological variation within transplanted community fractions. Amer. J. Bot. 51:1119- 1128. McMillan, C. 1965. Ecotypic differentiation within four North American prairie grasses. II. Behavioral variation within transplanted community fractions. Amer. J. Bot. 52:55-65. Metcalfe, C. R. 1960. Anatomy of the Monocotyledons. I. Gramineae. Oxford. Clarendon Press. Nees von Esenbeck, C. G. 1829. "Agrostologia Brasiliensis " in M artius’ Flora Brasiliensis. ( 2:208) 1829. Ostergren, G. and W. Heneen. 1962. A squash technique for chromosome morphological studies. Hereditas 48:332-341. Peattie, D. C. 1922. The Atlantic Coastal Plain element in the flora of the Great Lakes. Rhodora 25:57-70, 80-88. Pierce, G. J. 1974. The Coastal Plain floristic element in Michigan. Unpublished M.S., Western Michigan University. Raven, P. H. 1963. Amphitropical relationship in the floras of North and South America. Quart. Rev. Biol. 38(2): 151-177. ______and D. I. Axelrod. 1975. History of the flora and fauna of Latin America. Amer. Sci. 63(4):420-429. Riggins, R. 1977. A biosystematic study of the Sporobolus asper complex (Gramineae). Iowa State J. of Res.. 51(3):287-321. 96 Roberty, G. 1970. Monographique systematique des Andropogoneae du globe. Boissiera 9:216-239. Scribner. F. L. and C. R. Ball. 1898. Studies on American Grasses III. Miscellaneous notes and descriptions of new species. U.S.D.A. Bull 24 40-41. Stebbins, G. L. 1975. The role of polyploidy in the origins of North American grasslands. Taxon 24:91-106. 1975. 97
Table 4. Major taxonomic treatments of the members of the Schizachyrium scoparium complex. Not all taxa were treated by all authors.
HITCHCOCK (1950) GOULD (1967) ROBERTY (1960) (Andropogon)
A. scoparius S. scoparium S. scoparium var. frequens var. frequens var. neomexicanus var. neomexicanum var. septentrionalis var. septentrionalis var. virile A. littorale var. littorale A. divergens var. divergens A. maritimus S. maritimum var. m aritimum A. stolonifer S. stoloniferum var. stoloniferum A. cubensis S. cubense var. cubense A. gracilis S. gracile var. gracile A. sericatus S. sericatum A. rhizomatus S. rhizom atum A. niveus S. niveum 98
Table 5. Collections that provided chromosome counts for the taxa studied. Voucher specimens are on deposit at OS. Presented are state (country if not U.S.). county and voucher designation for each collection.
S. gracile-Florida: Dade. Fl-27. S. marihmum-Alabama: Baldwin, Ala-1; Florida: Bay. Fl-53; Franklin, Fl-50; Gulf. Fl-51. Fl-52. S. niveum-Florida: Highlands, FI-15. S. scoparium-Texas: Chambers, Tx-1; Oklahoma: Beaver. Ok-2; McCurtain, Ok-1; Georgia; Dekalb, Ga-4; Mentone, Ga-3; Kansas: Ford, Ks-2; Pratt, Ks-1; Seward, Ks-3; North Dakota: Grand Forks, Nd-1; Maine: Kennebec. Me-1; Androscoggin, Me-2; Canada: Ontario, Cn-1. S. scoparium var. divergens-Louisiana: Caddo, La-3; Morehouse, La-4, La-6; Natchitoches, La-2; Union, La-5; Vemon, La-1, La-7; Texas: Bowie, Tx-8; Cass. Tx-7; Newton, Tx-7 Tyler, Tx-6. S. scoparium var. littorale -Indiana; Porter. In-1; North Carolina: Carteret, Nc-1, Nc-2; New Hanover, Nc-3. Texas: Nueces, Tx-2; San Patricio, Tx-3; Refugio, Tx-4, Tx-5. S. scoparium var. polycladus -Alabama: Baldwin, Ala-2; Florida: Charlotte, Fl- 17; Citrus, Fl-46; Clay. Fl-1, Fl-25. Fl-29; Collier, FI-13. FI-14. Fl-28; Dade, Fl- 21, Fl-41; Desoto, Fl-16; Destin, Fl-54; Dixie, Fl-47; Franklin, Fl-49; Hernando, Fl-45; Highland. FI-18. Fl-39. Fl-40; Hillsborough. Fl-22; Indian River. Fl-8; Lake, Fl-36; Liberty, FI-19; Oceola, Fl-6, Fl-7; Okaloosa, Fl-55; Okechobee, Fi ll. Fl-38; Polk. Fl-44, Fl-37; Putnam. Fl-2, Fl-31; Seminole. Fl-5, Fl-34, Fl-35; St. Johns. Fl-26, Fl-30; St. Lucie, Fl-9, FI-10; Sumter. Fl-23; Taylor. Fl-20; Volusia. Fl-3, Fl-4, Fl-32, Fl-33; Wakulla, Fl-48; Walton. Fl-24, Fl-25; Georgia: Ocilla, Ga-1; Carlton, Ga-2. S. scoparium var. rhizomatum-Florida: Dade, Fl-42, Fl-43; Monroe, Fl-56, Fl- 57. S. sericatum-Florida: Monroe, FI-12. S. condensatum -Mexico: Vera Cruz. Stuessy. Crawford & Bruner, 1222, 1224, 1229, 1234, 1239, 1242, Oaxaca, Stuessy, Crawford & Bruner, 1272. Figure 12. Distribution of the Schizachyrium scoparium complex in North America.
99 1 00 a
CM
C Vtf Figure 13. Growth forms of members of the Schizachyrium scoparium complex.
101 littorale
scoparium rhizomatum niveum fundamental stoloniferum gracile organization cu b en se 8ericetum
i r
maritimum Figure 14. SEM photograph of the Schizachyrium caryopsis (S. scoparium), X I8. Figure 15. SEM photograph of the S. scoparium caryopsis, X50.
103
Figure 16. Typical leaf cross-section of the majority of the taxa of the Schizachyrium scoparium complex. Figure 17 Leaf cross-section of Schizachyriam gracile, and S. cubense. 1 06
16
eg- i t s a> © epidermis vascular, sderenchyma bundle
17 undifferentiated parenchyma Plate I. Chromosomes for the taxa of the Schizachyrium scoparium complex, a. Schizachyrium scoparium var. divergens. b. S. scoparium var. scoparium,.c. S. scoparium var. liltoralc. d. S. rhizomatum. e. S. maritimum. f. S. stoloniferum. (X450)
107 ’.*■ » . - 1 §4% ' S f t i *
a 1 £ s < ■ /
r«
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5 % h /> V ^'/K , «*» % S t> » . . ‘ » 'A * i • . ; « V * v * ‘ £»i%* £> * 5 ' j * V ' * \ \ i> 4'vx rf£ ~.ju #.* v »' ■’ *' - Plate II. Chromosomes for the taxa of the Schizachyrium scoparium complex, a. Schizachyrium niveum. b. S. condensatum. c. S. gracile. d. S. gracile (S. sericatum). (X450, except a. (X200)).
1 09 Plate II 110
M i # M r i m
* ^ v i>* a , x T f , * * £■*, **«► «*,
9 4- '** •* ^ -j* f ’-'• .
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*1 *-‘W Figure 18. Distribution of the Schizachyrium scoparium complex and the S. condensatum complex in the western Hemisphere. Locations of diploid S. condensatum indicated by the shaded circles. 1 12
S. scoparium
~n
S. condensatum Figure 19. Distribution of Schizachyrium gracile. Figure 20. Distribution of S. maritimum.
113 1 14
19
20 Figure 21. Features of Schizachyrium gracile. a. leaf sheath, blade cross-section, X7. b. Dispersal unit. XI5. c. Side-view of sessile spikelet, XI5.
115 21 116 Figure 22. Features of Schizachyrium maritimum, X5. a. Dorsal view of dispersal unit. b. Ventral view of dispersal unit. c. Dorsal view of rachis. d. Ventral view of rachis.
117 1 1 8
22 Figure 23. Features of Schizachyrium maritimum. a. Leaf sheath, ligule. collar. Note constricted collar, b. Plant base showing decumbent habit. XI.
119 1 20 23 Figure 24. Distribution of Schizachyrinm niveum.
121 1 22
24 Figure 25. Features of Schizachyrium niveum. a. Plant habit. X0.5. b. Dispersal unit. X5. c. Fertile lemma with awn, X5.
1 23
Figure 26. Distribution of Schizachyrium scoparium var. diveraens. Figure 27. Distribution of Schizachyrium scoparium var. littorale.
125 1 26
\A
27
—V
C" Figure 28. Features of Schizachyrium scoparium var. divergens.
127 1 28
28 Figure 29. Features of Schizachyrium scoparium var. littorale. a. Dispersal unit and fertile lemma, X5. b. Leaf sheath, ligule, blade, c. Basal portions of plant.
Figure 30. Distribution of Schizachyrium scoparium v ar. polyclaclus. Figure 31. Distribution of Schizachyrium scoparium var. rhizomatum.
131 1 32
V o ^ • •
31
m M
<9‘
& 0 Figure 32. Features of Schizachyrium scoparium var. polycladus. a. Range of development of pedicelled spikelets, X I5. b. Dispersal unit.
X5. c. Basal portion of plant (XI) showing rhizomes with intemodes not completely covered by leaves.
133 134 32
\
C Figure 33. Features of Schizachyrium scoparium var. rhizomatum. Basal portion of plant showing rhizome internodes completely covered by scale leaves, XI.
135 1 36 F ig u re 3 4 . Distribution of Schizachyrium cubense.
1 37 138
— — v SUMMARY Phcnctic, ecological, geographical and cytological data were used in this study in order to determine relationships among the taxa of the Schizachyrium scoparium complex in North America, construct a reliable taxonomy for the group and investigate the origins of S. scoparium. Examinations of chromosome numbers, the external morphology of caryopses, and leaf anatomy reveal little information that can be used to determine relationships among the species in the complex. Taxonomic interpretations rely upon the understanding of the variations in morphology, ecology and geography. Principal components, cluster and discriminant analyses suggest that specific and varietal limits that have been based upon one or few characters prove unreliable. Within S. scoparium, the majority of individual characters show no particular geographical correlation and the same character state can be found in many parts of this species' range. The phenetic analyses also demonstrate that four taxa, S. gracile, S. maritimum, S. niveum and S. scoparium are morphologically, ecologically and, for the most part, geographically distinct from one another. Taxonomically, these taxa are recognized at the species level. Within S. scoparium four varieties are recognized that are morphologically similar but ecologically and geographically quite distinct: vars. divergens, littorale, rhizomatum 139 140 The transfers of S. stoloniferum and S. rhizom atum to the varietal level under S. scoparium create two new taxonomic combinations, respectively S. scoparium var. rhizom atum and S. scoparium var. polycladus Study of the chromosomes finds no diploid taxa in this complex in the southeastern United States and dictates the formation of a new hypothesis to explain the origin of Schizachyrium scoparium. An original hypothesis, dependent upon southeastern diploids, must be modified. The phenetic analyses suggest that S. scoparium, a tetraploid, is closely related to S. condensatum a species of Central and South America. The presence of diploid and tetraploid populations in S. condensatum along with current interpretations of the probable phytogeographical history of this species, support a hypothesis that S. scoparium is derived from S. condensatum. APPENDIX A
Data matrix for all OTUs used in phenetic studies of Schizachyrium. Character codes are presented along the top margin and explanation of them is given in Table 1, Chapter I. OTUs are labelled as follows: Type specimen of S. scoparium var. ducis = Ductyp, type specimen of S. scoparium var. villosissimum = Villtyp, type specimen of S. scoparium var. septentrionalis Septyp,= type specimen of S. scoparium = Sctyp, type specimen of S. scoparium var. virile = Virtyp, S. scoparium var. divergens = Div —, S. scoparium var. Jrequens = Fre —, S. scoparium var. littorale = Lit—, S. scoparium var. neomexicanum Neo= —, S. scoparium var. septentrionalis Sep= —, S. scoparium var. virile = Vir S. gracile = GRA —, S. m aritim um = MAR S. niveum = NIV —. S. rhizomatum = RHI —, S. stoloniferum = STO —. S. condensatum var. gracilipes = GRP —, S. condensatum = CON —. S. plumigerum = PLU —, S. lactiflorus = LAC —, S. microstachyum (S. Am erican condensatum) = MIC
141 IDENTITY FERLEMLN LEMS IF ID FRTAWNLN PEDICLLN PEDSPKLN PSAVNLEN SPATHV© PEDUNCLN CALLUSLN CONIO 3 0 8 3 13 5 3 3 1 3 0 0 1 3 9 0 0 3 CON1 1 3 0 8 3 12 3 4 3 1 0 0 5 1 3 7.0 0 3 CON12 3 5 8 6 9 3 3 0 1 0 0 0 1 3 1 2 0 3 CON13 3 3 8 6 1 7 0 3 3 1 3 0 3 2 0 2 0 0 3 LACt 3 0 8 3 9 3 3 .0 1 3 0 .3 2 0 2 3 0 3 LAC2 3 0 8 3 8 0 3 .0 2 0 0 3 2 0 4 0 0 3 LAC3 3 0 8 3 too 3 0 1 3 1.0 2 0 2 0 0 3 LAC 4 2 3 8 0 7 3 3 0 1 0 1 3 2 3 9 .0 0 3 LACS 2 3 8 0 7 .3 2 3 1 3 0 3 1 3 1.2 0 3 LAC6 3 0 8 6 9 3 3 0 1 3 1.0 1.3 8 0 0 3 LAC7 3 0 9 7 9 .0 3 3 1 3 0 .3 1 3 3 .0 0 3 LAC8 2 3 8 0 8 3 3 .0 1 3 1.3 1 3 1.2 0 3 LAC9 3 0 8 3 1 0 0 3 .0 1 3 0 3 1 3 9 .0 0 3 LACIO 2 3 8 3 8 3 3 0 1 3 0 .0 1 3 8 0 0 3 LACI 1 2 0 8 3 10 3 4 0 1 3 0 0 1 3 2 3 0 3 LACt 2 8 0 6 3 9 3 S O 1 0 0 0 1.3 2 0 0 3 LACI 3 2 3 8 0 8 0 3# .0 1 0 1 5 1 3 1 0 0 3 LAC 14 3 0 8 3 9 0 3 0 2 0 0 3 1 3 2.0 0 3 LAC 13 3 0 8 3 9 3 3 0 1 3 0 .3 1.3 2 3 0 5 LAC 16 3 0 8 3 10 3 3 3 1 5 1.3 1 3 2 0 0 3 LAC 17 2 0 7 3 8 0 2 3 1 0 1 0 1 3 1 0 0 3 MCI 0 6 8 3 10 0 0 .3 0 6 3 3 6 3 3 3 1 0 M C 3 3 0 8 3 1 4 3 4 .3 1 3 0 3 2 0 2 3 0 .5 M C 3 3 5 8 6 16 U 6 0 2 0 0 .5 1 3 1 2 0 3 PLUI 4 0 8 7 14 0 8 0 3 3 4 0 1 5 7.0 1.0 PLU2 4 0 7 3 16 3 8 .0 4 0 3 3 1 3 1.8 1 0 PLU3 3 0 8 3 11 3 4 .0 2 0 0 .3 1 3 4 0 0 3 PLU4 3 0 8 0 1 0 5 3 .3 1.3 0 0 1.0 7 0 0 3 PLUS 3 3 7 3 1 4 4 .0 1 0 0 .0 1.5 1.7 0 3 PLU6 4 0 7 3 13 6 6 .3 3 3 1.3 1 3 2 6 1.0 PLU7 3 3 8 6 10 0 3 .0 1 0 0 .0 1 3 3 0 0 .7 PLUS 4 0 8 7 1 7 3 6 .0 2 .0 3 0 2 0 3 0 1.0 142 ©ENTITY FERLEMLN LEMBF© FRTAVNLN PEDCLLN PEDSPKLN PSAVNLEN SPATHV© PEDUNCLN CALLUSLN CONIO 3 .0 8 3 1 3 3 3 3 1 3 0 0 1 3 9 0 0 .3 c o m i 3 .0 8 3 1 2 3 4 3 1 0 0 3 1.3 7 .0 0 .3 CON 12 3 5 8 6 9 3 3 0 1 0 0 0 1 3 1 2 0 3 CON 13 3 3 8 6 17 0 3 3 1 3 0 .3 2 0 2 0 0 .3 LAC1 3.0 8 3 9 3 3 .0 1 3 0 .3 2 0 2 5 0 3 LAC2 3 .0 8 3 8 0 3 .0 2 0 0 .3 2 .0 4.0 0 3 LAC3 3 0 8 3 1 0 0 3 0 1.5 1.0 2 0 2 0 0 3 LAC 4 2 3 8 0 7 .3 3 .0 1 0 1.3 2 3 9 0 0 3 LACS 2 .3 8 0 7 .3 2 3 1 3 0 3 1 3 1.2 0 3 LAC6 3.0 8 6 9 .3 3 .0 1.3 1.0 1.3 8 0 0 5 LACT SO 9 7 9 0 3 .3 1 3 0 3 1.3 3.0 0 3 LAC0 2 3 8 0 8 3 3 .0 1 3 1.3 1 3 1 2 0 3 LAC9 3 0 8 3 1 0 .0 3 .0 1 3 0 3 1 3 9 .0 0 3 LACIO 2 3 8 3 8 3 3 .0 1 3 0 0 1 3 8 0 0 3 LA C 11 2 0 8 3 10 3 4 .0 1 3 0 0 1.3 2 3 0 3 LACI 2 1 0 • 1 9 .3 ' S O 1 0 0 .0 1 3 2 0 0 .3 LACI 3 2 3 8 0 8 0 34 .0 1 0 1.3 1 3 1.0 0 3 L A C I4 3 0 8 3 9 0 3 0 2 0 0 3 1 3 2 0 0 3 LAC IS 3 .0 8 3 9 3 3 0 1 3 0 .3 1 .3 2 3 0 .3 L A C I6 3 .0 8 3 1 0 3 3 .3 1 3 1.3 1 .5 2 0 0 3 LACI 7 2 0 7 3 8 0 2 .3 1 0 1.0 1 3 1 0 0 3 MCI 0 6 8 3 1 0 0 0 .3 0 6 3 3 6 3 3 3 1.0 M C 3 3 .0 8 3 1 4 3 4 .3 1 3 0 3 2 0 2 3 0 .3 M C 3 3.3 8 6 16 0 6 .0 2 0 0 .3 1.3 1 2 0 3 PLUI 4 .0 8 7 14 0 8 0 3 3 4 0 1 5 7 0 1.0 PLU2 4 0 7 3 1 6 3 8 .0 4 0 3 .3 1.3 1.8 1 0 PLU3 3 0 8 3 1 1 3 4 .0 2 0 0 5 1 3 4 .0 0 3 PLU4 3.0 8 0 1 0 5 3 .3 1.3 0 .0 1.0 7 .0 0 .3 PLUS 3.3 7 3 1.4 4 .0 1.0 0 .0 1.3 1.7 0 .3 PLU6 4.0 7 3 1 3 6 6 .3 3 .3 1.3 1 3 2 6 1.0 PLU7 3.3 8 6 to o 3 .0 1.0 0 .0 1 3 3 0 0 .7 PLUS 4 0 8 7 1 7 3 6 .0 2 0 3 .0 2 .0 3 .0 1.0 143 DENTITY RACEME XE RACEMLEN SPIKLNUM SPATHLEN RACHISLN RACHTPLN RACHP8LN RACPBPCL C ALSP8LN SESSPKLN C0N 10 0 2 3 6 19 4 3 0 3 2 3 6 0 0 3 3 0 CON! 1 0 3 3 6 2 0 3 0 0 3 2 3 5 0 0 3 3 0 CON! 2 1 44 9 17 3 0 0 3 1 3 6 0 0 0 3 0 CON! 3 1 4 3 9 2 3 3 3 0 3 2 3 7 3 0 7 3 0 LACI 0 11 3 0 11 3 0 0 4 2 0 3 0 0 3 4 0 LAC2 0 13 3 10 3 0 0 3 1 3 3 0 0 3 4 0 LAC3 0 13 3 12 3 0 0 3 2 0 3 0 0 3 4 5 LAC4 0 13 3 12 30 0 3 2 0 3 0 0 3 4 0 LACS 0 18 6 11 2 3 0 3 1 5 6 0 0 3 3 3 LAC6 0 13 6 13 3 0 0 3 2 0 3 0 0 3 4 0 LAC7 0 13 4 10 3 3 0 3 2 0 3 0 0 3 4 .0 LAC8 0 13 7 14 3 0 0 3 2 0 5 0 0 3 4 0 LAC9 0 2 2 8 13 3 0 1 0 2 0 7 3 0 3 4 0 LACIO 0 10 19 7 13 3 .0 0 3 3 0 0 3 4 0 LAC! 1 0 13 3 12 3 3 1 0 2 0 1 00 0 3 5 0 LACt 2 0 12 4 14 3 0 1 0 2 0 6 0 0 3 4 0 LAC13 0 15 7 10 3 0 0 3 2 0 3 0 1 0 3 5 LAC14 0 17 6 13 3 0 0 3 2 0 1 0 0 0 3 4 3 LAC15 0 12 3 13 3 0 0 3 1 3 100 0 3 4 3 LACI 6 0 16 6 10 3 3 0 3 1 3 3 0 0 3 4 3 LACI 7 0 34 7 14 2 3 0 5 2 0 3 0 0 3 4 0 MIC 1 0 3 3 9 4 6 3 0 1 0 0 3 60 1 0 9 5 MIC3 0 4 0 9 3 2 3 0 0 .3 2 3 7 3 0 3 4 0 MIC3 1 4 0 8 3 3 6 0 0 3 2 3 3 0 0 3 5 0 PLU1 0 3 0 3 2 6 7 3 0 3 2 3 100 1 0 8 5 PLU2 1 4 3 7 2 4 8 0 0 7 3 0 1 00 1 0 8 0 PLU3 1 2 6 7 13 33 0 3 3 0 7 3 0 3 4 0 PLU4 0 2 7 9 13 35 0 5 1 3 6 0 0 3 4 0 PLUS 0 32 9 23 40 0 5 20 30 0 3 4 0 PLU6 1 3 2 3 2 3 6 0 0 3 4 0 1 0 0 1 3 7 0 PLU7 0 29 5 3 0 3 3 0 3 2 3 1 0 0 1 0 3 5 PLU8 0 3 0 3 2 8 70 0 .3 3 3 7 3 1 5 5 0 DENTITY HEIGHT RHIZOMES BLADLENG BLADVDT SHEATLN LIGULLEN BLADE SHEATPU6 THROAPUB AURICPUB C 0N 10 8 9 0 1 6 3 5 .0 7 0 2 3 0 O O O CON11 7 3 0 14 3 3 .0 8 0 1 3 0 0 0 0 com 2 140 0 21 7 0 10 0 3 0 0 0 0 0 com 3 91 0 1 4 3 3 .0 1 0 0 1.3 0 0 0 0 LAC1 3 8 0 10 3 .3 7 0 1 3 0 0 0 0 LAC2 8 5 0 10 4 .0 3 0 1 3 0 0 0 0 . LAC3 9 9 0 12 3 .3 7 0 1 3 0 0 0 0 LAC4 91 0 12 4 3 10 3 2 3 0 0 0 0 LACS 6 2 0 12 4 3 8 3 2 0 . 0 0 0 0 LAC 6 5 3 0 2 3 4.0 1 1 3 2 3 0 0 0 0 LAC7 11 1 0 1 2 3 3 .0 7 3 1 3 0 0 0 0 LAC8 1 10 0 1 4 3 8 0 9 0 2 0 0 0 0 0 LAC9 73 0 9 0 8 0 6 3 2 0 0 0 0 0 LACIO 101 0 11 3 0 8 0 1 3 0 0 0 0 LACI 1 6 5 0 12 3 0 6 3 1 0 0 0 0 0 LACI 2 31 0 14 3 .0 7 .0 1 0 0 0 0 0 LACI 3 6 0 0 4 0 4 0 3 3 1 5 0 0 0 0 LACM 36 0 6 0 3 .3 3 0 1 3 0 0 0 0 LACI 3 9 5 0 13 4 0 8 3 1.3 0 0 0 0 LACI 6 8 0 0 1 1 3 3 .0 6 0 2 0 0 0 0 0 LACI 7 4 2 0 10 4 .0 5 3 2 0 0 0 0 0 MIC1 63 0 6 3 3 3 3 6 1.0 2 2 2 2 M C 3 167 0 12 4 0 7 .3 2 0 0 0 0 0 MIC3 7 6 0 8 0 2 .3 7 0 1 3 0 0 0 0 P IU I 4 3 0 10 2 0 3 3 2 0 0 0 0 0 PLU2 2 7 0 10 2 3 3 0 2 0 0 0 0 0 P IU 3 103 0 19 6 .0 8 0 2 0 0 0 0 0 PLU4 100 0 17 6 0 6 2 1 3 0 0 0 0 PLUS 5 0 0 7 0 3 .0 7 0 2 0 0 0 0 0 PLU6 33 0 8 7 3 0 5 .0 1 0 0 0 0 0 PLU7 30 0 12 2 3 3 .4 1.0 0 0 0 0 PLU8 33 0 7 0 2 3 SO 1 0 0 0 0 0 145 IDENTITY FERLEMLN LEMBFO FRTAVNLN PEDICLLN PEDSPKLN PSAVNLEN SPATHVO PEDONCLN CALLUSLN S T 0 4 0 3 0 2 0 8 .0 3 .0 2 0 0 0 1.3 3.7 0 3 ST041 6.3 3 0 1 2 0 6 .3 4 .0 2 0 2 .3 4 8 0 3 S T 0 4 2 4.3 2 2 9 3 4 .3 1 3 0 3 2 0 4 9 0 3 S T 0 4 3 4.0 2 3 6 3 3 .0 1.3 0 3 2 .3 6 7 0 .3 S T 0 4 4 4 3 2 2 9 0 3 .0 2 0 0 3 2 3 1.9 0 3 S T 0 4 3 3.0 3 0 1 0 0 4 0 3 0 1 3 2 3 6 0 0 3 S T 0 4 6 6 0 3 4 14 0 7 0 3 0 0 0 2 3 7 3 0 3 S T 0 4 7 4.3 3 4 9 0 4 3 2 0 1 0 2 3 3 0 0 3 S T 0 4 8 4 0 2 2 8 3 3 3 1 3 0 0 2 3 4 6 0 3 S T 0 4 9 3 5 31 1 1 0 5 .0 3 3 1 3 3 0 4 0 0 3 S T 0 3 0 5.0 3 0 13 0 3 .0 2 3 1 3 2 0 3 0 0 3 ST 051 3 0 3 3 9 0 4 .3 2 0 1.0 2 3 3 8 0 3 VILLTYP 4.3 3 3 8 .0 4 .0 1 3 1 3 2 3 4 2 0 3 VIR1 5.0 4 0 9 .3 3 .3 3 0 0 .3 1.3 4 7 0 5 VIR10 6 5 3 0 15 3 3 0 9 3 1 3 3 3 1.3 0 3 VIR2 3 0 3 0 11 0 3 .3 3 0 0 0 1.3 6 7 0 3 VIR3 6 3 3 0 13 0 4 .3 7 3 1 0 3 .3 3 0 0 .3 VIR4 6 0 2 3 11 3 3 0 7 0 0 0 2 3 3 9 0 3 VIR5 3.0 3 0 11 3 4 .3 3 3 0 3 2 .0 6 .0 0 3 VIR6 3 0 3 0 1 1 .3 4 .3 6 0 2 0 2 0 8.3 0 3 VIR7 6 0 16 9 3 3 .0 3 0 0 3 2 0 7.3 1 0 VIR8TYP 6 0 3 0 1 2 0 3 .0 0 6 1 0 1 3 3 0 0 3 VIR9 5.0 4 0 12 3 4 .3 4 3 2 0 2 0 7 0 0 3 CONI 4 0 4 3 13 0 3 0 2 0 2 .0 1.3 4 .0 0 3 C0N2 2 5 8 0 1 0 0 4 0 1 0 0 .0 3 .0 2 3 0 3 CON3 3 0 8 3 1 0 0 4 5 1.0 1 5 3 3 3 .3 0 3 C0N4 3 0 8 3 1 2 3 4 3 1 3 1 3 3 3 1.0 0 3 CONS 3 3 71 1 3 0 3 3 1 3 J O 3 0 2 3 0 3 CONS 3.0 8 3 1 3 0 4 3 1.3 0 0 1 0 2 .0 0 3 C0N7 3 0 6 0 11 0 4 .0 1 3 1 1 1 3 1.2 0 3 C0N8 3.0 6 0 11 0 4 .0 1 3 1 0 1 0 1.0 0 3 C0N9 4 0 8 7 13 3 4 3 1 3 3 .0 1 3 2.0 0 5 146 DENTITY RACEf- RACEMLEN SPIKLNUM SPATHLEN R ACHISLN RACHTPtN RACHPBIN RACPBPCL CALSPBIN SESSPKIN S T 0 4 0 1 3 6 10 3 7 3 0 1 0 2 .3 6 0 1 0 € 0 5T041 1 4 6 10 5 8 3 0 1 0 4 0 7 3 2 0 9 0 S T 0 4 2 2 6 3 9 4 2 4 3 1 0 2 0 7 3 1 0 6 3 S T 0 4 3 1 4 0 12 6 3 4 0 1.0 1 0 5 0 1 0 6 5 S T 0 4 4 0 2 5 6 2 3 4 0 1.0 2 3 6 0 1 5 7 3 S T 0 4 3 1 3 0 13 3 0 4 0 1.0 4 0 7 3 2 .0 7 0 S T 046 2 53 11 4 2 3 0 1.0 S 3 6 0 1 3 9 0 S T 0 4 7 0 37 10 3 6 3 3 1 0 3 0 6 0 1 0 6 0 S T 0 4 8 2 23 8 3 2 2 3 1 0 2 3 6 0 1 0 5 3 S T 0 4 9 0 6 2 13 8 3 7 0 0 3 3 3 6 0 0 3 6 5 S T 0 5 0 1 3 8 9 3 3 4 3 1 0 4 3 6 0 1 3 7 3 ST051 2 4 3 11 4 0 4 3 1 0 1 3 3 0 1 0 3 5 VLLTYP 1 3 9 12 3 3 3 3 0 3 2 3 3 0 1 3 6 0 VR1 1 4 3 12 4 3 4 0 1 0 2 .3 6 0 0 3 7 0 VIR10 0 4 2 8 3 3 3 0 1 0 2 0 3 0 1 3 10 3 VIR2 2 4 5 9 6 2 3 0 0 .3 2 0 6 0 1 0 7 0 V*?3 0 4 3 8 3 0 3 0 0 .3 2 3 4 0 1 0 9 0 VIR4 2 3 3 7 3 0 3 0 1.0 2 0 5 0 1 0 BO VIR3 2 4 2 9 31 3 0 1 0 1.3 3 0 1 0 7 0 VIR6 2 3 0 1 1 31 6 0 1.0 2 3 3 0 1 0 7 3 VIR7 2 3 7 7 4 7 5 0 1.0 1 5 3 0 1 0 8 0 VR8TY P 2 3 5 8 4 8 3 0 1.0 2 3 5 0 1 0 0 8 VIR9 2 3 0 13 4 3 2 3 1 0 2 3 3 0 1 0 7 0 C0N1 2 3 7 8 4 8 4 0 0 3 3 3 7 3 2 0 6 5 C0N2 1 3 0 6 3 3 4 0 0 3 3 0 6 0 0 3 4 5 C0N3 0 22 6 17 4 5 0 .3 2 3 6 0 0 3 4 0 C0N4 1 3 0 7 17 4 3 0 8 3 0 6 0 0 3 3 0 COM3 1 2 7 17 4 3 0 .8 3 0 6 0 0 .3 5 0 C0N6 1 3 0 7 2 0 3 0 0 5 3 .0 7 0 1 0 5 .0 C0N7 1 3 3 9 18 4 0 0 .5 2 3 3 0 0 3 4 0 C0N8 0 2 3 6 2 0 4 0 0 .3 2 3 6 0 0 3 6 0 C0N9 0 17 3 21 43 0 .3 2 0 100 1 0 6 0 1 47 IDENTITY HEIGHT RHIZOMES BIADLENGBLADVOT SHEATLN LIGULLEN BLADE SHEATPU8 THROAPUB AURIC PUB S T 0 4 0 118 I 2 2 3 0 0 9 7 1 3 0 I 1 I ST041 107 t 14 2 3 0 6 3 1.0 1 t 1 I S T 0 4 2 9 6 1 2 6 6 0 1 0 2 1.3 0 0 0 0 S T 0 4 3 6 9 t 1 3 9 3 0 8 0 1 0 0 0 0 0 S T 0 4 4 7 0 1 1 06 3 3 6 .3 1 0 0 1 1 1 S T 0 4 3 103 I 2 8 3 0 11 3 1 0 1 I 1 1 S T 0 4 6 130 1 3 9 9 .0 12 3 2 0 0 1 1 1 S T 0 4 7 9 3 1 13 5 4 .0 7 4 1 0 0 0 0 0 S T 0 4 8 126 1 14 3 3 .0 7 3 0 3 0 0 0 0 S T 0 4 9 120 1 2 3 6 0 12 8 2 0 0 I 1 1 S T 0 3 0 61 t 1 7 3 3 .3 6 3 0 3 0 0 0 0 ST031 2 1 0 1 2 9 6 0 1.1 1.0 0 0 0 0 VILLTYP 112 0 18 4 .3 0 .8 1.3 1 I I 1 VIR1 1 14 0 13 3 .0 3 .5 2 0 0 0 0 0 VIR10 8 8 0 4 3 4 .5 1 2 8 1 3 1 1 t 1 VIR2 8 0 0 17 3 3 3 7 3 2 0 1 1 1 1 VIR3 9 8 0 14 3 4 0 8 3 2 0 0 0 0 0 VIR4 6 6 0 3 0 3 4 .0 8 .7 1 3 0 0 0 0 VIR3 84 0 16 5 4 .0 3 8 1.0 1 1 1 1 VIR6 133 0 18 5 4 .0 8 0 1 3 0 0 0 0 VIR7 1 13 0 2 4 7 4 .3 9 8 1 0 1 1 I I V R 8T Y P 160 0 3 3 4 0 7 8 1 0 1 0 0 0 V R 9 100 0 21 5 .0 7 3 2 0 0 0 0 0 C0N1 125 0 2 6 4 .0 8 0 1 3 1 1 0 1 C0N2 1 03 0 2 3 5 8 0 1 2 3 2.3 0 0 0 0 C0N3 too 0 14 3 1 1 0 8 3 2 0 0 0 0 0 C0N4 141 0 1 7 3 7 5 9 3 1 3 0 0 0 0 COM3 127 0 17 1 0 0 11 3 2 0 0 0 0 0 C0N6 48 0 13 3 .0 7 .3 1 0 0 0 0 0 C0N7 1 26 0 4 5 8 .0 1 9 0 2 3 0 0 0 0 C0N8 73 0 21 3 .0 8 4 2 3 0 0 0 0 C0N9 1 24 0 21 3 .0 1 0 0 2.0 0 0 0 0 148 DENTITY FERLEMLN LEMBFD FRTAVNLN PEDCLLN PEDSPKLN PSAVNLEN SPATHVC PEDUNCLN CAUUSLN S T 08 4 0 3 8 1 1 3 4 0 1.5 0 3 3 0 4 8 0 3 S T 0 9 3 0 17 6 0 3 0 2 0 0 .3 3 3 2 3 0 3 STOIO 3 0 2 0 9 .0 4 .3 3 .5 1 0 3 0 4 3 0 3 ST01 1 3 0 30 t o o 4 .0 2 0 0 5 3 0 3 6 0 5 S T 0 I 2 3 .0 3 0 7 0 3 .3 2 0 1 3 3 3 6 7 0 3 S T 0 I 3 3 0 3 0 9 0 3 .0 2 3 0 3 3 0 4 0 0 3 STOt 4 3 3 3 6 12 0 4 0 2 3 2 0 3 0 4 .3 0 3 S T 0 1 3 4 3 4 3 1 0 0 4 0 2 3 0 0 4 0 1 0 0 3 S T 0 1 6 4 0 3 8 1 0 3 4 0 1 3 0 3 3 0 3 3 0 3 S T 0 1 7 6 3 31 11 0 6 3 3 .0 0 .5 4 0 4 3 0 3 S T 0 1 8 3 0 3 0 9 3 3 3 2 5 0 5 2 3 4 3 0 3 S T 0 I 9 4 0 3 8 7 0 3 3 2 0 0 .0 3 3 1.2 0 3 S T 0 2 0 4 0 3 8 9 3 4 0 1 3 0 0 0 .0 9 3 0 .5 ST021 5 3 3 6 11 3 4 .3 2 3 1 3 2 3 6 3 0 3 S T 0 2 2 3 0 3 0 1 1 3 3 3 3 0 1 3 2 3 4 0 0 3 3 T 0 2 3 3 0 2 0 12 3 4 .0 2 0 0 3 2 0 3 6 0 3 S T 0 2 4 3 0 2 0 1 1 0 3 0 2 3 1 0 2 0 4 8 0 5 S T 0 2 3 6 0 2 3 13 3 6 .0 2 3 0 .3 2 5 4 0 0 3 S T 0 2 6 3 3 18 7 0 3 0 2 3 0 0 2 3 4 2 0 .3 S T 0 2 7 3 0 2 0 10 0 4 .0 2 0 1 0 2 0 4 2 0 3 S T 0 2 8 5 .0 3 0 1 1 3 4 0 3 0 0 .3 2 0 3 .7 0 3 S T 0 2 9 4 .0 2 3 1 1 3 3 3 2 0 1 .0 2 5 3 .4 0 3 S T 0 3 0 3 .0 3 0 8 3 4 3 2 0 0 0 2 0 3 0 0 3 ST031 3 .0 3 0 7 3 3 .3 2 3 1 0 2 .0 2 7 0 3 S T 0 3 2 5 0 4 0 1 1 3 3 .3 2 3 1.0 2 0 1.7 0 .3 S T 0 3 3 5 3 18 10 3 3 .0 2 0 0 5 3 0 3 .0 0 3 S T 0 3 4 3 .0 4 0 6 .0 3 0 2 0 1 0 2 0 3 8 0 3 S T 0 3 3 4 3 3 0 7 0 3 0 2 3 1.0 2 3 3 .4 0 3 S T 0 3 6 5 0 2 0 9 .0 4 .3 2 3 1 3 2 3 4 2 0 3 S T 0 3 7 4 3 2 2 8 3 4 3 3 .0 0 3 2 0 4 0 0 3 S T 0 3 8 6 3 3 8 13 0 6 .3 3 3 0 3 2 0 9 3 0 3 S T 0 3 9 3 0 3 0 11 3 3 0 2 3 0 .3 7 0 3 6 0 3 149 DENTITY RACEMEXE RACEMLEN SPKLNUM SPATHLEN RACHISLN RACHTPLN RACHP8LN RACP8PCL CALSPBLN SESSPKLN S T 0 8 2 5 5 12 4 5 4 0 0 7 2 5 5 0 1 0 5 0 S T 0 9 t 3 0 9 3 5 3 5 0 .5 4 0 6 0 1 5 5 0 S T 0 1 0 2 4 5 13 4 5 5 0 1 0 3 5 6 0 1 5 8 0 ST011 t 3 6 10 3 6 4 0 1 0 2 5 6 0 1 5 6 5 S T 0 1 2 2 4 3 10 5 0 5 0 1 0 2 0 5 0 0 5 6 5 STO t 3 1 4 0 10 4 0 5 0 0 5 3 5 6 0 1 0 7 0 S T 0 1 4 t 4 2 9 4 7 4 5 1.0 4 0 5 0 1 5 8 5 S T 0 1 5 0 3 5 9 3 3 3 0 1 0 4 0 5 0 0 5 6 0 S 7 0 t 6 0 4 5 9 5 3 4 5 0 5 4 0 6 0 1 0 6 0 STO t 7 0 4 0 7 15 7 0 1.0 3 5 7 5 0 5 t o o S T 0 I 8 t 6 0 12 5 0 6 0 1 0 2 5 5 0 1 5 7 0 STO t 9 t 3 7 10 3 0 4 0 0 5 3 5 6 0 1 0 5 0 S T 0 2 0 0 4 0 10 3 0 35 0 5 4 0 7 5 2 0 5 5 ST 021 2 3 0 9 4 3 4 0 1 0 3 5 6 0 1 0 7 0 S T 0 2 2 1 3 5 9 3 8 4 5 1.0 3 5 5 0 1 5 8 0 S T 0 2 3 2 3 0 7 4 8 4 5 1.0 2 5 6 0 1 5 7 0 S T 0 2 4 2 3 5 9 4 3 3 5 1 0 3 0 5 0 1 0 8 0 S T 0 2 5 t 3 5 8 5 2 5 0 1 0 2 5 6 0 1 0 9 0 S T 0 2 6 t 4 2 14 5 8 5 0 1 0 1 5 6 0 1 0 7 0 S T 0 2 7 2 5 0 4 3 3 8 4 0 1.0 2 5 7 5 1 5 6 0 S T 0 2 8 1 3 4 8 3 8 3 0 1 0 2 0 5 0 0 5 7 0 S T 0 2 9 1 4 0 7 4 0 5 0 1 0 3 0 6 0 1 5 7 5 S T 0 3 0 2 3 5 9 3 2 4 0 0 5 2 5 5 0 1 0 7 0 S T 0 3 t 2 41 8 4 6 5 0 1 0 2 0 5 0 1 0 7 0 S T 0 3 2 1 3 0 8 3 5 4 0 1.0 2 0 5 0 1 0 7 0 S T 0 3 3 t 4 5 9 5 5 5 0 1.0 4 0 6 0 1 5 8 0 S T 0 3 4 2 2 5 8 3 5 4 0 1 0 2 5 5 0 0 5 7 0 S T 0 3 5 t 4 7 10 4 2 4 5 1 0 1 5 5 0 1 5 7 5 S T 0 3 6 2 3 3 9 3 8 4 0 1.0 2 0 6 0 0 5 7 0 S T 0 3 7 2 3 5 10 3 3 3 5 1 0 2 5 6 0 1 0 6 5 S T 0 3 8 2 5 0 9 5 2 5 5 1 0 3 5 8 0 2 0 9 0 S T 0 3 9 2 4 3 10 4 0 5 0 1.0 3 0 7 5 1 0 7 5 1 50 DENTITY HEIGHT RHIZOMES BLADLENO BLADVDT SHEATLN L CULLEN BLADEPUB SHEATPU8 THROAPUB AURIC PUB S T 0 8 181 1 23 3 .0 18 0 0 3 0 0 1 1 ' S T 0 9 12 0 3 0 3 0 120 2 0 0 0 0 0 S T 0 I 0 1 03 18 3 0 9 3 2 0 0 0 0 0 S T 0 1 I 118 2 0 3 .0 11 3 1 0 0 0 0 0 S T O I2 1 63 2 3 3 3 8 3 2 0 0 1 1 1 S T 0 1 3 9 3 20 6 0 100 2 0 1 1 1 1 S T 0 1 4 9 3 26 6 .0 1 3 0 2 0 1 1 0 0 S T 0 1 3 3 8 12 3 0 5 3 1.0 0 1 0 1 S T 0 1 6 9 0 2 2 3 0 3 7 1 3 0 0 0 0 S T 0 1 7 6 5 18 4 3 7 0 1 3 0 0 0 0 S T 0 1 8 1 3 0 2 2 2 3 9 7 1 0 0 0 0 0 S T 0 1 9 1 03 17 3 .3 9 3 1 0 0 0 0 0 S T 0 2 0 9 3 2 3 3 3 .3 9 0 1 0 0 0 0 0 ST021 9 3 2 0 6 3 7 3 2 3 0 0 0 0 S T 0 2 2 9 6 13 3 3 8 3 2 0 0 0 0 0 S T 0 2 3 1 1 2 19 3 0 6 7 2.0 0 1 1 1 S T 0 2 4 8 8 21 3 3 0 9 0 2 3 1 1 1 1 S T 0 2 3 1 3 3 2 7 .3 3 .0 1 3 0 1.3 0 1 1 1 S T 0 2 6 13 9 1 3 5 4 .3 1 1 3 1 3 0 0 0 0 S T 0 2 7 1 3 8 20 3 0 9 .0 1.0 1 1 1 1 S T 0 2 8 1 36 2 3 4 0 9 6 1 0 0 1 1 • 1 S T 0 2 9 7 3 1 1 6 3 3 6 5 1.0 0 0 0 0 S T 0 3 0 8 9 12 4 .0 7 9 1 0 0 0 0 0 ST031 3 8 10 3 4 0 3 2 1 0 0 1 1 1 S T 0 3 2 74 1 6 7 4 .0 9 8 1.3 0 0 0 0 S T 0 3 3 3 8 1 7 3 4 .0 10 0 1.3 0 0 0 0 S T 0 3 4 9 0 14 3 .0 6 4 1 3 0 0 0 0 S T 0 3 3 91 13 S O 7 3 1.0 0 0 0 0 S T 0 3 6 9 9 14 4 .0 8 0 1 0 0 0 0 0 S T 0 3 7 9 3 1 8 3 3 0 7 7 2 .0 0 0 0 0 S T 0 3 8 9 9 14 8 3 0 7 2 1.5 0 0 0 0 S T 0 3 9 71 18 6.0 2 0 0 0 0 0
12 3 1 51 DENTITY FERLEMLN LEMBFDFRTAVNLN PEDCLLN PEDSPKLN PSAVNLEN SPATHVC PEDL1NCLN CALLUSLN RH12 4 5 3 3 6 0 4 3 3 5 0 .2 3 0 4 0 1 0 RHI1 3 0 2 0 1 0 0 4 .0 2 3 0 2 2 8 4 3 1 0 RHI2 3 3 0 3 3 3 0 2 3 0 0 2 0 4 3 1 0 RHI3 3 0 3 4 4 3 3 0 2 0 0.1 2 0 4 3 1 0 RHI4 4 0 0 3 0 3 .5 2 0 0 1 2 0 3 2 1 C RHI3 4 .3 0 2 3 4 0 2 0 0 .0 2 0 3.0 1 0 RHI6 3.3 4 2 4 3 3 3 2 0 0.1 3 3 4.3 1 0 RHI7 3 .0 4 0 6 .3 3 0 2 3 0 2 2 3 3 3 1 0 RHI7 4 .0 3 3 6 .3 3 .0 2 3 0 0 3 0 3 7 0 3 RHI8 4 .3 4 8 7 3 4 .0 3 0 0 1 4 0 3 8 1 .0 RH19 5 .0 2 0 8 3 3 3 2 3 0 .2 3 0 3.2 1 0 S C 1 0 3 0 3 0 1 2 3 4 5 2 0 2 0 1 3 4 0 0 3 SCI 2 6 0 3 4 1 3 3 3 3 3 0 0 0 1 3 6 6 0 3 SCTYP 4 3 4 3 1 2 0 3 0 1 0 2 3 1 0 4 1 0 3 S E P 10 5 .0 2 0 8 .0 4 3 1 3 0 .5 1 3 2 7 0 3 SEP11 7 .0 2 9 1 3 0 6 5 6 0 4 0 2 0 8 3 0 3 SEP2 6 0 4 2 16 5 6 .0 4 0 1 0 2 0 3 8 0 5 SEP3 3 0 3 0 12 3 7 5 3 0 1.3 2 3 1.3 0 3 SEP4 5 3 4 3 1 2 3 6 .0 3 0 0 .3 1 0 6 0 0 3 SEP5 5 .0 3 0 11 3 6 0 2 3 0 3 2 0 7 3 0 3 SEP6 3 3 18 13 0 6 3 3 0 1 0 2 0 4 3 0 3 SEP7 5 0 3 0 15 0 7 .0 2 5 1 3 2 0 3 2 1 0 SEP8 6 0 3 0 1 2 3 5 .3 3 0 2 0 2 0 3 7 0 3 SEP9 30 30 133 3 0 3 0 2 3 3 0 3.3 0 3 SEPTYPE 3 0 4 0 1 2 0 6 .0 2 0 2 0 2 3 7 3 0 .3 ST01 5 0 10 9 0 3 .0 2 3 1 5 3 0 3 0 0 3 S T 0 2 4 0 2 3 1 0 0 3 .0 2 .0 1.0 1 3 3 8 0 3 S T 0 3 3 0 15 8 0 3 .0 3 0 2 .0 2 0 4.0 0 3 S T 0 4 5 .0 4 0 1 0 3 4 3 2 0 2 0 2 0 4 0 0 3 S T 0 5 5 .3 2 7 11 3 3 .0 . 3 0 0 .5 2 .0 3 2 0 3 S T 0 6 4.3 3 3 1 0 0 3 .0 2 0 0 .5 2 0 3 8 0 3 S T 0 7 5 0 3 0 11 0 4 .0 3 0 1 5 2 3 4 .3 0 5 152 )
DENTITY RACEMEXE RACEMLEN SPIKLNUfi SPATHLEN RACHISLN RACHTP1.N RACHP8LN RACPBPCL CALSPBLN s e s s p k l n RH12 2 3 3 13 4 3 4 0 1 0 0 3 3 0 1 3 6 3 RHI1 2 3 3 14 4 0 3 3 1 0 0 3 3 0 1.0 6 3 RHI2 1 2 0 6 3 0 3 0 1 0 0 2 6 0 1 0 4 0 RHI3 1 2 0 6 2 8 30 1 0 0 .2 6 0 1 0 3 0 RHI4 1 23 6 3 0 3 5 1 0 0 2 7 3 1 3 5 3 RHI3 1 2 3 7 3 2 3 3 0 3 0 .2 7 3 1 0 3 3 RH16 2 3 3 9 2 6 4 0 0 3 0 2 3 0 1 0 3 0 RHI7 2 3 5 12 3 3 4 3 0 3 0 2 3 0 1.0 6 0 RHI7 2 3 3 3 3 0 4 0 0 8 2 0 3 0 0 8 6 0 RHI8 1 3 3 3 4 0 3 0 1 0 0 3 73 1 0 6 0 RHI9 2 3 3 6 3 3 3 3 1 0 0 2 6 0 1 0 7 0 S C 10 2 3 3 8 3 6 3 0 2 3 2 0 7 3 0 3 7 0 SCI 2 2 4 0 8 4 3 3 0 1 0 4 0 3 0 1 0 3 3 SCTYP 1 4 8 8 4 3 4 0 0 .3 2 0 3 0 1 0 6 5 SEP 10 2 3 2 7 2 8 4 0 1 .0 2 3 6 0 1 0 7 0 S E P 1 1 2 6 3 9 6 0 6 0 1 0 4 3 6 0 2 0 1 0 SEP2 2 3 3 6 4 6 3 0 1 0 SO 6 0 1 0 8 5 SEP3 11 3 0 11 3 0 3 2 0 2 3 100 1 0 1 0 SEP4 2 3 8 6 3 0 6 5 1.3 2 3 5 4 1 0 7 3 SEP3 2 3 0 7 4 0 4 3 1 0 3 0 6 0 1 0 7 0 SEP6 2 2 8 6 3 2 3 3 1 0 0 3 4 0 0 3 8 0 SEP7 1 4 2 7 5 7 6 5 1.0 3 .0 3 0 1 3 9 0 SEP8 1 3 2 7 4 7 3 0 0 .3 2 3 7 0 1 0 7 3 SEP9 1 3 6 8 4 8 4 0 1 0 3 0 7 3 1 0 7 0 SEPTYPE 2 4 6 8 3 0 3 0 1.0 3 0 6 0 1 0 7 3 ST01 1 3 0 8 6 0 3 0 1 0 3 3 3 0 2 3 7 0 S T 02 2 3 0 9 3 2 3 0 1.0 3 0 7 3 1 0 6 0 S T 03 2 5 0 12 3 3 4 0 1 0 3 3 7 3 1 .0 7 0 ST 04 2 4 0 11 3 3 4 0 1 0 3 0 7 3 1 0 7 0 S T 03 2 3 0 8 4 2 3 3 1.0 3 .0 6 0 1 0 7 .0 S T 06 2 3 5 7 4 4 3 0 1 0 2 0 6 0 1 0 7 0 S T 07 2 2 8 6 3 5 33 1 0 3 0 3 0 1 3 7 3 1 53 IDENTITY HEIGHT RHIZOMES BLADCENO BLADVDT SHE ATLN LIGULLEN BLADEPUBSHEATPUB THROAPUB AURICPUB RH12 6 0 1 11 2 .3 3 3 1 0 0 0 0 0 RHI1 74 1 13 4 0 6 0 1 0 0 1 0 0 RHI2 6 0 1 14 5 1.3 3 .3 0 3 0 0 0 0 RHI3 6 0 1 1 3 6 2 0 5 4 0.3 0 0 0 0 RHI4 6 0 1 13 2 3 4 2 0 3 0 0 0 0 RHI3 6 3 1 17 2 .3 3 2 0 3 0 0 0 0 RHI6 7 3 1 9 5 2.3 3 7 0 3 0 0 0 0 RHI7 6 4 1 15 2.0 3 3 0 5 0 2 0 0 RHI7 6 5 1 13 3 3 4 6 1 0 0 0 0 0 RHI8 8 7 1 1 1 3 3 3 6 7 0 3 0 0 0 0 RHI9 8 5 1 19 3.0 1 1 4 0 3 0 0 0 0 s c t o 5 8 0 7 2 3.0 3 6 0 3 0 0 0 0 SC 12 6 9 0 1 8 5 3 0 7 3 1 0 0 0 0 0 SCTYP 9 0 0 14 4 .3 7 0 1 0 0 1 0 0 SEP 10 38 0 7 2 2 0 4 2 1 0 0 0 0 0 SEP1 1 9 0 0 19 3 4 3 9 0 1 0 0 0 0 0 SEP2 4 7 0 11 3 4 .0 7 0 1 0 0 0 0 0 SEP3 100 0 3 3 7 0 1 3 0 3 0 0 0 0 SEP4 4 7 0 9 3 1 3 3 4 0 3 0 0 0 0 SEP3 5 0 0 11 3 0 3 3 1 0 0 0 0 0 S t P 6 32 0 6 5 3 0 5 3 1 0 0 0 0 0 SEP7 61 0 19 5 3 3 7 3 1 0 0 0 0 0 SEP8 70 0 16 3 0 1 0 1.0 0 0 0 0 SEP9 4 0 0 1 5 3 2 .3 7 .0 1 0 0 0 0 0 SEPTYPE 6 0 0 19 4 0 1 0 0 1.3 0 0 0 0 ST01 112 1 2 3 3 0 14 0 1 0 1 1 1 1 S T 0 2 102 1 3 3 4 .0 6 2 0 3 0 1 0 0 S T 0 3 143 1 27 4 0 11 0 1 0 0 1 0 0 S T 04 175 1 26 3 0 9 0 1 3 0 1 0 0 STOT 194 1 3 3 5 3.0 13 7 2 3 0 1 1 1 S T 0 6 6 6 1 19 3 3 10 0 0 5 0 0 0 0 S T 0 7 75 1 2 2 3 4 .0 8 2 1 3 0 1 1 1 1 54 DENTITY FERLEMLN LEMBFD FRTAVNLN PEOICLLN PEDSPKLN PSAVNLEN SPATHVIO PEDUNCLN CALLUSLN MAR8 8 0 2 3 11 0 7 .0 3 3 0 .3 4 0 3 3 0 3 MAR9 7 0 21 11 3 6 .0 7 0 1 0 4 .0 3 7 0 5 MARIO 6 0 2 3 1 1 0 6 0 4 3 1 0 3 3 4 3 0 3 MAR1 1 6 3 31 1 2 3 3 3 3 .3 1.3 4 0 4 .0 0 3 MARI 2 7 0 13 1 0 0 6 .0 6 0 1.3 3 0 4 .6 0 3 M AR13 7 0 3 3 8 0 3 .3 4 3 0 3 4 0 3 0 0 3 HARM 6 3 2 3 1 3 0 6 .0 7 0 0 3 5 0 3 .0 0 5 M A RIS 7 0 2 9 10 0 6 0 6 5 1 3 5 0 3 3 0 3 MARI 6 7 3 3 6 8 .0 3 .0 3 3 0 .3 6 0 3 0 0 3 M AR17 7 .3 3 6 10 0 3 0 0 3 0 3 3 0 4 3 0 3 M AR18 6 0 6 3 10 0 3 0 6 .0 3 3 6 3 3 3 0 3 NEOI 6 0 5 0 13 0 3 0 4 3 1 0 3 0 4 .3 0 3 NEOIO 6 .0 3 0 1 3 0 6 0 3 0 0 0 2 3 2 3 0 3 NE02 7 3 7 5 14 0 3 .3 3 0 1.0 2 3 3 0 0 3 ME03 3 3 6 0 12 0 6 .0 3 0 1 0 2 0 3 0 0 3 NE04 3 0 4 0 14 0 3 .0 2 3 1 0 2 0 3 3 0 3 NE05 6 0 3 0 1 0 0 3 .0 3 3 1 0 2 0 3 3 0 3 NE06 3 3 4 3 8 3 3 .0 2 0 1 0 1 3 8 0 0 5 NE07 3 0 7 3 14 3 3 0 5 0 1 3 0 3 1 3 0 3 NE08 6 0 5 0 1 0 0 5 .0 4 .0 1.3 2 0 3 2 0 3 NE09 6 0 3 0 12 3 6 .3 3 0 1.3 1 3 9 0 0 3 NP/1 3 3 8 3 10 3 0 .3 0 2 0.1 2 0 4 .6 0 1 NIV2 4 3 9 0 11 3 6 .3 0 2 0 2 2 0 2 2 1 0 N r/3 4 0 8 7 12 5 0 3 0 2 0 .2 2 3 4 0 1 0 NT/4 4 0 8 7 12 3 0 3 0 2 0 2 2 3 4 0 1 0 N r/3 4 0 8 7 1 2 0 6 3 1 0 0 1 2 3 2 0 1 0 N r/6 4 0 8 7 11 0 3 0 1 3 0.1 3 0 2 6 1 0 NW7 4 3 8 6 1 2 0 3 0 2 0 0.1 3 5 3 0 1.0 N r/8 4 0 2 3 13 0 5 0 2 3 4 0 2 0 2 8 1 0 N r/9 3 0 9 0 11 0 3 .0 1 3 1 3 1 5 2 3 1 0 RHIO 3 5 3 8 8 3 4 .3 2 3 0 2 2 3 4 8 1 0 RH11 3 0 4 0 9 0 4 3 3 3 0.1 2 .3 3 0 1 0 1 55 DENTITY RACEf' RACEMLEN SPIKLNUM SPATHLEN RACHISIN RACHTPLN RACHP8LN RACPBPCL CALSPBIN SESSPKLN MAR8 1 2 5 4 3 2 5 5 1 0 3 3 5 0 1 0 1 1 0 M AR9 1 4 0 8 4 0 4 5 1 .5 3 5 44 0 5 10 0 M AR10 1 4 4 9 4 8 4 0 1 .5 3 0 7 3 0 5 9 0 MARI 1 1 3 8 7 4 3 4 3 1.0 3 5 7 8 0 3 9 0 MARI 2 1 5 0 8 5 0 5 0 1.3 3 3 7 0 1 0 0 10 3 MARI 3 1 4 5 7 5 5 4 0 1.3 4 3 6 0 0 3 9 0 MARI 4 1 4 3 10 5 7 4 0 1.0 4 0 7 3 1 0 9 5 M AR15 1 5 3 10 5 5 4 0 1 0 4 .3 5 0 0 0 9 3 MARI 6 1 4 9 9 5 3 4 5 1.3 6 0 6 0 1 0 9 0 M ARI 7 1 3 5 8 4 6 4 0 1 0 3 3 3 0 1 0 10 0 MARI 8 0 3 5 9 4 6 5 0 1 0 4 3 6 0 1 0 9 5 NEOI 2 6 5 12 6 5 5 0 1 0 4 0 100 1 0 8 0 NEOIO 2 3 8 7 3 5 6 0 1 0 2 0 100 i 0 8 0 NE02 1 3 8 7 3 7 5 5 1 0 2 0 100 1 0 8 0 NE03 2 4 7 9 4 0 5 0 1 0 2 3 8 3 1 0 7 3 ME 04 2 4 5 9 5*0 5 0 1 0 3 3 100 1 0 1 0 0 NE05 1 5 5 11 5 0 5 0 1 .0 3 0 100 1 5 7 0 NE06 2 4 0 8 4 0 4 0 1 0 3 3 8 3 1 0 6 5 NEC7 1 4 5 10 5 5 5 0 0 3 3 .0 100 1 0 7 0 NE08 2 3 3 6 3 0 5 0 1 0 4 0 100 1 0 8 0 NE09 I 5 0 6 6 0 6 5 0 3 2 .0 1 00 1 0 9 3 NIV1 I 3 8 8 3 6 5 0 1 0 0 .2 100 1 0 3 .3 NIV2 ' 2 4 5 6 3 5 7 0 0 3 0 2 100 1.0 6 0 NIV3 2 37 7 4 0 5 0 1 0 0 2 100 1.0 6 3 NIV4 0 3 0 5 3 2 5 0 1 0 0 2 100 1 0 6 5 NIV3 0 3 0 6 2 5 7 0 0 5 0 2 100 1 0 6 0 NIV6 0 30 6 3 2 5 0 1 0 0 2 100 1 0 5 0 NIV7 0 3 0 6 3 2 6 0 0 3 0 3 100 1 0 6 .0 NIV8 1 2 6 7 4 0 3 0 0 3 6 0 100 0 3 6 3 NIV9 1 3 5 6 2 9 5 5 1 0 2 3 100 0 3 6 0 RHIO 2 3 7 8 3 4 4 5 1 0 0 .2 6 0 1 0 6 0 RH1 1 2 4 0 10 4 0 4 0 1 0 0 .3 6 0 1 5 7 5 1 56 DENTITY HEIGHT RHIZOMES BIADLENGBLADVDT SHEATLN LK3U.LEN BIAOEPUB SHEATPUB THROAPUB AURICPU8 MAR8 3 6 1 143 4 .0 8 8 1 .0 2 2 2 2 MAR9 7 8 I 7 6 3 .0 3 .7 0 .3 2 2 2 2 MARIO 4 7 1 11 4 .0 3 .0 0 3 2 2 2 2 MARI t 4 0 I 7 3 4 0 3 9 0 ,7 2 2 2 2 M AR12 4 8 t 9 8 4 .0 4 0 0 3 2 2 2 2 MARI 3 71 1 1 3 3 4 .0 6 8 1 0 2 2 2 2 MARI 4 6 8 t 12 2 4 .0 3 9 1 0 2 2 2 2 M A R IS 61 1 13 4 4 .0 3 7 0 7 2 2 2 2 MARI 6 4 9 1 13 3 4 .3 3 .6 0 .7 2 2 2 2 MARI 7 3 7 1 9 0 4 .3 2 8 0 3 2 2 2 2 MARI 8 6 3 1 6 3 3 3 3 6 1.0 2 2 2 2 NE01 9 0 0 8 3 4 .0 3 0 1.0 0 0 0 0 NEOIO 6 0 0 14 3 .0 5 0 1 0 0 0 0 0 NE02 6 3 0 13 4 0 3 0 1.0 0 0 0 0 NE03 1 30 0 1 3 3 4 .3 6 3 2 0 0 0 0 0 NE04 7 0 0 18 3 .0 6 0 1 3 0 0 0 0 HE 05 8 5 0 14 4 0 7.0 1 0 0 0 0 0 ME06 6 3 0 14 3 .0 3 0 1 0 0 0 0 0 NE07 6 5 0 12 3 .0 6 0 1 0 0 0 0 0 NE08 9 7 0 3 6 3 .0 8 0 1.0 0 0 0 0 NE09 6 9 0 14 3 2 .0 7 3 0 5 0 0 0 0 NM 7 3 0 6 2 0 3 4 1.0 0 0 0 0 NW2 7 0 0 8 3 .5 3 7 1.0 0 0 0 0 N N 3 6 2 0 7 2 .0 4 5 1.0 0 0 0 0 N V 4 7 7 0 7 3 3 .3 4 3 1.0 0 0 0 0 NK'S 6 5 0 10 3 .3 3 0 0 3 0 0 0 0 N V 6 3 8 0 7 6 4 .0 3 3 0 .3 0 0 0 0 N N 7 6 9 0 9 3 .0 4 .3 0 3 0 0 0 0 NW8 8 8 0 2 7 4 .0 8 0 1.0 1 1 1 1 N N 9 4 9 0 9 0 4 .0 4 0 1.0 0 0 0 0 RH10 6 0 t 19 3 0 6 2 0 .3 0 0 0 0 RH11 6 0 I 17 3 .0 6 3 0 3 0 0 0 0 1 57 ©ENTITY FERLEM1N LE M B F© FRTAVNLN PEDICLLN PEDSPKLN PSAVNLEN SPATHVID PEDUNClN CAUUSIN GRA12 3 0 3 0 11 3 3 0 0 5 0 0 2 3 4 3 1 0 SERI 2 4 4 3 11 0 3 .0 1 0 0 .0 0 2 3 0 1 0 SER2 3 0 3 4 1 2 0 4 .3 2 0 0 0 0 2 3 .7 1 0 SER3 3 0 3 0 1 3 0 4 0 2 0 0 2 0 2 3.3 1 0 SERA 3 0 3 4 11 0 3 0 1 0 0 0 2 3 3 0 0 3 SER3 3 0 3 3 1 4 0 3 0 2 0 0 0 2 3 4 4 0 .3 SER6 3.3 4 0 14 3 4 3 2 0 0 0 2 3 3 .3 0 3 SER7 4 3 3 0 1 3 0 3 3 2 0 0 0 2 0 4 3 0 3 SER8 3 0 3 3 1 2 0 4 .3 1 0 0 0 2 3 3.0 0 3 GRP1 3 0 8 3 7 3 3 .0 1 0 0 0 3 0 3 0 3 0 GRP2 3.0 8 3 6 5 3 .0 1.3 0 0 2 3 2 3 0 3 GRP3 3 0 8 3 6 3 3 0 1 3 0 0 2 3 2 0 0 3 GRP4 3 0 8 3 7 3 3 0 1 0 0 0 3 0 3 0 0 3 GRP5 3 0 8 3 3 .0 3 .0 2 0 0 .3 2 0 2 .0 OS GRP6 3 0 8 3 7 .0 3 0 1.3 0 0 3 0 4 0 0 3 GRP7 3.0 8 3 8 3 2 .3 1 0 0 0 3 0 4 0 C 3 L IT 1 7 0 14 1 0 0 7 .3 4 0 1 0 3 0 6 5 0 3 LIT2 6 3 31 1 3 4 6 0 4 0 1 3 3 3 4.3 0 3 LIT3 5 0 2 0 1 1 3 3 .3 2 3 3 0 3 0 3 3 0 .3 LIT4 6 0 3 3 1 3 5 6 0 2 3 3 .3 4 0 3.3 0 .3 LIT5 6 0 3 0 1 6 6 0 3 0 2 0 2 0 6 2 0 3 IIT 6 6 3 3 9 1 3 0 3 0 2 0 1.0 2 0 6 2 0 3 LIT7 6 0 4 0 1 7 0 6 0 3 0 2 0 2 0 3.2 0 3 LIT8 6 0 3 4 1 6 3 6 .0 3 0 1 0 2 0 6 0 0 3 LIT8 3.0 3 0 1 2 0 3 .0 3 0 2 0 2 0 4 .7 0 3 LIT9 6 0 4 2 1 3 0 6 0 3 3 1 3 2 3 6 .0 0 3 MAR2 B.O 2 3 9 .0 6 0 6 3 1 3 4 0 5 .0 0 3 MAR3 7.3 4 0 1 2 3 3 .3 8 3 0 0 5 0 4 .2 0 3 MAR4 7.3 3 4 1 0 0 3 .0 7 0 0 0 4 .3 4 .0 0 3 MARS 7.3 4 0 9 3 . 6 3 8 0 0 .0 4 3 3.0 0 3 MAR6 7 3 3 4 9 0 3 .3 7 0 1 0 3 3 4 0 0 .3 MAR7 7 0 3 4 1 2 0 6 3 6 0 0 3 5 2 3 .0 0 3 1 58 IDENTITY RACEMEXE RACEMLEN SPIKLNUMSPATHLENRACHISLN RACHTPIN RACHP8LN RACPBPCl CALSPBlN SESSPXIN GRA12 1 3 5 9 5 6 3 0 0 5 0 6 100 2 3 4 5 SERI 1 3 0 7 4 2 2 5 0 5 0 5 100 2 3 4 0 SER2 1 3 5 11 4 2 3 0 0 3 0 6 100 2 3 6 0 SER3 1 2 5 7 6 2 3 0 1 0 0 6 100 2 0 5 5 SER4 1 3 5 7 41 2 5 0 5 3 0 100 2 3 6 0 SER5 1 3 0 8 4 2 3 0 0 3 3 3 100 2 3 4 3 SER6 1 3 5 7 6 0 3 0 0 3 3 3 100 2 3 5 0 SER7 1 2 5 9 5 6 35 0 3 6 3 100 2 0 5 5 SER8 1 3 7 11 4 3 30 0 3 3 3 100 2 0 4 3 GRP1 0 19 5 10 3 0 0 3 1 3 4 0 2 0 4 5 GPP 2 0 2 0 7 18 3 0 1 0 1 3 5 0 0 3 5 0 GRP3 0 2 0 8 16 2 5 1 0 1 0 3 5 0 .0 4 0 GRP4 0 16 7 18 3 0 1 0 1.3 5 0 0 0 4 5 GRP5 0 2 2 7 21 3 0 1 0 1 3 5 0 0 3 4 5 GRP6 0 15 5 13 3 0 0 3 1 3 5 0 0 3 3 0 GRP 7 0 2 3 8 17 3 0 0 3 1.5 5 0 0 3 4 3 LIT1 1 7 7 11 1 07 7 0 1 0 2 3 71 1 0 9 0 LIT2 2 4 0 7 5 7 5 5 1 0 6 0 7 5 1 3 10 0 LIT3 2 3 4 7 4 7 4 0 1 0 3 3 7 5 1 3 8 0 LIT4 0 3 0 6 4 4 4 5 1 0 6 0 6 0 l 0 8 0 LIT5 0 3 5 7 4 7 5 0 1 0 3 0 75 1 3 8 5 LIT6 1 4 5 8 4 5 5 0 0 3 4 0 100 1.0 7 .0 LIT7 1 3 5 8 4 7 4 0 1 0 3 0 6 0 2 3 9 0 LIT8 1 3 0 5 5 5 7 0 1 0 4 0 100 1 0 10 0 LIT8 1 3 5 7 4 7 4 0 1.0 3 5 100 1.0 6 0 LIT9 4 8 9 5 0 5 0 1.0 3 3 100 1 0 8 .3 MAR2 1 41 7 5 0 5 0 1 3 2 3 6 0 0 5 1 0 3 M AR3 1 3 6 6 4 4 5 0 1 0 3 3 7 5 7 0 9 2 M AR4 1 4 0 8 41 5 0 1 3 3 3 7 5 7 0 9 2 M AR5 1 6 3 11 6 6 4 5 1 3 4 0 6 0 7 0 10 0 MAR6 1 4 2 10 4 5 4 5 1 3 3 0 6 0 0 3 9 3 MAR 7 1 5 4 1 1 5 2 4 5 1.0 4 3 6 0 1 0 9 0 1 59 IDENTITY HEIGHT RHIZOMES BLADLENG BLADV SHEATLN LIGULLEN BLADE SHEATPUB THROAPue AURICPUB GRAI2 6 3 0 2 3 1 0 3 2 0 3 1 0 1 I SERI 6 0 0 2 8 1.0 4 3 0 .5 1 0 1 I SER2 6 3 0 2 4 1 0 3 4 0 .3 1 0 1 I SER3 4 3 0 2 2 1 0 4 .3 0 .3 1 0 I I SERA 6 3 0 2 9 1 0 3 3 0 3 1 0 I I SER3 4 0 0 2 3 1 0 4 .7 0 .5 1 0 1 I SER6 3 9 0 2 9 1 0 3 0 0 3 1 0 I I SER7 4 2 0 2 2 1 0 4 3 0 3 1 0 1 I SER8 3 3 0 2 4 1 0 4 7 0 3 1 0 1 1 GRP1 31 0 3 3 3 0 2 2 1 3 0 0 0 0 GRP2 9 0 0 3 0 2 0 3 3 1 0 0 0 0 0 GRP 3 1 12 0 7 3 4 .0 3 .0 2 0 0 0 0 0 GRP4 1 00 0 6 0 2 3 3 3 1 0 0 0 0 0 GRP3 7 2 0 7 0 3 .0 3 .4 1 3 0 0 0 0 GRP6 1 3 0 0 5 5 3 .0 3 3 1 0 0 0 0 0 GPP7 4 3 0 8 5 3 0 5 0 2 0 0 0 0 0 L IT 1 1 5 6 0 1 4 3 6 0 8 7 1 5 2 2 2 2 LIT2 7 8 0 21 6 3 1 0 3 2 .0 0 2 0 0 LIT3 6 8 0 2 6 3 3 0 13 0 2 .0 0 2 0 0 LIT4 7 5 0 3 4 3 6 0 10 0 2 0 2 2 2 2 LfTS 3 2 0 12 3 .0 8 .0 2 0 2 2 2 2 LIT6 3 9 0 9 0 3 .0 3 0 1 0 2 2 2 2 LIT7 9 0 0 2 2 4 0 9 0 1 0 0 0 0 0 LIT8 3 7 0 19 4 0 12 0 1 0 0 2 2 2 Lrre 3 7 0 11 3 3 0 8 7 2 0 0 0 0 0 LIT9 8 2 0 24 4 .0 11 3 2 .0 2 2 2 2 MAR2 71 1 8 3 4 0 7 0 0 .3 2 2 2 2 MAR3 5 8 12 4 .0 3 9 1.0 2 2 2 2 MAR4 6 3 9 3 4 3 6 1 1.0 2 2 2 2 MARS 6 5 16 3 4 .0 7 9 1.0 2 2 2 2 MAR6 6 0 11 4 .3 3 0 0 7 2 2 2 2 MAR7 7 7 134 4 .0 6 3 0 .7 2 2 2 2 1 60 IDENTITY FERLEMLN LEMBIFID FRTAVNLN PEDCLLN PEDSPKLNPSAVNLEN SPATHV© PEDONCLN CALLUSLN DIV1 5 0 5 0 14 0 6 0 2 .0 2 3 1 5 6 3 0 3 DIV2 5 5 3 0 9 0 5 0 4 3 0 3 2 5 4 0 0 5 DIV3 4 5 3 3 1 3 5 3 .0 3 3 0 3 3 0 3 2 0 5 DIV4 6 0 2 5 9 0 4 5 3 3 1.0 3 0 4 2 0 5 DIV3 5 0 3 0 12 0 6 .3 2 3 0 5 2.0 2 0 0 3 DIV6 5 0 3 0 1 1 0 5 0 1 5 1 3 2 0 2 5 0 .3 DIV7 6 0 3 4 1 1 0 5 0 3 5 1.3 1 5 8.3 0 3 DIV8 6 0 20 13 0 3 .3 2 0 0 3 1 5 8 0 0 5 DIV9 5 5 17 1 0 5 4 .3 0 3 1 0 1 5 4,3 0 3 DIVIO 6 0 3 3 9 5 3 .0 7 0 1 0 2 0 6 2 0 3 DUCTYP 5 0 4 0 11 5 4 .0 3 0 1 0 2 5 7 2 0 3 FRE1 4 0 2 5 1 3 0 4 .3 1 3 0 .0 2 5 4 2 0 3 FRE2 6 0 4 0 17 0 6 3 3 0 3 0 1 5 7.0 0 .3 FRE3 4 5 4 4 1 5 5 5 .0 3 0 1 3 2 5 3 8 0 3 FRE4 6 0 3 0 1 3 0 5 .0 3 0 2 0 3 0 6 3 0 3 FRE5 3 0 4 0 1 0 0 3 0 2 3 0 0 3 5 2 2 0 3 FRE6 3.5 3 6 1 2 5 6 0 2 3 0 0 20 5 2 0 5 FRE7 3 0 4 0 12 0 3 .3 2 0 0 0 20 6 5 0 5 FPE8 3.0 6 0 14 5 3 .0 2 5 0 3 1 5 6 0 0 .5 FRE9 3 0 3 0 1 4 5 3 .0 1 3 1.5 2 0 6 3 0 5 FRE10 4.5 4 5 9 0 3 .0 2 0 2 0 1.5 4.4 0 5 GRA1 3 0 3 0 19 0 4 .3 3 0 0 0 1 5 3 7 1 0 0R A 2 3 0 5 0 11 0 4 0 1 0 0 0 20 3 7 1 0 GRA3 3 5 4 3 17 0 4 .3 2 3 0 .0 2 5 6 0 1 0 GRA4 2 5 4 3 14 5 4 0 2 0 0.1 3 0 3 4 1 0 GRA5 3 0 5 0 1 4 5 3 5 2 .0 0 2 3 .0 4.7 1 0 GRA6 2 5 4 0 1 0 5 3 .3 1 3 0 2 3 0 9 2 1 0 GRA7 4 0 5 0 1 7 5 4 .3 2 0 0 2 3 5 9 1 1 0 GRA8 3 0 5 0 1 5 0 4 0 1 5 0 3 3 0 7 4 1 0 0R A 9 S O 34 12 0 3 .3 1 0 0 0 2 0 3 2 1 0 GRAIO 2 5 20 14 5 3 5 2 0 0 2 3 0 4 6 1 0 GRAI 1 S O 3 4 14 0 3 0 1.3 0,1 3 5 7.7 1 0 CENT IT Y RACE11EXE RACEMLEN SPIKLNUMSPATHLEN RACHISLN R ACHTPLN RACHP8LN RACPBPCL CALSP8LN SESSPKLN DIV1 1 4 7 12 6 2 3 3 1 0 2 0 60 1 0 7 .0 DIV2 1 4 0 9 4 8 4 0 1.0 2 0 30 1.0 7 3 DIV3 2 4 0 9 4 0 3 0 1.0 3 0 3 0 1 .0 8 0 D1V4 2 3 0 11 41 4 0 1.0 1 3 5 0 1 0 8 0 DIV3 2 31 7 2 2 3 3 1 .0 2 0 5 0 1 0 7 C D1V6 1 3 0 ■7 4 0 4 3 0 3 4 3 7 3 1 0 7 0 DIV7 2 5 0 12 3 3 4 0 1 0 2 0 3 0 1 0 6 0 DIV8 2 4 6 7 4 6 6 0 0 3 3 0 7 3 1 .0 9 0 D1V9 2 3 3 11 4 3 3 5 1 0 2 0 3 0 1 0 7 0 DIVIO 2 4 0 9 4 3 3 0 1 0 2 0 6 0 1 0 8 0 DUCTYP 1 4 6 11 71 4 0 1 0 2 3 7 3 1 3 7 0 FRE1 2 3 7 13 3 7 3 3 0 3 3 0 7 3 1 0 6 0 FRE2 2 3 3 6 4 4 3 0 0 .8 4 0 100 1 0 9 0 FRE3 0 3 4 8 4 2 3 3 0 .8 3 3 3 0 1 0 6 3 FRE4 2 3 7 8 3 0 4 0 1 0 3 3 7 3 2 3 7 3 FRE3 0 3 6 9 4 2 4 0 1 0 3 0 7 3 1 0 8 0 FRE6 1 36 7 4 3 4 3 1 0 3 3 6 0 1 0 8 0 FRE7 1 4 3 9 4 8 4 0 1.0 2 3 7 3 1 0 6 0 FRE8 1 4 0 8 4 3 2 3 1 0 3 0 6 0 1 0 6 0 FRE9 2 3 0 9 3 6 6 0 0 3 2 0 3 0 1.0 6 3 FRE10 1 3 3 9 4 6 3 0 0 5 2 3 7 3 1 0 7 0 GRA1 0 3 0 6 6 0 3 0 1 0 0 4 100 2 3 7 0 0RA 2 2 3 4 9 31 4 0 0 5 0 6 100 1 0 3 0 GRA3 1 4 0 1 1 3 2 4 0 0 3 0 6 100 2 3 3 0 GRA4 0 4 3 13 6 2 3 0 0 .3 0 6 100 2 3 4 0 GRA3 0 4 3 13 3 2 30 0 3 0 6 100 2 0 3 0 GRA6 2 4 2 12 8 0 30 0 3 0 7 100 2 0 4 0 GRA7 2 4 3 12 6 2 4 0 0 3 0 3 100 2 3 6 0 GRA8 1 4 3 13 6 6 30 0 3 0 6 100 2 0 4 3 GRA9 1 3 3 13 3 0 2 3 0 .3 0 3 100 2 3 4 0 GRA10 1 4 2 12 6 2 3 0 0 3 0 6 100 2 0 4 3 GRA 11 1 4 8 1 2 3 6 40 0 .3 0 3 100 2 3 4 0 1 62 IDENTITY HEIGHT RHIZOMES BLADLENO B L A D V 0T SHEATLN LICULLEN BLADE SHEATPU8 THR0APU8 AURICPUB DIV1 5 7 0 8 5 3 3 6 2 1.0 0 0 0 0 DIV2 182 0 21 4 .0 8 3 2 .0 1 t 1 t DIV3 74 0 1 8 3 4 .0 7 0 2 0 1 1 t I DIV4 1 35 0 21 4 .0 8 3 2 .0 1 I 1 1 DIV3 8 7 0 19 3 0 9 .0 2 0 1 I 1 I DIV6 6 5 0 2 0 4 0 8 3 1 0 0 0 0 0 DIV7 109 0 24 4 3 .0 8 0 2 0 1 I 1 I DIV8 75 0 14 3 3 0 3 .6 1.0 0 0 0 0 o rv 9 110 0 2 9 3 0 7 0 1 0 0 0 0 0 DIV10 9 0 0 13 3 .0 5 3 1 0 1 1 t 1 DUCTYP 4 2 0 5 6 4 0 3 3 1 0 0 0 0 0 FREI 5 7 0 1 3 3 3 0 4 6 1 0 0 0 0 0 FPE2 8 2 0 16 3 3 8 4 1 0 0 0 0 0 FRE3 6 7 0 13 7 3 .0 6 2 1 0 0 0 0 0 FPE4 6 7 0 2 0 2 3 4 6 0 3 0 0 0 0 FRE5 5 2 0 18 4 0 9 0 1 0 0 0 0 0 FRE6 7 2 0 1 8 3 4 0 0 9 1 0 0 0 0 0 FRE7 6 4 0 10 2 .3 6 2 1 3 0 0 0 0 FPE6 51 0 7 3 4 0 3 3 2 0 0 0 0 0 FRE9 3 3 0 11 4 0 6 0 1 0 0 0 0 0 FREIO 102 0 14 3 .0 6 .0 1 0 0 0 0 0 GRA1 5 5 0 2 6 0 .5 8 .0 0 7 1 0 GRA2 35 0 10 3 1.0 6 2 0 3 1 0 OR A3 30 0 6 0 1 0 2 7 0 3 1 0 GRA4 4 7 0 11 1 0 4 3 0 3 1 0 GRAS 44 0 13 3 1.0 3 3 0 3 1 0 GRA6 7 8 0 2 6 3 1 0 3 7 0 3 1 0 GRA7 5 0 0 1 7 3 1 0 3 3 0 3 1 0 GRA8 70 0 2 9 1 0 7 .0 0 3 1 0 GRA9 5 6 0 14 1 0 4 2 0 .5 1 0 GRAIO 35 0 8 3 1 .0 4 2 0 3 0 0 GRAI 1 3 6 0 11 3 1.0 3 6 0 3 1 0 1 63 LITERATURE CITED
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