MORPHOMETRIC ANALYSIS AND MONOGRAPH OF MONARDA SUBGENUS CHEILYCTIS (LAMIACEAE) By Nathan Joel Sammons A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Plant Biology 2011 ABSTRACT MONOGRAPHY AND MORPHOMETRIC ANALYSIS OF MONARDA SUBGENUS CHEILYCTIS (LAMIACEAE) By Nathan Joel Sammons Monarda subgenus Cheilyctis (Lamiaceae) is a monophyletic group comprised of 16 species of annual and perennial herbs, subshrubs, and shrubs. It has a temperate North American distribution that extends from southern Canada, across much of the United States, into Mexico as far south as Michoacán. The group is most easily characterized by strongly arching upper corolla lips, inserted stamens, and verticillate glomerules. A multivariate morphometric analysis was performed to assess species circumscriptions. Fifty morphological characters were measured from 386 herbarium specimens. Gower’s similarity coefficient was used to perform unweighted pair-group method using arithmetic averages (UPGMA) and principal coordinates analysis (PCOA). The results revealed 11 distinct clusters that correspond to previously accepted species (M. austromontana, M. citriodora, M. clinopodioides, M. fruticulosa, M. humilis, M. maritima, M. mexicana, M. pectinata, M. punctata, M. stanfieldii, and M. viridissima) and five clusters that represent new species or combinations (see below). the data elucidate the characters important to the Boxplots of circumscription of the species. A monograph of this group, including a morphological cladistic analysis, is also presented. Sections covering taxonomic history, morphology, phylogeny, and taxonomy, as well as a key to the species of the subgenus are provided. In addition to a full synonymy and description, phenology, distribution, a specimen image, and a list of representative specimens is also provided for each species. Two species are described (Monarda n. sp. 1, Monarda n. sp. 2), and three new combinations are proposed (M. arkansana ined., M. occidentalis ined., M. villicaulis, ined.) ACKNOWLEDGMENTS There are a number of people and agencies I wish to acknowledge for their contribution to this research, but before I proceed in listing them I wish to express particular gratitude to several individuals whose assistance during these long years has been invaluable. I thank my major professor and mentor, Alan Prather, who has given me help, advice, interminable patience, and inspiration; and also my research committee members Jeff Conner, Andy Jarosz, and Tao Sang whose insights into plant science have refined and enhanced this project. I also wish to thank the curators of the herbaria ARIZ, FLAS, CAS, GH, KNK, MEXU, MICH, MO, MSC, NCU, NMC, OKL, RM, TAMU, TEX, UARK, UNM, and US for graciously making their material available. Many thanks are also due to Paul Taylor, whose grants funded two collection trips to Texas, and to Cecile Gordon and Jack Sammons, the best field assistants in all of Texas. My sincere appreciation also goes to Jan Szyren, and Richard Triemer for aid in digitizing the dissection slides. I also extend special thanks to Rachel Carter for digging me out again and again and always seeing things from the exact right perspective; and Robert Misner for giving, so abundantly, the encouragement, support, and understanding that has made this effort possible. iv TABLE OF CONTENTS LIST OF TABLES…........................................... vii LIST OF FIGURES….......................................... viii CHAPTER 1 INTRODUCTION TO THE GENUS AND THE DISSERTATION…........... 1 CHAPTER 2 MORPHOMETRIC ANALYSIS OF MONARDA SUBGENUS CHEILYCTIS.......... Introduction….......................................... Materials and methods…................................. Results…............................................... Discussion…............................................ 5 6 7 20 57 CHAPTER 3 MONOGRAPH OF MONARDA SUBGENUS CHEILYCTIS.................. 71 Introduction….......................................... Taxonomic history…..................................... Morphology…............................................ Species Concept........................................ Phylogeny…............................................. Keys….................................................. Taxonomy….............................................. Monarda austromontana.................................. Monarda citriodora..................................... Monarda clinopodioides................................. Monarda mexicana....................................... Monarda pectinata...................................... Monarda n. sp. 1....................................... Monarda arkansana ined................................. Monarda fruticulosa.................................... Monarda humilis........................................ Monarda maritima....................................... 72 74 75 82 82 102 106 107 113 119 124 129 133 137 141 146 150 v Table of Contents (Cont’d) Monarda Monarda Monarda Monarda Monarda Monarda occidentalis ined.............................. punctata....................................... stanfieldii.................................... villicaulis ined............................... viridissima.................................... n. sp. 2....................................... 154 159 167 171 176 181 OVERVIEW OF ONLINE MONOGRAPH…............................. 186 APPENDICES............................................... 196 CHAPTER 4 Appendix A. Morphometric data for Monarda section Cheilyctis. Character numbers correspond to those presented in the Materials and Methods section............ Appendix B. Morphometric data for Monarda section Aristatae. Character numbers correspond to those presented in the Materials and Methods section........... LITERATURE CITED.......................................... vi 197 261 296 LIST OF TABLES Table 1 2 Page Characters used in the phylogenetic analysis of Monarda subgenus Cheilyctis........................ 90 Data matrix of cladistics characters for species of Monarda subgenus Cheilyctis and two outgroups (M. bradburiana and M. russeliana). The character number matches those presented in table 1.......... 94 3 Morphometric data for Monarda Section Cheilycits... 198 4 Morphometric data for Monarda Section Aristatae.... 262 vii LIST OF FIGURES Figure 1 2 3 4 5 6 7 Page Synopsis of the Monarda section Aristatae UPGMA dendrogram based on Gower similarity estimates of 116 taxa. Clusters of individuals of the same species (or intraspecific rank) are represented by a single branch, and the number of individuals in that branch is indicated in parentheses, if more than one.................................. 22 Group one in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity......................... 23 Group two in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity......................... 24 Group three in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity......................... 25 Group four in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity......................... 26 PCoA plot of the first three principal Coordinates. The variability accounted for by each axis is 19.5% for the first, 14% for the second, and 10% for the third. Outlined regions correspond to groups discussed in text.............................................. 28 PCoA plot of the first three principal coordinates. The variability accounted for by each axis is 19.5% for the first, 14% for the second, and 10% for the third. This is a rotation of the same plot shown in Fig. 6. Arrow highlights the the single specimen of Monarda n. sp. 1.................................. 29 viii List of Figures (Cont’d) Figure 8 9 10 11 12 13 14 15 16 Page Box plots of Petiole Length (mm), Serration Distance (mm), and Leaf Length (cm) of Monarda section Aristatae......................... 31 Box plots of Leaf Widest Point (cm), Width of Lower Bract (mm), and Widest Point of Lower Bract (mm) of Monarda section Aristatae........... 32 Box plots of Length of Upper Bract (cm), Width of Upper Bract (mm), and Widest Point of Upper Bract (mm) of Monarda section Aristatae......................................... 33 Box plots of Calyx Length (mm) and Calyx Lobe Width (mm) of Monarda section Aristatae........... 34 Box plots of Lower Lip Width (mm) and Lateral-Limb Length (mm) of Monarda section Aristatae......................................... 35 Synopsis of the Monarda section Cheilyctis UPGMA dendrogram based on Gower similarity estimates of 170 OTUs. Clusters of individuals of the same species (or intraspecific rank) are represented by a single branch, and the number of individuals in that branch is indicated in parentheses........ 38 Group one in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity......................... 39 Group five in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity......................... 41 Group two in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity......................... 42 ix List of Figures (Cont’d) Figure 17 18 19 20 21 22 23 24 Page Portion of group 3 in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity......................... 43 Second branch in group three in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity...... 44 Group four in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity......................... 45 PCoA plot of the first two principal coordinates. The variability accounted for by each axis is 18.1 for the first and 12 for the second. Groups encircled by the dotted lines are discussed in the text................... 48 PCoA plot of the first three principal coordinates. The variability accounted for by each axis is 18.1% for the first, 12% for the second, and 5.2% for the third. a = M. punctata from Florida and southeast U.S., b = M. humilis, c = M. stanfieldii, d = M. villicaulis ined.................................. 49 Box plots of Internode length (mm), Internode Width (cm), and Petiole Length (mm) of Monarda section Cheilyctis........................ 50 51 Box plots of Length of the Serrated Margin (mm), Leaf Tooth Size (mm), and Leaf Length (cm) of Monarda section Cheilyctis................ 52 Box plots of Length of Upper Bract (cm), Width of Upper Bract (mm), and Leaf Width (cm) of Monarda section Cheilyctis................ 53 x List of Figures (Cont’d) Figure 25 26 27 28 29 30 Page Box plots of Number of Flowering branches, Calyx Length (mm), and Calyx Orifice Pubescence Length (mm) of Monarda section Cheilyctis........................................ 54 Box plots of Calyx Lobe Width (mm), Lower Corolla Lip Width (mm), and Mid-Lip Length of Monarda section Cheilyctis..................... 55 Box plots of Mid-Limb Width (mm), Lateral-Limb Length (mm), and Lateral-Limb Width (mm) of Monarda section Cheilyctis........................................ 56 Box plots of Upper Corolla Lip Length (mm), Upper Corolla Lip Width (mm), and Upper Limb Length (mm) of Monarda section Cheilyctis......... 57 Trichome types. A. Unicellular hairs along the stem of M. punctata (Correll 18925, TEX). B. Multicellular hairs on the pilose nodal region of M. maritima (Johnston 53254.22, TEX). C. Glandular punctations on a canescent adaxial leaf surface of M. fruticulosa (Garcia 110, TEX). D. Villous abaxial leaf surface of M. villicaulis ined. (Bourdo 306, MSC). For interpretation of the references to color in this and all other figures, the reader is referred to the electronic version of this dissertation........... 79 Calyx lobe shapes and vesture types. A. M. clinopodioides (Lawson 40, OKL) with aristate calyx lobes and long lobe margin hairs; B. M. humilis (Higgins 7737, NMC) with acute calyx lobes and glabrous margins; C. M. austromontana (Maysilles 8272, TEX) showing sparsely packed, short calyx lobe margin hairs; D. M. maritima (Churchill 90-930, MSC) with lanceolate calyx lobes; E. M. citriodora with orifice and calyx tube hairs................. 82 xi List of Figures (Cont’d) Figure 31 32 Page One of four equally most parsimonious cladograms for Monarda subgenus Cheilyctis. Numbers represent unambiguous state changes for the characters listed in table x. Outgroups are M. bradburiana and M. russeliana........................................ 89 Strict consensus tree of Monarda subgenus Cheilyctis derived from four equally parsimonious cladograms (114 steps, CI = 0.711, RI = 0.708). Taxonomic sections are indicated by vertical bars, and numbers above branches indicate bootstrap values. Outgroups are M. bradburiana and M. russeliana........................................ 90 33 Herbarium specimen of M. austromontana. Van Devender 96-378 (ARIZ)............................ 111 34 Herbarium specimen of M. citriodora. Massey 2070 (OKL)........................................ 117 35 Herbarium specimen of M. clinopodioides. Dubrule 307 (TAMU)................................ 122 36 Herbarium specimen of M. mexicana. Maysilles 7771 (TEX)........................................ 127 37 Herbarium specimen of M. pectinata. Ballinger s.n. (TEX)........................................ 131 38 Herbarium specimen of Monarda n. sp. 1. Yen 2681 (CAS)........................................ 136 39 Herbarium specimen of M. arkansana. Moore 400459 (UARK)..................................... 140 40 Herbarium specimen of M. fruticulosa. Runyon 3991 (RM)......................................... 144 41 Herbarium specimen of M. humilis. Goodrow 522 (UNM)............................................. 149 xii List of Figures (Cont’d) Figure Page 42 Herbarium specimen of M. maritima. Johnston 53254.22 (TEX).................................... 153 43 Herbarium specimen of M. occidentalis. Correll 33017 (TEX)............................... 157 44 Herbarium specimen of M. punctata. Moore 400459 (UARK)..................................... 163 45 Herbarium specimen of Monarda stanfieldii. Lundell 9029 (TEX)................................ 170 46 Herbarium specimen of M. villicaulis. Hestbeck 10 (MSC)................................. 174 47 Herbarium specimen of M. viridissima. Correll 19593 (GH)........................................ 179 48 Herbarium specimen of Monarda n. sp. 2. Herring 349 (FLAS)................................ 184 xiii CHAPTER 1 INTRODUCTION TO THE GENUS AND THE DISSERTATION 1 Monarda L. (Lamiaceae) is a temperate North American genus with a natural distribution that extends from southern Canada, across much of the United States, and into Mexico as far south as Michoacán (Scora 1967). It has bright colored flowers and is commonly grown ornamentally. The genus has agricultural value as companion plants used to encourage pollinator visits and Monarda punctata L. has been shown to be a good choice for habitat manipulation projects that focus on the maintenance of natural enemy arthropod populations to minimize the use of poisons on adjacent crop fields (Frank, Shrewsbury et al. 2008). Monarda is perhaps most widely utilized for its phytochemistry though it is not cultivated expressly for such purposes. Essential oils and extracts (chiefly thymol, carvacrol, and phenolic monoterpenes) from species in this genus have long been recognized as having therapeutic and bioactive properties (Pammel 1911) and they have been used to treat nausea, swelling, rheumatic pain, fever, and coughs (Chevallier 1996). Thymol, in particular, has been used to kill parasites such as ascarids and hookworms and is used as an antiseptic in soaps, mouthwashes, and toothpastes. Dorman and Deans (2004) show that M. citriodora Cerv ex Lag., oils are strong antioxidants, and M. fistulosa L. extracts, long used as a stimulant and to remove colic pain, is recommended as a 2 treatment to Seborrhea (Zhilyakova, Novikov et al. 2009) because it demonstrates antibacterial, antimycotic, and antiinflammatory properties. Yamada et al (2010) recently isolated seven novel glycosides from M. punctata and found that carvacrol isolated from M. punctata has an inhibitory effect on the hydrolysis of triacylglycerols. They further suggest it as a candidate for treating and preventing metabolic syndrome. Despite this interest in Monarda, the taxonomy of this group remains problematic. Even with Scora’s (1967) detailed and comprehensive examination of the genus, new work has emerged regarding its pollination ecology (Whitten 1981; Cresswell 1990), new interest has arisen concerning evolution within the genus (Prather, Monfils et al. 2002; Keith 2003), and there has been continued debate on its taxonomy (Correll and Johnston 1979; Turner 1994; Prather and Keith 2003). In this light it is time to reexamine the portion of the genus (Monarda subgenus Cheilyctis) which continues to be characterized by high degrees of taxonomic uncertainty. The objectives of this research are to 1) construct a current taxonomic hypothesis of Monarda subgenus Cheilyctis, 2) produce a traditional monograph for the subgenus, and 3) create a web-based monograph providing additional features not available in a traditional paper-bound manuscript. 3 The remaining chapters of this dissertation address the three above-mentioned objectives. Chapter 2 contains a morphometric analysis that uses descriptive statistics to examine the variation among the taxa of the subgenus as well as multivariate statistics to examine species distinctiveness and groupings. A morphometric analysis was chosen over a DNA-based analysis because of the availability of herbarium material used in the study. To collect DNA samples from a similar number of individuals across the same distributions would have been prohibited by time and financial constraints. A species list is also presented in chapter 2, and characters that are useful in developing that circumscription are discussed. Chapter 3 contains the traditional monograph for Monarda subgenus Cheilyctis. It covers morphology, taxonomic history, phylogeny, keys, synonymy, phenology, distribution, specimen images, species descriptions, and representative specimens for each taxon in the group. The final chapter presents an overview of the online monograph and provides a list of features it contains. It also discusses some other works that highlight internet-based technologies in plant systematics. 4 CHAPTER 2 MORPHOMETRIC ANALYSIS OF MONARDA SUBGENUS CHEILYCTIS 5 INTRODUCTION Though the taxonomy of Monarda subgenus Cheilyctis has had an incremental increase in the number of species in the last 200 years, the taxonomic history of the group is best characterized by a persistent shifting in the ranks of the existing taxa. In the last five decades, even though there has only been one new species and three new varieties (Scora 1965; Scora 1967; Correll 1968; Turner 1994) there have been 10 new combinations (Waterfall 1950; Cory 1953; Shinners 1953; Scora 1965; Scora 1967; Turner 1994; Prather and Keith 2003), each of which involved rank shifts between species, subspecies, or variety. Of the 11 varieties of Monarda punctata published in Scora’s (1967) treatment of the genus, only M. punctata var. maritima had not already received at least one rank change. Monarda punctata var. maritima would later be elevated to species by B.L. Turner (1994). Much of the taxonomic juggling these taxa have received stems from the fact that the majority are regionally endemic and morphologically very similar, or from the fact that the traits being used to separate the groups are often less conspicuous (calyx lobe width, angle of stem indumentum). The hurdles, then, in building a complete taxonomy of this group lies in developing a solid understanding of what role the morphological 6 variation plays at both the inter and intraspecific levels of the subgenus. While there have indeed been detailed works examining the morphological differences present among the species of the group, these putative differences have not been studied quantitatively. The goals of this investigation are to 1) evaluate the morphological variation between the species of the subgenus, and among the varieties of M. punctata, in particular, and 2) propose and test my circumscription of the taxa in this subgenus. MATERIALS AND METHODS Taxon sampling The following is (based on preliminary observations of herbarium specimens) my putative species taxonomy for Monarda subgenus Cheilyctis: Section Cheilyctis M. fruticulosa Epling M. humilis (Torrey) Prather & J.A. Keith M. maritima (Cory) B.L. Turner M. occidentalis ined 7 M. punctata L. var. punctata var. arkansana (McClintock and Epling) Shinners var. correllii B.L. Turner var. intermedia (McClintock and Epling) Waterfall var. lasiodonta Gray M. villicaulis ined M. stanfieldii Small M. viridissima Correll Section Aristatae M. austromontana Epling M. citriodora Cervantes ex Lagasca var. citriodora var. parva Scora M. clinopodioides Gray M. pectinata Nuttall M. mexicana Epling Species and variety identity for each specimen was consistent with Turner (1994) for all Texas collections, Prather and Keith (2003) for Monarda humilis and Scora (1967) for all other collections. The new combinations I propose in my taxonomy 8 (M. occidentalis ined, M. villicaulis ined) and M. austromontana were also identified as above, though they are all identified as intraspecific taxa by those authors, but treated as species in my taxonomic circumscriptions. Traits were measured from 381 specimens in anthesis from the following herbaria: ARIZ, FLAS, CAS, GH, KNK, MEXU, MICH, MO, MSC, NCU, NMC, OKL, RM, TAMU, TEX, UARK, UNM, and US standardized abbreviation from Index Herbariorum (The New York Botanical Garden 2007). These taxa represent populations from across the distribution of Monarda subgenus Cheilyctis. Herbarium specimens were measured and treated as an independent operational taxonomic units (OTU) (Appendix A, B). I sampled 30 specimens of each OTU (species/variety of Monarda subgenera Cheilyctis and Aristatae) where possible. Due to lack of equal representation in the loan/collections, this was not an option for all taxa. The number of specimens I measured per taxon is as follows: 27 for M. austromontana ined, 25 for M. citriodora var. citriodora, 3 for M. citriodora var. parva, 30 for M. clinopodioides, 30 for M. fruticulosa, 15 for M. humilis, 20 for M. maritima, 25 for M. occidentalis ined, 28 for M. pectinata, 15 for M. punctata var. arkansana, 30 for M. punctata var. intermedia, 30 for M. punctata var. lasiodonta, 30 for M. punctata var. punctata, 20 for M. stanfieldii, 30 for M. villicaulis ined, and 25 for M. viridissima. 9 Character sampling The characters selected for the analysis were chosen because they 1) were considered important by previous investigators of Monarda 2) demonstrated variability between and within taxa or 3) have been of historical interest in the taxonomy of Lamiaceae. Measurements were taken either by direct measuring (with a linear rule), measuring using a dissection scope and ocular micrometer, or measurements made from a digital image taken through a dissection scope (list of traits measured by each method is discussed below). For the traits that were imaged, measurements were taken using tpsDig2 (Rohlf 2004). This software package measures the distance between user-specified landmark points on a digital image, and, given an accurate scaling standard, converts the measurements to a standard measuring unit (μm or mm). The techniques employed to measure a specific character were used consistently among all individuals. Quantitative traits. Most vegetative characters were measured using a binocular dissection scope with an ocular micrometer. Large characters (leaf and stem dimensions) were measured with a 10cm plastic rule. All measurements were taken at standardized reference points (see below). The leaves and nodes that were 10 measured were found at the second node subtending the basal glomerule. The internodes were those which immediately subtended the above mentioned node. 1. Internode Length – taken from the penultimate node subtending the inflorescence to the next lower node. 2. Internode Width – taken from at the mid-point of the penultimate internode subtending the inflorescence. 3. Leaf Length – taken from the tip of the leaf to the base of the petiole. 4. Leaf Width – taken from the widest point of the leaf. 5. Leaf Widest Point – distance from the base of the petiole to the widest point of the leaf. 6. Petiole Length – taken from the base of the lowermost portion of the leaf blade to the leaf node. 7. Length of Serrated Margin – taken from tip of leaf to basal most serration. 8. Serration Distance – distance from the distal point of attachment of one tooth to the distal point of attachment of an adjacent tooth. Measurement taken at the approximate midpoint of the serrated region of the leaf. 9. Leaf Tooth Size – taken from the distal point of attachment to the tip of the tooth. 11 10. Length Laminar Tissue – taken from tip of petiole to tip of leaf. 11. Number of Flowering Branches – number of inflorescences on the plant. 12. Lower Bract Length – taken from the tip of the lower bract to the attachment point. 13. Lower Bract Width – taken from the widest point of the lower bract. 14. Lower Bract Widest Point – distance from the base of the lower bract to the widest point. 15. Upper Bract Length – taken from the tip of the upper bract to the attachment point. 16. Upper Bract Width – taken from the widest point of the upper bract. 17. Upper Bract Widest Point – distance from the base of the upper bract to the widest point. All floral characters were measured using tpsDig. The image files were generated by taking an image with a digital camera (Nikon cool-pix) mounted onto a dissecting microscope. Wet mounts of the flowers were prepared by teasing the corolla from the calyx, flattening the upper and lower corolla lips, exposing the stigma and stamens, and flattening the lobes of the upper and lower corolla lips. The floral tissues were suspended on 12 the slide in permount and covered with a 0.1 mm cover slip. Slides were preserved by sealing the cover slip with fingernail polish. Raw jpg images were converted to tps files using tps_util software. The images were scaled by taking a digital image of a 10cm plastic rule at the same magnification as the floral slides. This image was used to calibrate tpsDig before taking the measurements. The following is a list of the standard points where each floral character was measured. 18. Calyx Length – taken from the base of the calyx to the base of a calyx lobe. 19. Calyx Width – taken from one side of the flattened calyx to the other, or when the calyx is open, the distance across half of the calyx veins. 20. Calyx Orifice Pubescence Length – taken from the hairs at the base of the adaxial side of the calyx lobe. 21. Calyx Lobe Length – taken from the base of the calyx lobe to the tip of the calyx lobe. 22. Calyx Lobe Width – taken from the middle of the calyx lobe. 23. Calyx Lobe Margin Pubescence Length – taken from the hairs on the margin of the calyx lobe. 13 24. Lower Corolla Lip Length – taken from the upper lip/lower lip connection fold to the terminus of the midlimb of the lower lip. 25. Lower Corolla Lip Width – taken from one lateral margin of the lip to the other lateral margin of the lip at the base of the limbs. 26. Mid-limb Length – taken from the tip of the middle limb of the lower corolla lip to the base of the cleft along the midpoint of the middle limb 27. Mid-limb Width – taken from one lateral margin of the middle limb of the lower corolla lip to the other lateral margin of the middle limb at the widest point of the midlimb. 28. Lateral-limb Length – taken from the tip of the lateral limb of the lower corolla lip to the base of the cleft along the midpoint of the lateral limb. 29. Lateral-limb Width – taken from one margin of the lateral limb of the lower corolla lip to the other margin of the lateral limb. 30. Upper Corolla Lip Length – taken from upper lip/lower lip connection fold to the tip of the upper lip. 31. Upper Corolla Lip Width – taken from one lateral margin of the lip to the other lateral margin of the lip at roughly half-way up the lip. 14 32. Limb Length (cleft depth) – taken from the tip of the upper corolla limb to the base of the cleft along the midpoint of the limb. 33. Limb Width – taken from one lateral margin of the upper corolla limb to the other lateral margin of the limb at the base of the limb. 34. Hair Length - taken from the hairs on the outer surface of the upper corolla lip. Qualitative traits. These characters were observed with a compound dissection scope to detect the presence (scored as ―1‖) or absence (scored as ―0‖) of hair types on the surface of various plant parts. In the characters described below ―<3‖ and ―>5‖ refer to the presence of hairs comprised of less than three cells or greater than five cells, respectively. Organ selection was consistent with the methods used in the quantitative traits (as described above). 35. Internode <3 – examined on the facial surface of the internode. 36. Internode >5 – same as above. 37. Nodal Ridges <3 – examined at the edges of the node, which are the corners formed by fibrous bundles. 15 38. Nodal Ridges >5 – same as above. 39. Inter-ridge Region <3 – examined on the facial surface of the node. 40. Inter-ridge Region >5 – same as above. 41. Interpetiolar Plane <3 – examined at the equatorial region of the node. 42. Interpetiolar Plane >5 – same as above. 43. Petiole <3 – examined on the petiole of the selected leaf. 44. Petiole >5 – same as above. 45. Leaf Midvein <3 – examined on the midvein of the selected leaf. 46. Leaf Midvein >5 – same as above. 47. Abaxial Surface <3 – examined on the laminar region of the abaxial side of the leaf. 48. Abaxial Surface >5 – same as above. 49. Calyx Facial Tissue <3 – examined between the veins of the calyx tubes 50. Calyx Facial Tissue >5 – same as above. Data Analysis Cluster Analysis. All quantitative data were standardized to zero mean and variance in an effort to reduce the effects of 16 different scales of measurement for the different characters. The standardized data were then used to create a similarity matrix based on the Gower coefficient (Gower 1971; Sneath and Sokal 1973). This metric was selected because of its flexibility in using both quantitative and qualitative data (Gower 1971; Binns, Baum et al. 2002; Podani and Schmera 2006). Standardization and the computation of the similarity matrix were performed using the statistical package, MVSP (Kovach Computing Services 1998). Using NTSYSpc 2.1 (Rohlf 2000), dendrograms were generated with the unweighted pair-group method using arithmetic averages (UPGMA) clustering algorithm (Sneath and Sokal 1973; SAHN module with UPGMA as clustering method). Ordination. Data were standardized as above, and the Gower similarity matrix was used to complete a Principal Coordinate Analysis (Sneath and Sokal 1973; Reyment, Blackith et al. 1984; Marcus 1996; Anderson and Willis 2003). This ordination method was chosen over the more commonly used Principal Components Analysis (PCA) because it can be applied to many different kinds of similarity or distance measures. It is implicit with PCA that a covariance or correlation matrix be used, which is not the case for this investigation (see above for explanation). PCOA is also preferred because it focuses on the relationship among individuals as opposed to the relationship among characters (as 17 in PCA) and evaluates OTU’s individually instead of under the assumptions of group membership (Reyment, Blackith et al. 1984; Marcus 1996; McCauley and Ballard 2007). NTSYSpc 2.1 was used to double-center (convert the triangular Gower matrix into a square matrix) the data matrix (DCENTER module with square distances selected), to calculate the eigenvectors and eigenvalues (EIGEN module with four dimensions selected and a vector scaling of SQRT(LAMBDA)) and to examine the axes. Descriptive statistics. Boxplots were generated using the default boxplot command in R (R Development Core Team 2009). The upper and lower region of the boxes correspond to the upper (75th percentile) and lower (25th percentile) quartiles, respectively. The line in the box is the median, and the dashed lines represent one standard deviation above and below the mean. Outliers are represented by open-faced circles. Because the sections of Monarda subgenus Cheilyctis are monophyletic (Prather et al. 2002) the analyses were performed separately for each section, to keep the interpretation of the results from becoming too cumbersome. The taxonomic hypotheses for the subgenus are based on the phenetic species concept (Sokal and Crovello 1970; De Queiroz 2007), which assigns individuals into groups (species) which are defined by morphological gaps. My species circumscriptions will 18 be tested based on how well the individuals of those putative species are separated morphologically. If there are sufficient morphological gaps between individuals of two species, then they will cluster on different branches of the UPGMA dendrogram. Likewise, individuals of different species will cluster away from each other in the multidimensional space of the PCOA map. However, if complete agreement between the analyses does not occur, either the UPGMA or the PCOA must suggest that a taxon is distinct from the others and there must be some support from individual morphological characters (possession of a novel qualitative trait or non-overlapping quartiles (as described above) for a quantitative traits or the taxon will not be recognized. It should also be noted that interpretations of the two analyses may have varying degrees of subjectivity. Because branches connect at discrete locations in the UPGMA dendrograms, interpretation of groups is a very direct process. In contrast, clusters in PCOA are more subject to interpretation, as gaps between taxa or groups of taxa are dependent on 1) how many dimensions are being shown (with all but the first three dimensions unusable because of our inability to perceive beyond three dimensions) and 2) at what angle the groups are being observed. Taxa appearing clustered together in two dimensions can actually be separated by very large gaps when adding a third 19 dimension, and taxa appearing clustered at one angle can be seen not to be by rotating the view to a different angle. Furthermore, the incapacity to utilize data beyond the third dimension may result in more conservative groupings because it actually employs less of the data for demonstrating putative gaps. The UPGMA tends to be more sensitive to outliers as well, and large outliers can result in large branch lengths. As a result of the aforementioned properties of the analyses, discrete clusters in the PCOA analysis tend to show less distance between them than they do in the UPGMA analysis. RESULTS Section Aristatae UPGMA The dendrogram of section Aristatae shows four major groups (Fig. 1 shows a synopsis of the entire dendrogram) as well as a long branch that represents the single specimen from Chihuahua, Mexico (referred hereafter as Monarda n. sp. 1) and another long branch that encompasses the two specimens of M. mexicana. Where there are groups of taxa belonging to the same species, all those individuals are represented by one branch. Groups 1 and 2 (Figs. 2 and 3) are comprised entirely of individuals of M. clinopodioides and M. pectinata, respectively. 20 Group 3 (Fig. 4) is comprised of specimens of M. citriodora var. citriodora and M. citriodora var. parva. Group 4 (Fig. 5) is comprised of all of the M. var. austromontana specimens. Monarda n. sp. 1 and M. mexicana can also be seen in Figure 5. 21 Figure 1. Synopsis of the Monarda section Aristatae UPGMA dendrogram based on Gower similarity estimates of 116 taxa. Clusters of individuals of the same species (or intraspecific rank) are represented by a single branch, and the number of individuals in that branch is indicated in parentheses, if more than one. 22 Fig. 2. Group one in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity. 23 Figure 3. Group two in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity. 24 Figure 4. Group three in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity. 25 Figure 5. Group four in the UPGMA dendrogram of Monarda section Aristatae based on the Gower coefficient of similarity. 26 PCOA A scatter plot (Fig. 6) of the first three principal coordinates shows four main clusters of taxa. The first cluster is comprised of M. pectinata with the two specimens of M. mexicana found just below that cluster. The second contains M. austromontana and the single specimen of Monarda n. sp. 1. The third is comprised exclusively of M. clinopodioides, and the fourth cluster contains specimens of both M. citriodora var. citriodora and M. citriodora var. parva. A rotation of that plot (Fig. 7) reveals further separation between the specimens of M. mexicana and M. pectinata and Monarda n. sp. 1 from M. austromontana. For this analysis the first three principal coordinates accounted, cumulatively, for 19.5%, 33.5%, and 43.5% of the total variation, respectively, which is consistent with PCOA values of similar analyses (McCauley and Ballard 2007; Pereira, Perez et al. 2007; Morawetz and Wolfe 2011). The remaining principal coordinates each summarize 6.5% or less of the total variation. 27 Figure 6. PCoA plot of the first three principal coordinates. The variability accounted for by each axis is 19.5% for the first, 14% for the second, and 10% for the third. Outlined regions correspond to groups discussed in text. 28 Figure 7. PCoA plot of the first three principal coordinates. The variability accounted for by each axis is 19.5% for the first, 14% for the second, and 10% for the third. This is a rotation of the same plot shown in Fig. 6. the single specimen of Monarda n. sp. 1. 29 Arrow highlights the Descriptive Statistics Box plots showed 13 characters that may be useful (defined as non-overlapping upper and lower quartiles (Pereira, Perez et al. 2007) in differentiating the taxa in this section. Vegetative traits (Figs. 8, 9 and 10) include Petiole Length, Serration Distance, Leaf Length, Leaf Widest Point, Width Lower Bract, Widest Point of Lower Bract, Length Upper Bract, Width Upper Bract, Widest Point of Upper Bract. Floral traits (figs 11 and 12) include Calyx Length, Calyx Lobe Width, Lower Lip Width, Lateral Limb Length. 30 Figure 8. Box plots of Petiole Length (mm), Serration Distance (mm), and Leaf Length (cm) of Monarda section Aristatae. 31 Figure 9. Box plots of Leaf Widest Point (cm), Width of Lower Bract (mm), and Widest Point of Lower Bract (mm) of Monarda section Aristatae. 32 Figure 10. Box plots of Length of Upper Bract (cm), Width of Upper Bract (mm), and Widest Point of Upper Bract (mm) of Monarda section Aristatae. 33 Figure 11. Box plots of Calyx Length (mm) and Calyx Lobe Width (mm) of Monarda section Aristatae. 34 Figure 12. Box plots of Lower Lip Width (mm) and Lateral-Limb Length (mm) of Monarda section Aristatae. 35 Section Cheilyctis UPGMA The dendrogram of section Cheilyctis shows five major groups (Fig. 13 shows a synopsis of the entire dendrogram). Detailed regions of the tree are referenced in the figures below. Groups 1 (Fig. 14) and 5 (Fig. 15) are comprised of only M. fruticulosa and M. maritima, respectively, while groups 2, 3, and 4 contain a number of different taxa. Group 2 (Fig. 16) contains three main clusters (subgroups) that correspond to M. humilis, M. stanfieldii, M. occidentalis ined, and a small cluster of four individuals of various M. punctata varieties. Group 3 shows three clusters: two clusters of M. punctata varieties (Fig. 17) and one large cluster that contains mostly M. punctata var. arkansana and M. villicaulis ined plus two samples of M. punctata vars. (Fig. 18). Finally, group 4 (Fig. 19) contains two main clusters as well as a single specimen of M. punctata var. correllii which is attached at the base of the two main clusters. The first of the two above-mentioned clusters corresponds to M. viridissima. The other branch is comprised of seven specimens of M. punctata var. punctata (hereafter referred in the text as Monarda n. sp. 2) found in Florida and the southeast U.S. which have leaves that are greatly reduced in 36 size compared to other specimens of M. punctata var. punctata in that region. A more detailed discussion of additional traits that differentiate these taxa is found in the Discussion section. 37 Figure 13. Synopsis of the Monarda section Cheilyctis UPGMA dendrogram based on Gower similarity estimates of 170 OTUs. Clusters of individuals of the same species (or intraspecific rank) are represented by a single branch, and the number of individuals in that branch is indicated in parentheses. 38 Figure 14. Group one in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. 39 Monarda maritima is found in southeastern Texas along the Gulf of Mexico. In the UPGMA tree it is found clustered among other OTU’s of the same rank (Fig. 15). 40 Figure 15. Group five in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. 41 Figure 16. Group two in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. The number ―1‖ represents two specimens of M. punctata var. lasiodonta and one specimen of M. punctata var. correllii. 42 Figure 17. Portion of group 3 in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. 1=var. arkansana, 2=var. lasiodonta, 3=var. correllii, 4=var. intermedia, 5=var. punctata. 43 Figure 18. Second branch in group three in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. 1=var. arkansana, var. lasiondonta, var. punctata; 2=M.arkansana. 44 Figure 19. Group four in the UPGMA dendrogram of Monarda section Cheilyctis based on the Gower coefficient of similarity. 1= M.punctata var. punctata 45 PCOA A scatter plot (Fig. 20) of the first two principal coordinates shows seven main clusters of taxa. is comprised of all of the M. maritima. The second group contains the Monarda n. sp. 2 specimens. of the M. viridissima samples. The first group Group 3 contains all The fourth group has all of M. fruticulosa as well as some M. humilis. The rest of M. humilis is found in the fifth group, along with all of the specimens of M. stanfieldii and M. occidentalis ined. Group six contains several specimens of M. punctata var. lasiodonta, M. punctata var. punctata, M. punctata var. intermedia, M. punctata var. correllii, and M. punctata var. arkansana. Group seven contains all of M. villicaulis ined as well as some individuals of M. punctata var. arkansana. Adding the third coordinate (Fig. 21) demonstrates that M. maritima and Monarda n. sp. 2 (groups 1 and 2 from figure 20) are actually separated by a larger gap than appears from the Fig. 20. Monarda n. sp. 1 (―a‖ in Fig. 21) can be seen with better separation from M. maritima (―b‖ in Fig. 21). This is also the case for the M. humilis individuals (―c‖ in Fig. 21) from groups 4 and 5. In figure 21 they can be seen clustered together, though they are still in close proximity to M. 46 occidentalis (not labeled) which are positioned immediately below the M. humilis individuals. group 7 should be divided further. Figure 21 also reveals that Monarda villicaulis ined (―d‖ in Fig. 21) is clustered separately from the M. punctata var. arkansana (―e‖ in Fig. 21) in that original grouping. For this analysis the first three principal coordinates accounted, cumulatively, for 18.1%, 30.1%, and 36.3% of the total variation, respectively. The remaining principal coordinates each summarize 5.3% or less of the total variation. 47 Figure 20. PCoA plot of the first two principal coordinates. The variability accounted for by each axis is 18.1 for the first and 12 for the second. Groups encircled by the dotted lines are discussed in the text. 48 Figure 21. PCoA plot of the first three principal coordinates. The variability accounted for by each axis is 18.1% for the first, 12% for the second, and 5.2% for the third. a = M. punctata from Florida and southeast U.S., b = M. humilis, c = M. stanfieldii, d = M. villicaulis ined. 49 Descriptive statistics Box plots showed 21 characters (with non-overlapping quartiles) that may be useful in differentiating the taxa in this section. Vegetative traits (figs 22, 23, 24) include internode length, internode width, petiole length, length of serrated margin, leaf tooth size, leaf length, leaf width, upper bract length, upper bract width. Inflorescence traits (figs. 25 - 28) include number of flowering branches, calyx length, calyx orifice pubescence length, calyx lobe width, lower corolla lip width, mid-limb length, mid-limb width, lateral-limb length, laterallimb width, upper corolla lip length, upper corolla lip width, upper limb length. order: Box plots display the taxa in the following M. arkansana, var. correllii, M. fruticulosa, M. humilis, var. lasiodonta, M. maritima, M. occidentalis, var. punctata, Monarda n. sp. 2, M. stanfieldii, M. villicaulis, M. viridissima. 50 Figure 22. Box plots of Internode length (mm), Internode Width (cm), and Petiole Length (mm) of Monarda section Cheilyctis. 51 Figure 23. Box plots of Length of the Serrated Margin (mm), Leaf Tooth Size (mm), and Leaf Length (cm) of Monarda section Cheilyctis. 52 Figure 24. Box plots of Length of Upper Bract (cm), Width of Upper Bract (mm), and Leaf Width (cm) of Monarda section Cheilyctis. 53 Figure 25. Box plots of Number of Flowering branches, Calyx Length (mm), and Calyx Orifice Pubescence Length (mm) of Monarda section Cheilyctis. 54 Figure 26. Box plots of Calyx Lobe Width (mm), Lower Corolla Lip Width (mm), and Mid-Lip Length of Monarda section Cheilyctis. 55 Figure 27. Box plots of Mid-Limb Width (mm), Lateral-Limb Length (mm), and Lateral-Limb Width (mm) of Monarda section Cheilyctis. 56 Figure 28. Box plots of Upper Corolla Lip Length (mm), Upper Corolla Lip Width (mm), and Upper Limb Length (mm) of Monarda section Cheilyctis. 57 Discussion The following discussion is organized into two sections. The first section is a discussion, taxon by taxon, of the taxonomic conclusions made concerning my putative species circumscriptions (from the Methods section). Those will be addressed in the following order: 1) taxa that are supported by both UPGMA and PCOA, 2) taxa that are supported by one analysis but ambiguous or not supported in the other 3) taxa that receive no support from either analysis The second section is a discussion of the contribution of morphological traits to the taxonomy of Monarda subgenus Cheilyctis. It covers characters that were shown in the box plot analyses and briefly summarizes the historical importance of those features as well as their utility in my current taxonomy for the subgenus. Taxonomic conclusions TAXA SUPPORTED BY BOTH UPGMA AND PCOA. Of the species presented in my putative taxonomy (Methods section), M. austromontana Epling, M. clinopodioides, M. maritima, M. villicaulis ined, M. 58 viridissima, M. fruticulosa, Monarda n. sp. 2, and M. pectinata have clear support by both UPGMA and PCoA. In both analyses these taxa cluster exclusively together in accordance with my hypothesis that they are indeed species. Of these, M. clinopodioides, M. pectinata, M. maritima, M. viridissima, and M. fruticulosa are all currently accepted species. Monarda austromontana Epling and M. villicaulis ined were considered varieties of M. citriodora and M. punctata, respectively, and Monarda n. sp. 2 is a spec. nov. from Florida and the southeastern U.S. Monarda austromontana is most easily recognized by its reflexed lower bracts but also has the longest (> 1.2 mm) lateral limbs of section Aristatae. Monarda villicaulis is most easily recognized by the qualitative indumentum traits (Adaxial Surface >5, Abaxial surface >5, Inter-ridge Region >5) that give the species villous laminar and stem surfaces making the plants appear silvery. Furthermore, they are geographically isolated with a distribution in the northeastern regions of the US, extending only as far south and west as Missouri. Monarda n. sp 2 is best distinguished from the other taxa in this section by smaller (<4cm long, <1 cm wide) ovate leaves (Figs. 23, 24), thin and spindly stems (measured as internode width, Fig. 22) with occasional prostrate growth, and short calyces (Fig. 25) that make the flowers appear smaller 59 than the other species in section Cheilyctis. It is found in central Florida and the southeastern U.S. TAXA SUPPORTED BY UPGMA, AMBIGUOUS IN PCOA. Monarda n. sp. 1, M. humilis, M. mexicana, M. stanfieldii, and M. occidentalis ined also all cluster together (or have clear separation via a small similarity coefficient with any other OTU's as in the case of the single specimen of Monarda n. sp. 1) in the UPGMA but have a limited number of OTU’s that are overlapping in the PCoA. Even considering this overlap in the PCoA analysis these five taxa warrant species status as they all have unique, identifying morphological features (discussed below). Monarda n. sp. 1 is most distinct from the other members of Monarda section Aristatae by virtue of it being densely canescent on all vegetative surfaces. This gives the taxon a distinctly grey appearance. It also has the shortest calyces (<5 mm long) (Fig. 11) in the section and bears further differences from the taxon it overlaps (M. austromontana Epling) in bract shape and pigmentation characters. Monarda humilis has the shortest upper corolla lips of the section (Fig. 28), and is distinguished from the taxon it overlaps (M. occidentalis ined) by several additional features. It has smaller leaves and teeth (Fig. 23, 24), and a smaller length of the margin that is 60 serrated (Fig. 23). It also has shorter upper bracts (Fig. 24), wider calyx lobes (Fig. 26) and wider mid (Fig. 26) and lateral (Fig. 27) limbs. Monarda mexicana has very finely serrulate leaf (Fig. 8) and bract (not shown) margins, the widest (> 15 mm) lower bracts in the subgenus (Fig. 22), and wide (>1.7 mm) calyx lobes (Fig. 26), and lower corolla lips (Fig. 26). Monarda stanfieldii has long petioles (Fig. 22), wide leaves with large dentate margins (Figs. 23, 24), and has flowers with long calyces (Fig. 25) and narrow lower corolla lips (Fig. 26). Monarda occidentalis ined is a more heavily branched species (as is M. humilis (Fig. 25)), lacking in calyx orifice hairs (Fig. 25), with smaller lower corolla lip limbs (Fig. 27) and narrow upper corolla lips (Fig. 28). There are some additional features that are distinctive of M. stanfieldii, M. occidentalis ined, and M. humilis that proved to be too problematic to include in this investigation. The first is the uniquely dense calyx orifice hairs seen in M. stanfieldii which were excluded, along with all density measures of indument, due to difficulty of data collection. The second is the large number of flowers clustered in each verticilaster of M. occidentalis ined. I refrained from counting this for each OTU as the sampling would require the destruction of entire glomerules for each specimen included in the analysis. 61 Finally, I avoided collecting data pertaining to color and pigmentation patterns between taxa, as age differences, drying methods, and other discrepancies between the handling of the herbarium specimens would lend to differential fading of the pigments. However, these traits are often invoked in separating the taxa in this section, and a good example is in separating M. humilis from its closest geographic neighbor, M. occidentalis ined. The former has purple floral bracts and white corollas with purple spots whereas the later has white floral bracts and yellow corollas with purple spots. These three species also have somewhat distinct distributions. M. occidentalis has the largest geographic distribution, extending south from southern Kansas through western Oklahoma across western Texas and eastern New Mexico and down into northern Mexico. M. stanfieldii is found in central Texas, east of M. occidentalis, and M. humilis is found in New Mexico on the western margin of the distribution of M. occidentalis. TAXA LACKING SUPPORT FROM EITHER ANALYSIS. Though individuals of M. citriodora var. parva clustered together in the UPGMA, the branch that separated them from M. citriodora var. citriodora was very shallow (larger coefficient of similarity between those 62 groups of OTU's). In fact, there are some branches of M. pectinata and M. austromontana that are similarly long. Their positions in the PCoA analysis were quite close to them as well. Since there weren’t any solid morphological traits that corroborate species recognition, the conservative conclusion is that they remain part of M. citriodora. With the exception of the M. punctata var. arkansana in group 3 (Fig. 18, discussed in detail below), there are large clusters in both analyses formed of varieties of M. punctata. Because of this I maintain that those taxa indeed belong to the species M. punctata and stress that it is a highly variable species. To further highlight this variation I make note of eight additional specimens of M. punctata that came out in unexpected locations along the dendrogram. Four of them (lasio61, punc298, punc160, punc159) show up in group 3 (Fig. 18) with M. arkansana ined, and four (corr464, corr470, corr471, lasio70) show up in group 2 (Fig. 16) with M. occidentalis ined. One possible explanation for those specimens to come out with M. occidentalis ined is because they have abnormally large (compared to the other taxa of M. punctata) bract dimensions. Similarly, the specimens that come out with M. arkansana ined have larger leaf dimensions, which they have in common with M. arkansana. However, in the PCoA they are clustered with the 63 rest of M. punctata, reinforcing their placement in that species. Of additional interest in M. punctata (especially evident in the UPGMA) were the two groups of M. punctata var. arkansana. One group (Fig. 17) shared a branch with M. punctata, and the other (Fig. 18) shared one with M. villicaulis. Close inspection of the specimens and Scora’s representative specimen list revealed that this pattern is an artifact of Scora being unfamiliar at the time of his publication with the variation in stem pubescence found in M. punctata var. punctata or of that taxon’s wider geographic distribution. In his key (which was used in assigning the specimens in this study to taxon) he separates M. punctata var. arkansana from M. punctata var. punctata based on the former as having stems with many horizontal bristles, leaf blades from 60 - 80 mm long, and being from Arkansas, whereas he recognized the latter as having stems with few horizontal bristles, leaf blades from 40 - 50 mm long, and being from the Atlantic and Gulf seaboards. The M. punctata var. arkansana individuals that share the branch with the remaining varieties of M. punctata (Fig. 17) are in fact M. punctata var. punctata that are found in Arkansas, with stems having many horizontal bristles. In fact, Scora examined the specimens in 1984 and annotated them as such. 64 Based on the cluster analyses and some unexpected morphological distinctiveness identified by the boxplots, I have elected to elevate M. punctata var. arkansana to M. arkansana ined. The species has the longest upper corolla lips (Fig. 28) of section Cheilyctis, as well as the longest lower corolla mid limb (Fig. 27) and upper corolla limb (Fig. 28). It also is distinguished from M. punctata, expressly, by its larger leaves and more densely pubescent surfaces (though there are specimens of M. punctata var. punctata that are exceptions to this). Despite the differences between M. arkansana and the rest of section Cheilyctis, I would still like to stress that this is a tentative conclusion, because the levels of variation of these traits are quite high within M. punctata. In summary of the changes to my proposed taxonomy, two new species (Monarda n. sp. 1, Monarda n. sp. 2) will be recognized and three new combinations (M. arkansana ined, M. occidentalis ined, and M. villicaulis ined). The following is my final taxonomic list of the species in Monarda subgenus Cheilyctis: Section Aristatae Monarda n. sp. 1 M. austromontana Epling M. citriodora Cervantes ex Lagasca 65 M. clinopodioides Gray M. pectinata Nuttall M. mexicana Epling Section Cheilyctis Monarda n. sp. 2 M. arkansana ined M. fruticulosa Epling M. humilis (Torrey) Prather & J.A. Keith M. maritima (Cory) B.L. Turner M. occidentalis ined M. punctata L. M. stanfieldii Small M. villicaulis ined M. viridissima Correll Contribution of Morphological Traits to the Taxonomy of Monarda subgenus Cheilyctis The traits most commonly employed for the circumscription of the species of this subgenus have been vestiture, leaf shape and dimensions, calyx shape and dimensions, and corolla pigmentation. Though calyx orifice hair density and corolla pigmentation were not examined in this analysis, there were a 66 number of the above characters that were found to be unique to a single taxon and helped in circumscribing my recognized taxa. Among these were stem and leaf pubescence as well as leaf dimensions. Below is a brief review of how these characters were used in previous treatments as well as a discussion of new insights into these characters and their impact on the taxonomy of this group. I also address additional characters from this study that showed promise for taxonomic utility. These include petiole length, leaf tooth size, length of serrated margin, and bract dimensions. Vestiture. Indumentum has played an important role in the taxonomy of this section from at least the inception of the first subspecies of M. punctata, M. punctata. subsp. villicaulis by Francis W. Pennell (Pennell 1919). Its utility stems from variation in hair type (bristly, canescent, ciliate, strigose, tomentose, villous, puberulent) density (dense, sparse, spreading) angle (down swept, horizontal, upswept) and placement (on stems, nodes, petioles, ab- and adaxial leaf surfaces, leaf margins, midveins, floral bracts, calyx lobes, calyx orifices), and has been invoked among various taxonomic keys for almost every taxon. Their treatment has been inconsistent, however, and there is some disagreement on aspects such as hair type and 67 location as well as how often it occurs across a given taxon and their overall importance as identifying traits. Epling (1935), McClintock & Epling(1942), Scora (1967), and Prather & Keith (2003) agree that M. fruticulosa has a canescent indument, whereas Turner (1994) refers to it as densely strigose. M. villicaulis ined has spreading hairs throughout according to Pennell (1919), but has a thinly pubescent upper leaf surface and dense covering of thin hairs along the lower leaf surface by McClintock & Epling (1942), Epling (1935), and Scora (1967). M. maritima has pilose indument upon the stem and leaf surfaces according to Turner (1994) and Prather & Keith (2003) but the taxon, sensu Scora (1967), has bristles on the stems and petioles and elongated stiffish hairs on the abaxial leaf surface. Leaf Dimensions. Most treatments of these taxa include circumscriptions based in part on leaf size and shape. McClintock & Epling (1942) treated M. fruticulosa as having narrow (not wider than 3 mm) linear leaves, and they treated the leaves of M. punctata var. immaculata as linear-lanceolate. Though Scora (1967) separated M. punctata vars. fruticulosa and immaculata from the other taxa in this section by their linear leaves, Turner, while not recognizing M. punctata var. immaculata, elevated M. punctata var. fruticulosa to M. 68 fruticulosa and also segregated it from the rest of the taxa based specifically on their narrow leaves. Leaf size is also a vital feature in the identification of other species in the subgenus as well. M. microfolia is readily identified by its small leaves, and M. arkansana ined by its larger leaves. Calyx lobe dimensions. Calyx lobe shape differences have also been used frequently to differentiate taxa in this section. The most prominent differences in this trait come from comparing the two sections. In fact, a defining feature of Monarda section Aristatae is the presence of long aristate calyx lobes, (vs. the shorter wider lobes of Monarda section Cheilyctis). Even within the sections, however, there is notable variation in these traits. Both Scora (1967) and McClintock and Epling (1942) separate M. punctata var. occidentalis from the other varies by their widely deltoid calyx teeth. In his taxonomic keys, Turner (1994) distinguishes M. punctata var. punctata from M. punctata var. correllii by the narrowly deltoid calyx lobes of the former and the broadly deltoid calyx lobes of the later. He also uses the ―broadly acute‖ calyx lobes of M. punctata var. occidentalis to distinguish it from M. punctata var. intermedia, which he indicates as having ―narrowly acute‖ lobes. The current treatment recognizes two species based, at least in part, on calyx lobe shapes. The first is M. occidentalis ined (for 69 reasons described above), the second is M. mexicana which demonstrates wider calyx lobes than any of the other species in Monarda section Aristatae. Petiole length. M. fruticulosa is without (or with a very minute (<0.2 mm)) petioles and M. maritima has petioles from 0.5 to 2.1 mm. The rest of the taxa have longer petioles (> 3 mm). Leaf tooth size. While a few taxa in Monarda section Cheilyctis (M. fruticulosa, M. humilis, M. microfolia, and M. viridissima) have teeth <0.3mm wide, most taxa in this section have teeth from 0.3 mm – 0.7 mm in width. Monarda stanfieldii, however, have the largest teeth from 0.8 - 1.3mm. Monarda mexicana, from Monarda section Aristatae has distinctly serrulate margins, unlike the larger teeth of the rest of that section. Length of serrated margin. The only taxon with definable differences in length of serrated margin is M. fruticulosa. In this species the serrated region of the leaves are smaller (<20% vs. >30%) than in the other taxa of section Cheilyctis. Bract dimensions. Inflorescence bracts have long been a main feature that taxonomists have used in developing a circumscriptive structure for the species in Monarda L. There is striking variability across the genus including shape, number, color, and degree of differentiation from the leaves. 70 In Monarda subgenus Cheilyctis there are two morphologically distinct types of bracts: the two basal, more foliar bracts, and the many upper bracts that are reduced in shape and different in color than the either the lower bracts that subtend them, or the true leaves of the plants. Between the sections one can compare the aristate tips of the upper bracts of Monarda section Aristatae with the acute to acuminate tips of the bracts of Monarda section Cheilyctis. There are also species that are noted for their bract color such as M. humilis and M. occidentalis ined (see taxonomic conclusions above) and those noted because of distinctive bract shapes, such as M. mexicana which has very wide lower bracts (also discussed above). 71 Chapter 3 MONOGRAPH OF MONARDA SUBGENUS CHEILYCTIS 72 Introduction Monarda subgenus Cheilyctis Rafinesque is comprised of 16 species distributed across the eastern and central United States and northern Mexico. In the U.S. it extends west into New Mexico and Colorado and in Mexico as far south as Hidalgo. Characteristically, they are herbaceous perennials and annuals that grow upright, branch at the base, and sometimes appear shrubby. Leaf arrangement is opposite and decussate. They have distinctly bilabiate flowers and a unique inflorescence comprised of sessile cymes (glomerules) that are subtended by two sets of bracts: a lower, petioled set and an upper, sessile set. The lower bracts are usually two in number and appear mostly green, whereas the upper set can occur as four to more than 40 in number and appear either green or highly colored. The species of subgenus Cheilyctis are distinguished from those of M. subgenus Monarda Scora by their taproots, strongly arching upper corolla lips, inserted stamens, and verticillastrate glomerules. Also, the species of M. subgenus Cheilyctis have a variable chromosome number (see below) whereas M. subgenus Monarda have n=18 chromosomes (or 16 in M. media). The subgenus has two sections, Cheilyctis and Aristatae. The species in Monarda section Cheilyctis are distinguished morphologically from those in section Aristatae by differences 73 in calyx lobe shape, calyx venation, and indumentum of the calyx opening (see morphology section for a discussion of these). The chromosome number of taxa in Monarda section Cheilyctis are n=11 (or 12 in M. humilis and M. villicaulis) while those in section Aristatae are n=9 (or 18 in M. pectinata). The taxa of Monarda section Cheilyctis are distributed from central U.S. to the eastern seaboard, with the majority of the species occurring in the states of Arkansas, Oklahoma, and Texas. They are often found in sandy soils on plains, prairies, beaches and river banks. The taxa of Monarda section Aristate are distributed across the southwestern United States and northern Mexico, with M. citriodora extending east into Florida, and M. pectinata extending north into Nebraska, across Colorado, and west into Arizona. The extent of the distributions of the species in this subgenus is highly variable. Some species, such as M. punctata and M. citriodora are found widespread across much of the distribution of the subgenus while others, such as M. fruticulosa and M. stanfieldii, are narrow endemics. This pattern is especially pronounced in section Cheilyctis. Several species, including M. fruticulosa, M. maritima, M. stanfieldii, and M. viridissima fit this distribution profile and are recognized (Keith 2003) as edaphic endemics. 74 Taxonomic history The taxonomic rank of many of the species and varieties of this subgenus has been a continued matter of debate for over eight decades (Epling 1935; McClintock and Epling 1942; Scora 1967; Turner 1994; Prather and Keith 2003). Monarda stanfieldii was once considered a subspecies of M. punctata by Epling (1935), a variety of M. punctata by Scora (1967), and not until 1994 did it achieve its currently accepted status of species by Turner (1994). Monarda punctata var. lasiodonta was considered a synonym of M. punctata var. occidentalis by Epling (1935), but both were treated as distinct subspecies by Mclintock and Epling(1942) and varieties by Scora (1967). Though these are only two examples, the taxonomic literature concerning Monarda subgenus Cheilyctis is replete with many more of this nature. To date, the most comprehensive treatments of this group were two reviews of the entire genus by McClintock & Epling (1942) and Scora (1967) who recognized six and five species of M. subgenus Cheilyctis, respectively. The objective of this monograph is to provide a current, in-depth treatment of this subgenus, including taxonomic changes and novelties that have been published since Scora’s 1967 monograph, as well as incorporate the extensive morphometric analyses performed as part of this dissertation (Chapter 2). 75 Morphology Duration/Vegetative Habit. With one exception (Monarda n. sp. 1) all the species in section Aristatae are annuals. The majority of the species from section Cheilyctis, however, are perennial, the annual species being M. humilis, M. occidentalis ined., M. punctata (which has some populations that are perennial), M. stanfieldii, and M. villicaulis (which may also have populations that are perennial). The perennial species can take on woody characteristics, and the more heavily branched taxa (M. fruticulosa, M. maritima, M. occidentalis, M. stanfieldii) can appear shrubby. Roots. Both annual and perennial species (see above) have a short tap root with numerous fibrous secondary roots which comprise the majority of the root system. The roots vary in size among the species, with the perennial taxa usually demonstrating larger tap roots than those that are annual. Stems. The stems are quadrangular and variously pubescent (see vestiture section) or glabrous. They have an upright growth pattern and may branch primarily at the base (M. maritima, M. pectinata), the middle region of the stem (M. arkansana, Monarda n. sp. 1, Monarda n. sp. 2, M. stanfieldii, and M. villicaulis ined), the apical region of the stem (M. austromontana and M. citriodora), or, most commonly, a combination of these. 76 Each species may also develop short (< 1 cm), axillary, lateral shoots which bear leaves in tightly clustered fascicles. Leaves. Species of subgenus Cheilyctis have leaves that are simple, with an opposite and decussate phyllotaxis, though leaves borne on the short axillary branches appear fascicular. Petiole length varies from sessile (M. fruticulosa) to 25 mm (M. stanfieldii). Leaf size and shape are useful characteristics for identifying taxa that have particularly small features (e.g. M. fruticulosa has small (10 - 40 mm long) linear leaves and Monarda n. sp. 2 has small (10 - 15 mm long) ovate to orbicular leaves. The remaining taxa have leaves that vary from lanceolate to ovate. Leaf margins are entire (in some M. fruticulosa) or serrate. The serrations do not always occur along the entire margin, and when present occur at the tip and extend from 15% (M. fruticulosa) to 100% (M. arkansana ined. and M. punctata) of the distance down the blade toward the base of the leaf. The widest part of the leaf occurs toward the middle of the leaf in some taxa and closer to the base of the leaf in others. The leaf bases range from acute to acuminate and are frequently ciliate. Vestiture. The vestiture of subgenus Cheilyctis consists of sessile glands and uniseriate non-glandular trichomes of various sizes and appearance (Fig. 29). The glands are usually sessile 77 and appear as punctate dots that are colored or translucent on the adaxial surface of the leaves and floral bracts and occasionally on the stems. The non-glandular trichomes range from short, rigid, unicellular hairs commonly found covering virtually all the plant surfaces of most species in this subgenus, to multicellular hairs (three to seven cells) that are clustered about the nodes, petioles, distal ridge of the upper corolla lobe, and on the margins of leaves, floral bracts, calyx opening (orifice), and calyx lobes. The multicellular trichomes can be curly or wavy forming a villous surface, or straight and flexible forming a pilose surface. Inflorescence. The inflorescences in this subgenus are very distinctive, as they are composed of multiple, axillary verticillate glomerules. They are important in distinguishing subgenus Cheilyctis from subgenus Monarda, as the latter has glomerules that are solitary and terminal, though variation among the species of this section is limited. 78 Figure 29. Trichome types. A. Unicellular hairs along the stem of M. punctata (Correll 18925, TEX). B. Multicellular hairs on the pilose nodal region of M. maritima (Johnston 53254.22, TEX). C. Glandular punctations on a canescent adaxial leaf surface of M. fruticulosa (Garcia 110, TEX). D. Villous abaxial leaf surface of M. villicaulis ined. (Bourdo 306, MSC). For interpretation of the references to color in this and all other figures, the reader is referred to the electronic version of this dissertation. 79 Inflorescence bracts. There are two main types of bracts that subtend the glomerules. The lowermost bracts are decussate to the distal most pair of leaves and are always two in number. They are usually green with a short petiole and often resemble the shape of the adjacent leaves, although they are usually reduced slightly in size, and can be variously pigmented. The blades vary from lanceolate to ovoid-lanceolate and their margins are entire or serrate. The widest part of the bract occurs toward the middle in some taxa and closer to the base in others. 40. The uppermost bracts vary in number between four and They vary in color from green to white to red and are sessile in either a single or several whorls. In section Cheilyctis the upper bracts are lanceolate with acute tips and are positioned in an upright to horizontal orientation. The bracts usually occur in a single whorl with <10 bracts per glomerule. In section Aristatae the upper bracts range from lanceolate to linear, are aristate, and have a horizontal to reflexed orientation. They usually occur in several whorls with >15 bracts per glomerule. Calyx. The orifice can be glabrous or diffusely (as in most species of section Cheilyctis) to densely bearded (species in section Aristatae and M. stanfieldii) with flexuous to stiff 80 white multicellular hairs. The lobes have varying shapes from deltoid to lanceolate and can be covered in hairs of varying length and density (Fig. 30). The calyx lobe tips also vary from acute (species in section Cheilyctis) to aristate (species in section Aristatae). Calyx indumentum and lobe shapes are commonly used in taxonomic keys (Scora 1967, Turner 1994, Prather 2003). Corolla. The two petals that form the upper lip may or may not form a minute cleft at the distal end of the lip. The lower corolla lip, formed from the fusion of the remaining three petals, has three distinct lobes: a middle lobe and two shorter lateral lobes. Between species the lobes vary in length (0.2-2 mm) and shape (tapered to clubbed) and (especially the middle lobe) may be reflexed upward relative to the horizontal portion of the lower corolla lip. Corolla color can be white, crème, yellow, or a range from pink to red, and can have white, yellow, or maroon spots. 81 Figure 30. Calyx lobe shapes and vesture types. A. M. clinopodioides (Lawson 40, OKL) with aristate calyx lobes and long lobe margin hairs; B. M. humilis (Higgins 7737, NMC) with acute calyx lobes and glabrous margins; C. M. austromontana (Maysilles 8272, TEX) showing sparsely packed, short calyx lobe margin hairs; D. M. maritima (Churchill 90-930, MSC) with lanceolate calyx lobes; E. M. citriodora with orifice and calyx tube hairs. 82 Species Concept Species discussed in this manuscript are defined according to the phenetic species concept (PSC)(Sokal and Crovello 1970, DeQueiroz 2007, McCauley 2007). They represent morphologically cohesive groups that are delineated by quantitative or qualitative gaps. Furthermore, intraspecific taxa are not formally treated (they are not given their own taxonomic descriptions, synonymies, etc.) in this monograph, as any significant morphological differences between them and other taxa would warrant species recognition. Instead, in the respective species descriptions I discuss any putative variation and specify the names and intraspecific ranks that are historically attached to the individuals that demonstrate that variation. Phylogeny A phylogenetic hypothesis for all 16 species of Monarda subgenus Cheilyctis is lacking. Prather et al (2002) presented an ITS phylogeny for the genus Monarda but included only six species from subgenus Cheilyctis. Likewise, for her master’s 83 thesis, Kieth (2003) examined patterns of speciation (using molecular phylogeny) in section Cheilyctis, but examined only six of the ten currently recognized species of that section. Therefore, a cladistics analysis was conducted to determine species relationships, provide an evolutionary framework for examining trait distributions, and identify autapomorphies for all currently recognized species of subgenus Cheilyctis. Cladistic analysis of Monarda subgenus Cheilyctis All 16 species of subgenus Cheilyctis were included in the analysis. The outgroups (M. bradburiana and M. russeliana) were selected based on their positions in the molecular phylogeny presented in Prather et al. (2002). Forty-three qualitative morphological characters were identified and assigned discrete states (table 1, 2) based on direct observations of herbarium specimens (see table 2 for list) made during the morphometric analysis discussed in Chapter 2. PAUP 4.0b10 (Swofford 2002) was used to apply the parsimony optimality criterion. All characters were given equal weight and treated as unordered, and multistate taxa were treated as polymorphic. Gaps were treated as missing data. Only 34 characters were parsimony informative (nine were autapomorphic). The heuristic search option was performed and resulted in four equally most parsimonious trees of 114 steps, consistency 84 indices (CI) of 0.711, and a retention indices (RI) of 0.708 (both CI and RI calculated excluding uninformative characters). Stability of the internal clades was tested with a bootstrap analysis of 1000 replicates. A representative cladogram (Fig. 31) and semi-strict consensus (Fig. 32) tree are presented. A semi-strict consensus tree was used to provide better resolution of section Aristatae because it incorporates topologies that are not contradicted by other equally parsimonious trees, though topological differences between the strict and semi-strict consensus trees are discussed below). The semi-strict consensus tree reveals two basal clades which correspond to Monarda sections Cheilyctis and Aristatae. There is strong support (BS = 99%) for the monophyly of the subgenus. Within section Aristatae the only relationship that contains bootstrap support >60% is between M. clinopodioides and M. pectinata (―4‖ from Fig. 32; BS = 64%). The remaining clades have bootstrap values <50%. The M. clinopodioides clade is defined by pectinate calyx lobe hairs and villous petioles. Another clade consists of M. austromontana and Monarda n. sp. 1 (―5‖ from Fig. 32; BS = <50%)) which share the apomorphies of lower bracts reflexed and lower bracts lanceolate. Monarda mexicana is potentially sister to all the remaining species of the section. The only topological differences between the four parsimony-equal trees for section Aristatae are the 85 relationships of M. citriodora to the above mentioned clades. In two of the trees it is sister to the M. clinopodioides/M. pectinata and M. austromontana/Monarda n. sp. 1 clades, and in the other two trees it is an unresolved polytomy with those two clades. Because of the latter, the difference between the semi- strict consensus and the strict consensus tree is the inclusion of a resolved M. citriodora in the semi-strict tree, instead of a polytomy in the strict consensus. In section Cheilyctis three clades are consistently identified among the most-parsimonious trees. The first (―1‖ from Fig. 32; BS = 71%)) is a clade of species with linear to linear-lanceolate leaves from south-central to southern Texas consisting of M. fruticulosa, M. maritima, and M. viridissima. The second (―2‖ from Fig. 32; BS = 73%) is a clade of species with glabrous calyces consisting of M. humilis, M. occidentalis, and M. stanfieldii. Monarda punctata is sister to the third (―3‖ from Fig. 32; BS = 75%) clade, which consists of species with villous indumentum on at least the abaxial leaf surfaces. This clade consists of M. arkansana, Monarda n. sp. 2, and M. villicaulis. The only topological differences between the four parsimony-equal trees for section Cheilyctis are the relationships of M. humilis, M. occidentals, and M. stanfieldii. In two of the trees M. stanfieldii is sister to M. humilis and M. occidentalis, and in the other two trees M. humilis is sister 86 to M. occidentalis and M. stanfieldii. For the representative tree I chose the phylogeny which depicts M. occidentalis as sister to M. humilis, but caution that their precise phylogenetic placement is not supported. Furthermore, the low bootstrap values (Fig. 32) for most of the nodes of the cladogram is a sign of concern, and is probably a result of the small data set relative to the number of taxa included in the analysis, as well as a moderate level of homoplasy. It is noteworthy that this current morphological phylogeny is largely congruent with the phenetic dendrogram presented in the previous chapter. Though evolutionary relationships should not be inferred from the dendrogram, it depicts many of the same relationships as seen in the current phylogeny. The only notable difference is the placement of Monarda n. sp. 2 with M. villicaulis and M. arkansana in the phylogeny, where it shares a branch with M. viridissima in the phenogram. The morphological phylogeny is also consistent with the subgenus Cheilyctis portion of the ITS phylogeny presented in Prather et al. (2002). Though they included only six species from this subgenus, their phylogeny depicts M. clinopodioides and M. pectinata as sister taxa, with M. citriodora as sister to that clade. The current phylogeny includes additional species, but the position of M. citriodora proximal to a clade containing M. clinopodioides and M. pectinata does not conflict with that 87 presented by the ITS tree. In the current phylogeny a clade consisting of M. austromontana and Monarda n. sp. 1 (not included in Prather (2002)) is sister to the M. clinopodioides clade, with M. citriodora sister to a clade encompassing all the above-mentioned taxa. The relationship between M. fruticulosa, M. viridissima, and M. punctata follow this same pattern. Both the ITS and this morphological phylogeny depict M. punctata proximal to a clade containing M. fruticulosa and M. viridissima. Though Keith (2003) did produce multiple cladograms based on ADH1 and ADH2, ncpGS, her objectives weren’t strictly consistent with those of the present phylogeny and thus the two should not be compared. 88 Figure 31. One of four equally most parsimonious cladograms for Monarda subgenus Cheilyctis. 89 Figure 32. Strict consensus tree of Monarda subgenus Cheilyctis derived from four equally parsimonious cladograms (114 steps, CI = 0.711, RI = 0.708). Taxonomic sections are indicated by vertical bars, and numbers above branches indicate bootstrap values. Outgroups are M. bradburiana and M. russeliana. 90 Table 1. Characters used in the phylogenetic analysis of Monarda subgenus Cheilyctis. 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. Stamens exerted (0); stamens inserted (1) Glomerules terminal (0); glomerules verticillastrate (1) Corolla weakly arched (0); corolla strongly arched (1) Leaf shape lanceolate (0); leaf shape linear (1); leaf shape linear-lanceolate (2); leaf shape ovate (3) Leaf tip acuminate (0); leaf tip acute (1) Leaf base round (0); leaf base acute (1); leaf base acuminate (2) Leaf margin serrate (0); leaf margin subentire (1) Leaf attachment petiolate (0); leaf attachment Subsessile (1); leaf attachment sessile (2) Upper bract tip shape acuminate (0); upper bract tip shape acute (1); upper bract tip shape aristate (2) Upper bract shape elliptical (0); upper bract shape lanceolate (1); upper bract shape oblong (2) Calyx lobe shape aristate (0); calyx lobe shape acuminate (1); calyx lobe shape lanceolate (2); calyx lobe shape acute (3) Abaxial leaf surface without minute (<3 cells) hairs (0); adaxial leaf surface with minute hairs (1) Abaxial leaf surface without long (>5 cells) hairs (0); adaxial leaf surface villous with long hairs (1); adaxial leaf surface pilose with long hairs (2) Midvein without minute (<3 cells) hairs (0); midvein with minute hairs (1) Midvein without long (>5 cells) hairs (0); midvein villous with long hairs (1); midvein pilose with long hairs (2) Petiole without minute (<3 cells) hairs (0); petiole with minute hairs (1) Petiole without long (>5 cells) hairs (0); petiole villous with long hairs (1); petiole pilose with long hairs (2) Interpetiolar plane without minute (<3 cells) hairs (0); interpetiolar plane with minute hairs (1) 91 Table 1 continued. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. Interpetiolar plane without long (>5 cells) hairs (0); interpetiolar plane villous with long hairs (1); interpetiolar plane pilose with long hairs (2) Nodal ridge without minute (<3 cells) hairs (0); nodal ridge with minute hairs (1) Nodal ridge without long (>5 cells) hairs (0); nodal ridge villous with long hairs (1); nodal ridge pilose with long hairs (2) Internode without minute (<3 cells) hairs (0); internode with minute hairs (1) Internode without long (>5 cells) hairs (0); internode villous with long hairs (1); internode pilose with long hairs (2) Calyx surface without minute (<3 cells) hairs (0); calyx surface with minute hairs (1) Calyx surface without long (>5 cells) hairs (0); calyx surface with long hairs (1) Plant perennial (0); plant annual (1) Calyx lobe cilia absent (0); calyx lobe cilia short (<3 cells) (1); calyx lobe cilia long (>5 cells) (2) Calyx lobe cilia density sparse(0); calyx lobe cilia density pectinate (1) Calyx lobe face without long (>5 cells) hairs (0); calyx lobe face with long hairs (1) Calyx orifice hair density sparse (<5 hairs) (0); calyx orifice hair density medium (>10 hairs <20); calyx orifice hair density dense (>40 hairs) Lower bract color green (0); lower bract color white (1) Upper bract color purple (0); upper bract color green (1); upper bract color white (2) Lower bract margin serrate (0); lower bract margin entire (1); lower bract margin serrulate (2) Upper bract cilia density sparse (<3 hairs)(0); upper bract cilia density dense (>6) hairs (1) Upper bract cilia position basal only (0); upper bract cilia position along the entire margin (1) 92 Table 1 continued. 36. 37. 38. 39. 40. 41. 42. 43. Upper bract margin entire (0); upper bract margin serrulate (1) Upper corolla lip without spots (0); upper corolla lip with spots (1) Lower corolla lip without spots (0); lower corolla lip with spots (1) Corolla color white, purple, or pink (0); corolla color yellow (1) Calyx color purple (0); calyx color green (1) Upper corolla lip without long (>5 cells) hairs (0); upper corolla lip with long hairs (1) Lower bracts strongly reflexed (0); lower bracts horizontal (1) Lower bracts linear (0); lower bracts lanceolate (1) 93 Table 2. Data matrix of characters used in the phylogenetic analysis of Monarda subgenus Cheilyctis and two outgroups (M. bradburiana and M. russeliana). The character number matches those presented in table 1. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 94 4 2 2 1,2 0 0 3 0 0 0 0 0 0 0 0 0 0 5 0 0 0 0 0 0 0,1 1 1 0 0 1 0 1 1 1 6 2 2 2 1 1 1 1,2 2 2 2 2 2 2 1 0 0 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 7 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 8 1 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 9 0 0 0 0 0 0 0 1 2 2 2 2 2 0 0 0 95 10 0 1 0 1 1 1 0 0 1 3 1 1 1 1 0 0 11 2 2 1 3 3 2 2 1 0 0 0 0 0 0 0 0 12 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 13 1 1,0 0 0 0 0 1,0 1 0 0 0 0 0 0 0 0 14 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 15 1 1 0 0 0 0 1,0 1 0 0 0 0 0 0 0 1 96 16 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 17 1 1 0 0 0 0 1,0 1 1 0 0 1 0 0 0 1 18 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 19 1 1,0 0 0 0 0 1,0 1 1 0 0 0 0 0 1 1 20 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 21 1 0 0 0 0 0 1,0 1 0 0 0 0 0 0 0 0 97 22 1 0,1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 23 1 0 0 0 0 0 1,0 1 0 0 0 0 0 0 0 0 24 1 1 1 0 0 0 1,0 1 1 1 1 1 0 1 1 1 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 25 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 26 0 0 0 1 1 1 0,1 0 1 1 1 1 1 0 0 0 27 2 2 2 2 1 2 0,2 2 2 2 2 2 2 1 2 2 98 28 1 1 1 1 1 1 0 1 1 0 0 1 0 0 0 0 29 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 30 1 0 1 1 1 2 1 1 2 2 2 2 1 2 1 1 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 31 0 0 0 0 0 0,1 0 0,1 0 0 0 0 0 0 ? ? 32 0 0 0,2 0 2 1,2 0 0 0 0 0 1 1 1 0 0 33 0 0 0 0 0 0 0 0 0 0 2 1 1 0 ? ? 99 34 1 0 0 0 0 0 0 0 1 1 1 1 1 0 1 1 35 0 0 0 0 0 0 0 0 1 1 1 1 1 0 1 1 36 0 0 0 0 0 0 0,1 1 0 0 0 0 0 0 0 0 Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 37 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 38 1 0 1 1 1 1 1 1 0 0 0 0 1 0 1 1 39 0 1 1 1 1 1 0,1 1 1 1 1 1 1 1 1 1 100 40 1 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0 41 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 0 42 1 1 1 1 1 1 1 1 1 1 0 1 1 0 ? ? Table 2 continued. M. maritima M. viridissima M. fruticulosa M. humilis M. occidentalis M. stanfieldii M. punctata Monarda n. sp. 2 M. clinopodioides M. citriodora M. austromontana M. pectinata M. mexicana Monarda n. sp. 1 M. bradburiana M. russeliana 43 0 0 0 0 0 0 0 0 0 0 1 0 0 1 ? ? 101 Key to the species of Monarda subgenus Cheilyctis 1a. Calyx lobe tips attenuate to aristate, 2a. Lowest inflorescence bracts > 12 mm wide; pollen Yellow; leaf margins serrulate.........4. Monarda mexicana 2b. Lowest inflorescence bracts < 8 mm wide; pollen white; leaf margins serrate 3a. Stems and leaves canescent; floral bracts immediately subtending the calyces with acuminate tips, green................................6. Monarda n. sp. 1 3b. Stems and leaves distinctly green; floral bracts immediately subtending the calyces with narrowly cuspidate to aristate tips, greenish-purple to red. 4a. Floral bracts of many widths, ciliate along the entire margin, strongly reflexed, especially the lowest set; lateral limb of lower corolla lip > 1.5 mm long ...................................1. M. austromontana 4b. Floral bracts of consistent width (may be linearlanceolate or elliptic, but not both), ciliate along the base to mid-margin, horizontally spreading or weakly reflexed; lateral limb of lower corolla lip <1.4 mm long 5a. Floral bracts immediately subtending the calyces > 4 mm wide, abruptly narrowing to a cuspidate tip; stems, bracts, and calyx tubes purple pigmented 102 ....................................2. M. citriodora 5b. Floral bracts immediately subtending the calyces ≤ 4 mm wide, gradually narrowing to a cuspidate tip; stems, bracts, and calyx tubes green (rarely sparsely pigmented with purple).....5. M. pectinata 1b. Calyx lobe tips acute to acuminate 7a. Leaves linear or linear-lanceolate 8a. Fruticose; leaves linear, ≤ 3 mm wide; stems and leaves canescent; petiole absent or minute, < 1 mm long ......................................8. M. fruticulosa 8b. Suffruticose; leaves linear-lanceolate to lanceolate, > 4 mm wide; stems and leaves green; petiole > 1 mm long 9a. Stems, petioles, and leaves densely pilose, calyx orifice glabrous; southern coastal Texas .......................................10. M. maritima 9b. Stems, petioles, and leaves puberulent, calyx orifice pubescent; south central Texas......15. M. viridissima 7b. Leaves lanceolate or ovate 10a. Calyx orifice densely bearded by a cone of white trichomes; leaves broadly lanceolate .....................................13. M. stanfieldii 10b. Calyx orifice glabrous or ciliate, not densely bearded; leaves lanceolate to ovate 11a. Calyx teeth broadly acute; calyx tube glabrous or 103 with minute hairs on (but not between) the veins; flowers lavender, white, or cream 12a. Floral bracts purple to pink; corollas lavender to purple; calyx orifice pubescent ....................................9. M. humilis 12b. Floral bracts white to green; corollas white to Cream; calyx orifice glabrous ........................11. M. occidentalis ined. 11b. Calyx teeth narrowly acute; calyx tube with scattered hairs between (and on) the veins; flowers yellow 13a. Midvein and laminar portion of lower leaf with short, single-celled hairs or glabrous ..................................12. M. punctata 13b. Midvein and/or laminar portion of lower leaf with longer, villous, multicellular hairs 14a. Leaves 2.2 – 3.5 cm long; upper bract tip acute; calyx lobe tips acuminate ..........................16. Monarda n. sp. 2 14b. Leaves 3.2 – 10cm long; upper bract tip acuminate; calyx lobes lanceolate 15a. Lower leaf surface pubescent throughout; calyx lobe face pubescent; lower corolla lip < 2.8 mm wide..14. M. villicaulis ined. 15b. Lower leaf surface pubescent primarily on 104 and around midvein; calyx lobe face glabrous; lower corolla lip > 3.2 mm wide ......................7. M. arkansana ined. 105 Taxonomy Monarda L. Sp. Pl. 1: 22. 1753. Annual, perennial, or biennial herbs, shrubs, or subshrubs to 120 cm tall; upright, simple or branching from the base, middle of stem, or inflorescence. Vegetative parts aromatic from essential oils carvacrol, monardol, thymol; glabrous or variously pubescent with single and multicellular hairs. Leaves linear to ovate, sessile to petiolate, margins dentate, serrate, serrulate or entire. Inflorescence a solitary terminal glomerule or multiple verticillate glomerules, flowers few to many, subtended by one or two sets of foliar bracts. Flowers bilabiate; lower lip three lobed, the middle lobe longer than the two lateral lobes; upper lip of two equal-sized lobes. Stamens 2 (occasionally two additional aborted stamens), inserted or exerted. Monarda subgenus Cheilyctis Scora, Univ. of Cal. Publ. Botany. 41: - 71. 1967. Monarda section Cheilyctis Rafinesque, Med. Fl. 2: 37. 1830. Monarda section Coryanthus Nuttall, Trans. Am. Philos. Soc.5:186. 1837. –Type species: Monarda punctata. 106 Herbs, shrubs, and subshrubs with taproots. Inflorescence of verticillate glomerules. Flowers with arched upper corolla lips and inserted stamens. N = 9, 11, 12 or 18. Monarda section Aristatae Epling, Univ. Calif. Publ. Botany 20: 186. 1942. Monarda subsection Aristatae Epling, Madroño 3: 26. 1935. –Type species: Monarda citriodora. Annual herbs. Floral bract tips aristate. Calyx lobe tips aristate. Pollen white. N = 9 or 18. 1. Monarda austromontana Epling, Madroño 3: 29. 1935. Monarda citriodora Cerv. ex Lag. subsp. austromontana (Epling) Scora, Madroño 18: 120. 1965. Monarda citriodora Cerv. ex Lag. var. austromontana (Epling) B.L. Turner, Phytologia 77(1): 56. 1994.--TYPE: MEXICO. Chihuahua: on La Bufa Mountain near Cusihuiriachic, 1887, Pringle 1355 (holotype: US!; isotype: NY (digital image)!, F (digital image)!, CAS 107 (digital image)!, US (digital image)!). Habit annual, subshrub to 70 cm tall; upright, branching from the inflorescence and middle of stem. Stem indumentum variable; internode with short hairs, node with short hairs, nodal plane with short hairs. Penultimate vegetative internode 31 - 65 mm long, 0.9 – 2.5 mm wide. Leaves 32 - 72 mm long, 4.1 - 12 mm wide; petiole 0.8 - 10 mm long; serrations 0.34 mm wide; leaf shape lanceolate, base acuminate; margins serrate from apex to 80% length of laminar tissue, apex acuminate; petiole indumentum of long hairs; abaxial surface hirsute with mediumlong hairs around the midvein; adaxial surface glabrous; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode to 60 mm long, 2 mm wide; pubescent. Lower floral bracts 40-110 mm long, 1.2 - 5 mm wide; sessile; lower bract shape linear-lanceolate, base attenuate, margins serrate; apex acuminate; abaxial surface puberulent; adaxial surface puberulent; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface green. Upper floral bracts 8.5 – 15 mm long, 3 - 6 mm wide; upper bract shape linear to linear-lanceolate, base cuneate, margins serrated; apex aristate; abaxial surface pubescent; adaxial surface pubescent; margins ciliate; abaxial surface green; adaxial 108 surface purple (rarely green). Calyx tube 6.4 – 7.2 mm long, 1.71 – 2.65 mm wide, with minute and pilose hairs on outer surface. Calyx lobes 2 – 5.6 mm long, 0.06 – 0.07 mm wide, aristate, with hairs along the margins. hirsute with stiff white hairs. Calyx orifice densely Lower corolla lip 5.5 - 9.2 (10) mm long, 2.1 – 3.2 mm wide; mid limb to 4.5 mm long, 2.7 mm wide, expanding toward the tip; lateral limbs to 1.7 mm long, 1.6 mm wide. Upper corolla lip 4.7 – 7.5 mm long, 2.6 – 3.8 mm wide, lobes to 0.65 mm long and 0.83 mm wide, corolla violet to white. Stamens to 1.7 cm in length, anthers 0.8 long, 0.3 wide, pollen white. Pistil to 17 mm long. N = 9 (Scora 1967). Phenology. Flowering specimens have been collected from July through December. Distribution. In open and rocky areas of the Sierra Madre of Western Mexico and U.S. M. austromontana is most similar morphologically to Monarda n. sp. 1 and M. citriodora. It can be distinguished from the former by its lack of canescence and longer (> 3 cm) lower bracts and the latter by strongly reflexed (vs. horizontally spreading) lower bracts. Bracts shape is also useful in distinguishing M. austromontana from M. citriodora. In M. austromontana the bracts gradually taper to an aristate tip (vs. abruptly tapering as in M. citriodora), and M. austromontana 109 inflorescences can bear both broadly elliptic and linear lanceolate bracts on the same inflorescence, whereas the upper bracts of M. citriodora are of a consistent width. 110 Figure 33. Herbarium specimen of M. austromontana. Van Devender 96-378 (ARIZ). 111 Representative specimens examined: MEXICO. CHIHUAHUA. Proximidades al campo 1, ejido El Largo, Municipio de Madera, Benitez 1984 (MEXU); Aroyo de las Varitas, ejido El Largo, municipio de Madera, Bolanos 1448 (MEXU); in open, partialy grazed area near the airstrip, southeast of Creel, Bye 2684 (MEXU); Bocoyna, ejido San Ignacio Arareco, east of Gonogochic, Bye 8725 (MEXU); NW of Creel, in open rocky area S of the railroad tracks, Bye 1795 (MEXU); Mesa de Horcones, Laferriere 2017 (TEX); Nabogame, Mpio. Temosachi, Jenkins 89-308 (TEX); road to Basaseachic trhough Tomochic, 2.2 km W of Rio Tomochic bridge, at end of pavement just after crossing a small crek on another bridge, Lane 2767 (TEX); Culebra mountains, Martin s.n. (ARIZ); 2 km southeast of Cocheno, LeSueur 887 (TEX); Municipio Ocampo, Parque Nacional de Cascada Basaseachic, Spellenberg 9243 (MEXU); Mpio de Casas Grandes, Tenorio 1773 (MEXU); 42 km WNW of Colonia Juarez in "Canyon de la Piedra cantil", upper part of the Tinaja, Wilson 8457 (TEX); DURANGO. in Mimbres Canyon, 26 miles west of Durango, route 40, Correll 20146 (LL); Memelichic, meadow east of town, Jenkins 89-297 (ARIZ); Quebrada de San Juan, 26 road miles north of railroad at Coyotes, on road to San Luis, Maysilles 8272 (TEX); 2 miles east of Llano Grande 16 miles east of El Salto, 160 meters northeast of KM 1039, Scora 112 2716 (TEX); Sierra Madre Occidental, 5.1 rd. mi. by hwy. 40 SW of El Salto at Aroyo De Agua, Worthington 8910 (TEX); SINALOA. Pmio Badiraguato a 15 km al N. de Surutato rumbo a Sta. Rita, Vega 2541 (MEXU); SONORA. Yecora, Neff 8-17-91-13 (MEXU, TEX); El Tigre Canyon and Mountain above El Tigre Mine, east of Esqueda and Lago Angostura, Turner 2098 (ARIZ); Mesa el Campanero, Arroyo Largo, upper tributary of Barranca El Salto, Van Devender 96-378 (ARIZ); U.S.A. ARIZONA. Navajo: Alongsie Fish Hatchery road near junction with Diamond Creek road; Fort Apache Indian Reservation, Granfelt 6-224 (ARIZ). 2. Monarda citriodora Cervantes ex Lagasca Gen. Sp. Nov. 2. 1816. --TYPE: MEXICO. Based on specimens grown from seed sent from Cervantes from unpublished locality and date(lectotype: BM; lsolectotype: G-DC). Monarda citriodora Cervantes ex Lag. var. parva Scora, Madroño 18: 120. 1965.—TYPE: U.S.A. Texas: San Patricio Co., near Sinton, Rowell 4977 (holotype: WWF). Monarda citriodora Cervantes ex Lag. var. attenuata Scora, Madroño 18: 121. 1965.—TYPE: MEXICO. Coahuila: near 113 Musquiz, 1963, Scora 2340 (holotype: MICH; isotype: UCR). Monarda dispersa Small, Fl. S.E. U.S. 138. 1903.--TYPE: U.S.A. Missouri: Eagle Rock, no date, Bush 122 (holotype: NY (digital image)!). Monarda aristata Nutt., Trans. Amer. Phil Cos. 5: 186. 1837. --Type: based on a collection from ―Arkansa,‖ no date, Nuttall s.n. (holotype: BM). Habit annual, subshrub to 75 cm tall; upright, branching heavily from the inflorescence and rarely from the base or middle of stem. Stem indumentum variable; internode with short hairs densely packed, node with short hairs, nodal plane with short hairs. Penultimate vegetative internode (20) 40 – 80 (150) mm long, 0.8 - 3.5 mm wide. Leaves 30 - 80 mm long, 3 - 9 mm wide; petiole 3 – 12 mm long; serrations 0.2mm wide; leaf shape lanceolate, base acuminate; margins serrate from apex to 95% length of laminar tissue, apex acuminate; petiole indumentum of long hairs; abaxial surface hirsute with medium-long hairs around the midvein; adaxial surface glabrous; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode to 40 mm long, 0.7-1.1 mm wide; glabrous to pubescent with minute downward curled hairs. 114 Lower floral bracts 12 - 37 mm long, 2.2 - 9 mm wide; sessile; lower bract shape lanceolate, base attenuate, margins serrate; apex acuminate; abaxial surface glabrous; adaxial surface glabrous; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface green. Upper floral bracts 9 – 21 mm long, 3.2 – 7.1 mm wide; upper bract shape oblanceolate, base cuneate, margins serrate to entire; apex aristate; abaxial surface puberulent; adaxial surface puberulent; margins ciliate; abaxial surface green; adaxial surface purple (rarely green). Calyx tube 7.2 – 8.4 mm long, 1.4 – 2.4 mm wide, with hairs of various sizes on outer surface. Calyx lobes 1.2 – 4.6 mm long, 0.1 – 0.2 mm wide, aristate, with long hairs along the margins. Calyx orifice densely hirsute with stiff white hairs. Lower corolla lip 3.2 – 9.8 mm long, 1.7 – 3.5 mm wide; mid limb to 3.2 mm long, 2.1 mm wide, expanding toward the tip; lateral limbs to 1.8 mm long, 0.8 mm wide. Upper corolla lip 6.8 – 9.2 mm long, 2.7 – 7.1 mm wide, lobes to 0.8 mm long and 0.8 mm wide, corolla violet to white. Stamens to 18 mm in length, anthers 0.8 long, 0.3 wide, pollen white. Pistil to 18 mm long. N = 9 (Scora 1967). Phenology. Flowering material has been collected from April through September. 115 Distribution. Widespread from eastern Mexico to much of the southeastern and eastern United States extending as far north as Illinois. M. citriodora is morphologically most similar to the western species M. austromontana and M. clinopodioides. It is distinguished from the latter by a larger growth habit and more frequent branching pattern as well as the shape of the upper bract tips. Monarda citriodora bract tips taper dramatically to an aristate bristle, whereas M. clinopodioides bracts have a much more gradual tapering. Features that distinguish M. citriodora from M. austromontana are covered in the M. austromontana discussion above. M. citriodora exhibits substantial levels of morphological variability and most authors have assigned varietal names to illustrate that variation. Populations, typically from northern Mexico, that have more attenuate bract tips and darker green foliage have been assigned to M. citriodora var. attenuata, and populations in south eastern Texas with shorter calyx (5 – 7 mm vs. 7 – 15 mm long) and corolla tubes (8 – 11 mm vs. 11 – 18mm long) belong to M. citriodora var. parva. 116 Figure 34. Herbarium specimen of M. citriodora. Massey 2070 (OKL). 117 Representative specimens examined: U.S.A. OKLAHOMA. Atoka: 0.4 miles N. of Chockie, Johnson 254 (OKL); 3 miles E. of Atoka on Hwy. 7 E, Massey 2070 (OKL); Bryan: Durant; US 75, 1 miles north of RT. 78, Semple 566 (OKL); Carter: Criner Hills, 1.5 miles northeast of Brock, Goodman 7849 (OKL); Choctaw: Hwy 109 near junction with Hwy 70, S of railroad tracks and about 1 mile SW of Fort Towson, Dorr 2381 (TEX);Marshall: 5 miles south of Madill at crossing of routes 99 and 32, Ettner 77-1 (OKL); pasture along Buncombe Creek, 1.5 miles NW of Shay, Williams 331 (OKL); Mayes: Pin Oak Acres 5 miles E, 1 and 3/4 mile N of Mazie; Sec. 15, T19N, R19E; Atoka fm, First 7 (OKL); Murray: on SH 77D east of I-35 north of Turner Falls, Folley 564 (OKL); Roger Mills: Antelope Hills, 4.5 miles northeast of Durham, T. 17N., R 25W., Sect. 22, Goodman 8384 (OKL); Rogers: 7.5 miles N Sageeyah Siding; S12 T23N R16E, Vanderpool 759 (OKL); Woods: about 13 miles northwest of Alva; near Greenleaf Creek, Nighswonger 3143 (OKL); TEXAS. Brazos: College Station, Thalis 30 (TEX); Burnet: grasslands in Inks Lake State Park, along Clear Creek, Correll 25295 (LL); Calahan: along Tex. 36, about 14 miles southeast of Abilene, Henderson 63-938 (TEX); Collin: in field along Shelley road, between Renner and Plano, Correll 18997 (LL); Dallas: Stults Prairie, southwest corner of Coit 118 Road and Belt Line Road, Correll 16806 (TEX); Goliad: hwy 249 4 miles east of junction F.R. 81, Semple 578 (MO); Harris: 1.5 miles north of F.M. Road 1959 north of Webster, Correll 32917 (LL); Leon: Blackland prairie on upper Cretaceious marls on top of the Marquez salt dome, Tharp 54996 (TEX); Liberty: off highway 146, ca. 1.6 miles north of Moss Hill, Lundell 15083 (LL);Nueces: Chapman Ranch Road, Segers s.n. (RM); San Patricio: Welder Wild Life Foundation, Scora 2222 (TEX); 0.5 mile west of Ingleside on T. 361, Semple 599 (MO); five miles west of Aransas Pass in black clay along roadside, Webster 7075 (TEX); San Saba: 10 miles southwest of San Saba, Howell 2 (LL); Somervell: 2 miles west of Glen Rose, Correll 19007 (LL); Starr: about 4.1 airmiles W of junction F.M. 650 and S Rt. 83, WNW of Roma; Lower Rio Grande Valley NWR, Fronton Tract, Carr 13595 (TEX); Valverde: depression 3 miles southeast of Del Rio, route 277, Correll 19448 (TEX); Walker: Huntsville, Albers 35013 (TEX). 3. Monarda clinopodioides A. Gray, Syn. Fl. N. Amer. 2(1): 375. 1878.--TYPE: U.S.A. Texas: Dallas Co., near Dallas, June 1874, J. Reverchon 446 (lectotype: GH, isotype: GH). Habit annual, herb to 45 (75) cm tall; upright, branching mainly from base, occasionally from the middle of stem. 119 Stem indumentum variable; internodes puberulent with minute downwardcurled hairs, especially in the lower parts; node with short hairs, nodal plane with long hairs. Penultimate vegetative internode 23 - 72 mm long, 1.2 – 2.6 mm wide. Leaves 38 - 52 mm long, 5.2 - 12 mm wide; petiole 1.8 – 6.5 mm long; serrations 0.4 mm wide; leaf shape lanceolate , base acuminate; margins serrate from apex to 80% length of laminar tissue, apex acute; petiole indumentum of long hairs; abaxial surface puberulent; adaxial surface glabrous; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 5 - 10 mm long, 0.8 - 1.2 mm wide; puberulent. Lower floral bracts 20 - 50 mm long, 4 - 8 mm wide; sessile; lower bract shape lanceolate, base acuminate, margins serrate; apex attenuate to aristate; abaxial surface puberulent; adaxial surface puberulent; margin ciliate to 70% length of bract; abaxial surface green, adaxial surface green. Upper floral bracts 10 - 20 mm long, 4 –5 mm wide; upper bract shape lanceolate, base cuneate, margins ciliate; apex aristate; abaxial surface puberulent; adaxial surface puberulent; margins entire; abaxial surface green; adaxial surface green to purple. Calyx tube 6 – 8.5 mm long, 1.2 – 1.8 mm wide, with minute scattered hairs on outer surface. Calyx lobes 3 – 4.8 mm long, 0.1 – 0.3 mm wide, narrowly triangular with aristate tips, ciliate with multicellular trichomes. Calyx orifice densely hirsute with 120 stiff white hairs. Lower corolla lip 5.2 - 8 mm long, 1.5 - 2.5 mm wide; mid limb to 3.3 mm long, 2.2 mm wide, expanding toward the tip; lateral limbs to 1.3 mm long, 1.2 mm wide. Upper corolla lip 5.2 – 6.2 mm long, 2.2 – 3.5 mm wide, lobes to 1 mm long and 0.8 mm wide, corolla white to pink with violet pigmentation. Stamens to 2.5 cm in length, anthers 0.8 long, 0.3 wide, pollen white. Pistil to 25 mm long. N = 9 (Scora 1967). Phenology. Flowering material has been collected from April through June. Distribution. Southern Kansas south through much of Oklahoma and central to south-central Texas. Monarda clinopodioides is most closely related (see phylogeny), and morphologically similar, to M. pectinata. Both have reduced branching and upper floral bracts that gradually taper to an apical bristle. The two can be distinguished from each other because M. clinopodioides most commonly has purple bracts, calyx lobes and (rarely white) corollas, whereas M. pectinata have green bracts and calyx lobes and yellow corollas. Monarda clinopodioides is also typically taller, growing 30 - 60 cm in height, than M. pectinata, though there is a degree of overlap with the latter growing 20 – 40 cm high. 121 Figure 35. Herbarium specimen of M. clinopodioides. Dubrule 307 (TAMU). 122 Representative specimens examined: U.S.A. OKLAHOMA. Alfalfa: near end of active runway; UTM: (05 779 40 673), Proctor KEG0328 (OKL); Beaver: rt 23, 7 miles south of Beaver, Huff 1391 (OKL); Comanche: 1 km W of Pratt Hill on Deer Creek Rd.; ec 23, T3N, R13W, Thompson S0486 (OKL); 1 km E of SW corner of Quanah Range; Sec 13, T2N, R15W, Thompson S0520 (OKL); 0.5 km E of SW corner of Quanah Range; Sec 14, T2N, R15W, Thompson S0527 (OKL); Custer: 1 mile E, 05 miles S of SE edge of Arapaho, Sanders 132 (OKL); Dewey: 5 miles S of Seiling, Landon 172 (RM); Jefferson: 13 miles east of Ryan along HW 32, 5-10 miles north of the Red River, Crook 953 (OKL); McClain: Johnson's Pasture; 4 miles west of Jct. of I35 on HW 9; south side of road, Lawson 40 (OKL); Tillman: 7 miles north of Chatanooga, Crook 1009 (OKL); Woods: about six miles northwest of Alva; sandy grassland on bluf above Salt Fork River, Nighswonger 672 (OKL); northwest of Alva about 6.5 miles; roadside on bluffs south of river, Nighswonger 1221 (OKL); 2 miles east of junction US 64 and Ok. 14; east of Alva o US 64, then 7 miles N and 0.6 miles E. south of road, Pearce 57 (OKL); TEXAS. Bastrop: both sides of New Road, 0.3 mi N of East Loop Road, ca 3.8 airmiles SE of jct. US Rt. 290 and F.M. 696; Camp Swift Training Site (Texas National Guard); Elgin East Quadrangle, 30.268890, -97.263889, Carter 13788 (TEX); 100 123 meters north of Dan Sawicki Bog (north of Bastrop), Kutac s.n. (TEX); Brazos: about 0.5 miles east of highway 6, along the OSR, Ajilvshi 8400 (TAMU); Burleson: one mile north of Lyons on highway 60, Simpson 139 (TAMU); Dewitt: Albers 45Ph025 (TEX); Fannin: about 1 mile north of Monkstown, Route 79, Correll 27489 (LL); Fayette: along West Point-Muldoon road, Tharp 51-535 (TEX); Freestone: in sandy woods 1 mile southeast of Streetman, Correll 16407 (LL); Hamilton: in caliche soil abt 2 miles N of Hamilton, off of hwy 281, Correll 32847 (LL); Henderson: sandy post oak area about 5 miles southeast of Eustace, Correll 23400 (LL); Hopkins: in oak flatwoods about 2 miles west of Sulphur Springs, Correll 16741 (LL); Lampasas: 12 miles W. Lampasas city limits sign on FM 580, Dubrule 307 (TAMU); Lee: Giddings, Albers 48019 (TEX); Robertson: sandy soil in highway 1940 right of way, 5 miles W of OSR New Baden, Carter 91 (TAMU); NE 1/4 Calver Quad., U.S.G.S 192; 2.5 miles SE of Calvert along Tx. Hwy. 6, Mud Creek marsh, Starbuck 1926 (TAMU); Wichita: prairie hilside within 100 yards of shore of New City (Iowa Park) Lake, 3.5 miles W of Iowa Park, Mahler 1135 (TEX). 4. Monarda mexicana Epling Madroño 3: 26. 1935.--TYPE: MEXICO. Durango: no locality, no date, P.I. Garcia 399 (holotype: 124 US!). Annual herb to 50 cm tall; upright, rarely branching. Stem indumentum variable; internode puberulent with short downward curled hairs and few long hairs, node with short hairs, nodal plane with short hairs. Penultimate vegetative internode 3 – 4 mm long, 0.8 - 1.2 mm wide. Leaves 45 - 60 mm long, 8 - 12 mm wide; petiole 3 – 8 mm long; leaf shape lanceolate, base acuminate; margins subentire , ciliate toward base, apex acute; petiole indumentum of short; abaxial surface puberulent with occasional longer hairs around the midvein; adaxial surface glabrous; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode 5 – 10 mm long, 0.6 - 0.8 mm wide; hirsute. Lower floral bracts 35 - 45 mm long, 16 - 23 mm wide; sessile; lower bract shape ovatelanceolate, base attenuate, margins entire; apex acute; abaxial surface glabrate, with occasional long hairs; adaxial surface glabrous; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface green or green with purple at base of bract. Upper floral bracts 9 – 12 mm long, 3 – 5 mm wide; upper bract shape lanceolate, base cuneate, margins entire, ciliate; apex acuminate; abaxial surface glabrate; adaxial surface glabrate; abaxial surface green; adaxial surface green to violet. Calyx tube 5.8 – 6.2 mm long, 1.8 – 2.1 mm 125 wide, glabrous. Calyx lobes 1.2 – 1.5 mm long, 0.1 – 0.2 mm wide, narrowly triangular, ciliate along the entire margin. Calyx orifice hirsute with flexible white hairs. Lower corolla lip 5.6 – 7.4 mm long, 3.5 – 4.2 mm wide; mid limb to 2.2 mm long, 1 mm wide, toward the tip; lateral limbs up to 1.1 long, 0.9 wide. Upper corolla lip to 7.8 – 9.2 mm long, 3.2 - 4 mm wide, lobes up to 0.2 mm long and 0.16 mm wide, corolla white to violet. Stamens to 18 mm in length, anthers 1.2 mm long, 0.3 mm wide, pollen white. Pistil to 18 mm long. N = 9 (Scora 1967). Phenology. Flowering material has been collected from August. Distribution. Collections of this species are very limited in number, but all have been collected in the state of Durango, Mexico. The highest geographic resolution for the location of this species was obtained from an herbarium label of collections 126 Figure 36. Herbarium specimen of M. mexicana. Maysilles 7771 (TEX). 127 (475 and 476) of James Maysilles: between El Salto and Pueblo Nuevo, Durango. M. mexicana is most likely sister to all the remaining species of Monarda section Aristatae (see Phylogeny section). It is distinguished most notably by its very wide lower bracts which have entire margins, and by its leaves which have subentire margins. It also has glabrous calyx tubes (which, among section Aristatae, is unique to this species) and green upper bracts. Turner (1994) treats M. mexicana as a variety (var. austromontana) of M. citriodora. This was based primarily on the broad calyx lobes exhibited by this species and other individuals of M. citriodora (sensu Turner 1994) he is familiar with in the region. However, while certainly recognizing the large morphological variation inherent in Mexican species of this section, it is the author’s opinion that M. mexicana is a separate species based on additional characters auxiliary to the calyx lobe shape discussed above (see chapter 2). Representative specimens examined: MEXICO. DURANGO. Garcia 399 (photograph)(MICH); from El Salto, south along lumber road toward Pueblo Nuevo (about 60 air miles southwest of C. Durango) Maysilles 7771 (MICH, TEX). 128 5. Monarda pectinata Nutt., J. Acad. Phil., ser. 2, 1: 182. 1847.--TYPE: U.S.A. New Mexico: Santa Fe Co., near Santa Fe, no date, Gambel s.n. (holotype: K, according to McClintock and Epling (1942) and Scora (1967)). Habit annual, herb to 30 (55) cm tall; upright, branching primarily from base of stem. Stem indumentum variable; internodes puberulent with minute downward-curled hairs, especially in the lower parts; node and nodal plane with short hairs. Penultimate vegetative internode 20 – 70 (86) mm long, 1 – 3.5 mm wide. Leaves 3.2 – 6.5 mm long, 5.5 - 12 mm wide; petiole 2.2 – 5 mm long; serrations 0.1 mm wide; leaf shape lanceolate, base acuminate; margins serrate from apex to 80% length of laminar tissue, apex obtuse; petiole indumentum of long hairs; abaxial surface puberulent with small hairs across entire surface; adaxial surface glabrous; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 10 – 50 mm long, 0.6 – 1.2 mm wide; puberulent. Lower floral bracts 21 - 60 mm long, 4 -8 mm wide; sessile; lower bract shape lanceolate, base acuminate margins serrate; apex acuminate; abaxial surface puberulent; adaxial surface glabrous; margin serrate and ciliate ; abaxial surface green, adaxial 129 surface green. Upper floral bracts 8 – 23 mm long, 2.6 – 3.8 mm wide; upper bract shape lanceolate, base cuneate, margins serrate; apex aristate; abaxial surface puberulent; adaxial surface glabrous; margins ciliate; abaxial surface green; adaxial surface green occasionally with purple margins or greenpurple throughout. Calyx tube 6.1 - 10 mm long, 1.5 – 2.5 mm wide, with minute scattered hairs on outer surface. Calyx lobes 2.5 – 3.5 mm long, 0.1 mm wide, aristate, margins ciliate. Calyx orifice densely hirsute with stiff white hairs. Lower corolla lip to 8 (9.5) mm long, 3 mm wide; mid limb to 3.5 mm long, 1.75 mm wide, expanding toward the tip; lateral limbs to 1 mm long, 1 mm wide. Upper corolla lip 5 – 8 (11) mm long, 2 - 3.8 mm wide, lobes to 0.6 mm long and 0.4 mm wide, corolla white to pink. Stamens to 26 mm in length, anthers to 1.2 mm long, 0.8 wide, pollen white. Pistil to 26 mm long. N = 18 (Scora 1967). Phenology. Flowering material has been collected from May through October. Distribution. Western Texas and the panhandle of Oklahoma extending west through Arizona and Colorado and north through Nebraska. M. pectinata is most closely related (and morphologically similar) to M. clinopodioides. The features distinguishing the 130 Figure 37. Herbarium specimen of M. pectinata. Ballinger s.n. (TEX) 131 two species can be found in the discussion under M. clinopodioides. The two species are largely allopatric, but populations of M. pectinata from western Texas occur within 250 miles of the central Texas populations of M. clinopodioides Representative specimens examined: U.S.A. COLORADO. Boulder: Boulder, Daniels 13 (MSC); Larimer: Fort Collins, Lawsen 384 (MSC); un-named mountain 0.5 miles NW of Cherokee Park rd, T10N R71W S10, Neely 3566 (RM); Park: along route 24, 1 mile west of Wilkerson Pass, Lake George, Churchil 7482138 (MSC); NEBRASKA. Banner: pipo woodland near C-7 about 8 miles SE of Harrisburg, Long s.n. (RM); Dawes: in the sand hills near Ft. Robinson, Porter 9356 (RM); Devel: Rush Creek, Tracy 7991 (MSC); Garden: along dirt road in Crescent Lake National Wildlife Refuge; ca. 1 to 5 miles S of lakeside, Theiret 59653 (KNK); Thomas: Nebraska National forest, Halsey, Haynes 9692 (KNK); NEW MEXICO Catron: middle fork of Gila, Wooton s.n. (NMC); near Gallo Spring in western Catron Co., Wooton s.n. (NMC); Cibola: Craters, Wooton s.n. (NMC); Harding: route 39, 10 miles east of town, Higgins 6948 (MSC); TEXAS. Andrews: North Andrews City, Ballinger s.n. (TEX); Bailey: in deep sandy loam of woody pasture 4 miles north of Maple, Rosson 956 (MSC); Dawson: 11.6 miles west of Lamesa, Shinners 30061 (TEX); Garza: barditch highway 380, 16 air miles east of Post, Hutchins 1133 (TEX); Hockley: Rachaner 132 (TEX); 132 Jefferson: grassy roadside in Limpia Canyon, 9-10 miles above Ft. Davis, Correll 18391 (TEX); near base of north slope of Mt. Livermore, Correll 33768 (TEX); Davis Mountains; Madeira Canyon, Kugel 2098 (MSC); Davis Mountains, highway 118, Scenic Drive, 7 miles from cut-off to McDonald Observatory, Lundell 14224 (TEX); Davis Mts., Hw 166, SW slope of Sawtooth Mt., protected arroyo, under oak tres, Sanders 75084 (TEX); Musquiz Canyon; Davis Mountains; 8 miles south of Ft. Davis, Warnock 5672 (TEX); Lubbock: Demaree 7554 (TEX); Midland: rte 349, 5 miles north of Midland, Correll 22048 (TEX); Midland Draw, 10 miles eastsoutheast of Midland, LeSassier 225 (TEX). 6. Monarda n. sp. 1, sp. nov.--TYPE: MEXICO. Chihuahua: Cuauhtemoc-La Junta, 2000 m, 25 July 1994, C. Yen & E. Estrada 2681 (holotype: CAS!; isotype: CAS!). Habit perennial, herb to 50 cm tall; upright, branching from the middle of stem. Stem indumentum variable; internodes canescent with minute downward-curled hairs. Nodes with short hairs, nodal plane with long hairs. Penultimate vegetative internode 25 - 35 mm long, 2.5 - 3.5 mm wide. Leaves 30 - 45 mm long, 5 - 11 mm wide; petiole 0.5 - 4 mm long; serrations 0.3 mm wide; leaf shape lanceolate, base acute; margins serrate from 133 apex to 75% length of laminar tissue, apex acute; petiole canescent with occasional long hairs; abaxial surface canescent with occasional long hairs around the midvein; adaxial surface canescent; margin ciliate toward the base of the leaf. Basal inflorescence internode 25 - 45 mm long, 1.5 - 2.5 mm wide; puberulent. Lower floral bracts 15 - 33 mm long, 8 - 12 mm wide; sessile; lower bract shape linear to lanceolate, base attenuate, margins entire; apex acuminate; abaxial surface canesent; adaxial surface canescent; margin ciliate toward the base of the bract; both surfaces canescent. Upper floral bracts 4 - 8 mm long, 0.8 - 3 mm wide; upper bract shape lanceolate, base cuneate, margins entire; apex acuminate; abaxial surface canescent; adaxial surface canesent; margins ciliate. tube 2.5 - 5.5 mm long, 1 - 2 mm wide, puberulent. Calyx Calyx lobes 1.8 - 2.6 mm long, 0.1 mm wide, aristate, margins ciliate. Calyx orifice densely hirsute with stiff white hairs. Lower corolla lip 4.5 - 5.2 mm long, 2.1 - 2.8 mm wide; mid limb up to 1.6 mm long, 1.4 mm wide, expanding toward the tip; lateral limbs to 1 mm long, 0.8mm wide. Upper corolla lip 4.5 - 5.5 mm long, 2.8 - 3.2 mm wide, lobes to 0.1 mm long and 0.15 mm wide, corolla white. Stamens to 10 mm in length, anthers 1.5 - 2.5 mm long, 0.4-0.7 mm wide, pollen white. Pistil 8 mm long. Phenology. Flowering specimens have been collected in July. 134 Distribution. Sierra Madre mountains in the state of Chihuahua, Mexico. Monarda n.sp.1 is most closely related, and morphologically similar, to M. austromontana. The former can easily be distinguished from the latter by their canescent vesture which gives the stems and leaves a grey appearance. The species is known from only a single collection. Representative specimens examined: Mexico. CHIHUAHUA: Yen 2681 (CAS); between Cuauhtémoc and La Junta. 135 Figure 38. Herbarium specimen of Monarda n. sp. 1. Yen 2681 (CAS). 136 Monarda section Cheilyctis Epling, Univ. Calif. Publ. Botany 20: 186. 1942. Monarda subsection Cheilyctis Epling, Madroño 3: 22. 1935. Annual or perennial herbs. Floral bract tips acute to acuminate. Calyx lobe tips acute to acuminate. Pollen yellow. N = 11 or 12. 7. Monarda arkansana ined., comb nov. M. punctata var. arkansana (McClintock and Epling) Shinners, Field and Lab. 21(2): 90. 1953. M. punctata ssp. arkansana McClintock and Epling, Univ. Calif. Publ. Botany 20: 182. 1942. —TYPE: U.S.A. Arkansas: Hot Springs County, near Magnet Cove, no date, Demaree 15590 (holotype: LA). Habit annual or perennial, subshrub to 65 cm tall; upright, branching heavily from the middle of stem. Stem indumentum variable; internode with long villous hairs, node with long villous hairs, nodal plane with long villous hairs. Penultimate vegetative internode 21 – 76 mm long, 1.2 – 3.2 mm wide. 137 Leaves 46 – 86 (100) mm long, 8 – 18 mm wide; petiole (4) 6.5 – 17 (24) mm long; serrations 0.2 mm wide; leaf shape lanceolate to oblanceolate, base attenuate; margins serrate from apex to 15% length of laminar tissue, apex acute to acuminate; petiole indumentum of long hairs; abaxial surface villous along the midvein; adaxial surface glabrous; margin ciliate toward the base of the leaf; both leaf surfaces green. Inflorescence internode 5 - 18 mm long, 0.6 - 0.8 mm wide; villous or glabrous. Lower floral bracts 30 - 65 mm long, 5.2 – 12 mm wide; sessile; lower bract shape lanceolate, base attenuate, margins entire; apex acuminate; adaxial surface glabrous; adaxial surface villous; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface green. Upper floral bracts 13 – 30 mm long, 5 – 13 mm wide; upper bract shape oblanceolate, base cuneate, margins serrated; apex acuminate; abaxial surface villous; adaxial surface glabrous; margins glabrous; abaxial surface green; adaxial surface red to green. Calyx tube 6.2 – 9.4 mm long, 1.1 - 2 mm wide, with minute scattered hairs on outer surface. Calyx lobes 1.0 - 1.6 mm long, 0.2 - 0.4 mm wide, lanceolate, with minute hairs along the margins. Calyx orifice ciliate with flexible white hairs. Lower corolla lip 4.1 – 7 mm long, 3 – 4.1 mm wide; mid limb to 3.5 mm long, 1.3 mm wide, expanding toward the tip; lateral limbs to 1.3 mm long, 1 mm wide. Upper corolla lip 7.8 – 12 mm 138 long, 3.8 – 4.2 mm wide, lobes to 0.4 mm long and 0.3 mm wide, corolla yellow in color with violet pigmentation. Stamens to 22 mm in length, anthers 0.8 mm long, 0.2 wide, pollen yellow. Pistil to 21 mm long. Phenology. N = 11 (Scora 1967). Flowering material has been collected from July through October. Distribution. Central and southern Arkansas extending south into northeastern Texas and northeastern Louisiana. Representative specimens examined: U.S.A. ARKANSAS, Desha: access road to Dam #2 on Ark. River, Locke 2784 (UARK); Garland: Hot Springs, Demaree 410 (UARK); Hot Springs National Park, Sugar Loaf Mt., Demaree 15840 (UARK); Hot Springs, Foreman s.n. (US); Hot Springs, Letterman s.n. (US); 1 M west of Three Frks Walnut Creek on 270 north of Crystall Springs, Scora 1056 (US); Hempstead: McNab, Palmer 12665 (US); Hot Springs: SW 1/4, Sec. 2, T 5S, R 22W, Point Cedar Quadrangle, Roberts 471 (UARK); Jefferson:within 1/4 mi. of 3006 West Orlando Drive in SW Pine Bluff, Locke 26 (UARK); Miller: Texarkana, Buchhohy 335 (UARK); Polk: vicinity of Grannis; 25 mi. S. of Mena, Moore 400459 (UARK); Bog Springs; Whiskey Peak vicinity, Moore 410-213 (UARK); Big Fork, Moore s.n. (UARK); Saline: Bryant, Moore 400401 (UARK); Sebastion: Fort Smith, Bigelow s.n. (US); Union: east of El Dorado (Champagnole Rd.), Moore 68341 (UARK); along 139 Figure 39. Herbarium specimen of M. arkansana. Moore 400459 (UARK). 140 US route 167, 5 km N of AR route 7 junction S of town. Sec. 10, T 18S, R 15W, Thomas 111098 (MSC); beside U.S. 167 south of El Dorado and 3 miles north of Ark. 7 junction; Sec. 10, T18S, R15W, Thomas 111098 (UARK). 8. Monarda fruticulosa Epling, Madrono 3: 26. 1935. Monarda punctata L. var. fruticulosa (Epling) Scora, Uni. Calif. Publ. Bot. 41: 46. 1967.--TYPE: U.S.A. Texas: Duval Co., near Pena Station, September 1884, Harvard s.n. (holotype: US!). Perennial herb to 1 m tall; upright, branching from the middle or base of stem. Stem indumentum variable; internodes canescent with minute downward-curled hairs; node with short hairs, nodal plane with occasional longer hairs. Penultimate vegetative internode 10 – 60 mm long, 0.6 - 3 mm wide. Leaves 10 - 40 mm long, 1 - 4 mm wide; sessile or with minute (0.5 mm) petiole; serrations 0.2 mm wide; leaf shape linear, base acuminate; margins serrate from apex to 15% length of laminar tissue, apex acute; petiole indumentum of long hairs; abaxial surface canescent; adaxial surface canescent; margin occasionally ciliate toward the base of the leaf. 141 Basal inflorescence internode 10 – 30 mm long, 0.6 - 1.5 mm wide; caulescent. Lower floral bracts 10 – 40 (56) mm long, 2-5 (7) mm wide; sessile; lower bract shape linear, base attenuate, margins serrate; apex acute; abaxial surface canescent; adaxial surface canescent; margin ciliate toward the base; adaxial surface yellow to white. Upper floral bracts 8 - 20 mm long, 3 – 12 mm wide; upper bract shape elliptic, base cuneate, margins entire or minutely serrated; apex acuminate; abaxial surface canescent; adaxial surface canescent; margins entire or occasionally cilate; adaxial surface white to pink. Calyx tube 4 – 6.5 mm long, 1 - 2 mm wide, with long villous trichomes on outer surface. Calyx lobes 1 – 2 mm long, 0.1 – 0.4 mm wide, narrowly triangular, with long villous trichomes along the margins. Calyx orifice hirsute with flexible white hairs. Lower corolla lip 2 – 5.5 mm long, 1.5 - 3 mm wide; mid limb to 2.6 mm long, 1.5 mm wide, expanding toward the tip; lateral limbs to 2 mm long, 1.3 mm wide. Upper corolla lip 5 -7 (12) mm long, 2 – 3 (4.2) mm wide, lobes to 0.6 mm long and 0.5 mm wide, corolla white to pink with lower corolla lip with occasional maroon pigmentation. Stamens to 1.6 mm in length, anthers to 0.8 mm long, 0.4 mm wide, pollen yellow. (Scora 1967) 142 Pistil to 16 mm. N = 11 Phenology. Flowering material has been collected from March through December. Distribution. Southern Texas in sandy roadsides and pastures. M. fruticulosa is part of the narrow leaf clade (see phylogeny). It is most closely related to M. maritima, Monarda n. sp. 2, and M. viridissima, and though they all have narrow leaves, M. fruticulosa is the only species whose leaves are truly linear. It also has acuminate calyx lobes, but its most distinguishing feature is the canescence that covers all of its vegetative surfaces. 143 Figure 40. Herbarium specimen of M. fruticulosa. Runyon 3991 (RM). 144 Representative specimens examined. U.S.A. TEXAS. Brooks: north of Encino, C.L. Lundell & 8850 (TEX); State Highway 285, twenty miles east of Hebronville, Garcia 110 (TEX); 22 mi. W of Falfurias, Muller 8053 (TEX); ca 18 mi S of Falfurrias on Scot & Hoper Ranch, O'Brian 1163 (RM); San Manuel to Encino, Runyon 2628 (TEX); 6.05 M north of Encino on highway 281, south of Falfurias, Scora 2223 (US); North of Encino on Highway 281, south of Falfurias, Scora 2224 (US); Cameron: vicinity of Brownsville, Shiller 178 (US); Hidalgo: 8 miles north of San Manuel, Lundell 12792 (TEX); 8 miles north of San Manuel, Lundell 12796 (US); Jim Hogg: E side of F. M. 1017, 3.0 roadmiles S of jct. with smaller road at Agua Nueva. Agua Nueva SE Quadrangle, 26.513800, -98.364600, Carr 13206 (TEX); ten miles south of Hebbronville on road No. 1017, Johnston 541835 (TEX); 1 mile north of Agua Nueva, Salinas 180 (TEX); Kenedy: 10 miles south of Sarita, Correll 26913 (LL); King Ranch, roadside 5.5 mi. S. rest stop S. of Sarita, Dubrule 804 (TAMU); Sarita, Fisher 41125 (RM, US); Saltillo Pasture, Norias Division of King Ranch, Johnston 53254.21 (TEX); Las Norias, Runyon 3991 (RM, TEX); on highway between Las Norias and Armsrong, Runyon 4333 (US, TEX); Kennedy Ranch, Tharp 42-40 (TEX); Zapata: about 11 145 miles northeast of San Ygnacio, Corell 35452 (GH, LL); San Ignacio, Gamboa 158 (TEX). 9. Monarda humilis (Torrey) Prather & J. A. Keith, Novon 13: 1. 2003. Monarda punctata var. humilis Torrey, Report of an Expedition down to the Zuni and Colorado Rivers 166. 1853. --TYPE: U.S.A. New Mexico: no date, [S.W.] Woodhouse s.n. (holotype: GH). Habit annual, herb to 50 m tall; upright, branching (occasionally from the base) from the middle of stem. Stem indumentum puberulent with small downward curling hairs on the internodes, nodes and nodal plane, often red (especially toward top of plant). Penultimate vegetative internode 20 – 50 mm long, 0.8 - 1.2 mm wide. Leaves 20 - 50 mm long, 3 - 8 mm wide; petiole 1 – 8 mm long; serrations 0.3 mm wide; leaf shape lanceolate, base acute; margins serrate from apex to 25% length of laminar tissue, apex acute; petiole indumentum of short hairs but occasionally with longer ciliate hairs; abaxial surface puberulent occasionally with medium hairs around the midvein; adaxial surface puberulent; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 5 - 7mm 146 long, 0.6-0.8 mm wide, puberulent. Lower floral bracts 20 - 40 mm long, 4 - 10 mm wide, petiolate; lower bract shape lanceolate, base acute, margins serrate; apex acute; abaxial surface puberulent; adaxial surface puberulent; margin ciliate toward the base; abaxial surface green, adaxial surface green. Upper floral bracts 1.9 – 25 mm long, 1 –9 mm wide; upper bract shape lanceolate, base cuneate, margins serrate; apex acuminate; abaxial surface puberulent; adaxial surface puberulent; margins entire; abaxial surface green; adaxial surface purple to green. Calyx tube 4 – 7.5 mm long, 1 – 2.5 mm wide, glabrous or with small trichomes along the veins. Calyx lobes 0.5 - 1 mm long, 0.4 – 0.9 mm wide, with long flexuous hairs on the margins. Calyx orifice hirsute with long flexuous hairs. Lower corolla lip 2 - 4 mm long, 1 – 3.5 mm wide; mid limb to 2 (3) mm long, 1.4 (2) mm wide, expanding toward the tip; lateral limbs to 1.4 mm long, 1.5 mm wide. Upper corolla lip 4 – 5.5 (6.5) mm long, 2 - 3 mm wide, lobes to 0.3 mm long and 0.3 mm wide, corolla white with maroon margin and spots. Stamens to 15 mm in length, anthers to 1.7 mm long, 1.5 mm wide, pollen yellow. Pistil to 15 mm long. N = 12 (Ward and Spellenberg 1984) Phenology. Flowering material has been collected from June through September. Distribution. Central and western New Mexico 147 M. humilis is most closely related, and most morphologically similar to M. occidentalis and M. stanfieldii and is part of the glabrous calyx clade (see Phylogeny section). It is found in sandy regions of western and central New Mexico which is west of the distribution range of M. occidentalis. It is distinguished from these two species by its smaller stature and purple pigmentation that is found on the upper bracts, calyx, and corolla, but also occasionally along the stems. Monarda humilis has violet flowers with maroon spots instead of the yellow or white flowers (which may also have maroon spots) commonly associated with the rest of the clade. 148 Figure 41. Herbarium specimen of M. humilis. Goodrow 522 (UNM). 149 Representative specimens examined: U.S.A. NEW MEXICO. Catron: North Cebollita Mesa T8N R9W, Sec. 8, McIntosh 1006 (UNM); Cibola: T7N, R10W, Sec. 8, DeBruin 448 (UNM); Inscription Rock, Wooton s.n. (US); Mts. West of Grant's Station, Wooton s.n. (NMC); Inscription Rock, Wooton s.n. (US); McKinley: T10N R18W Sec. 18 Zumi Indian Reservaion. 1.5 mi. e of Blackrock, 0.5 mi. NE of Blackrock Reservoir, Brandt 1094 (UNM); Rio Arriba: West of San Larenzo, Wooton s.n. (NMC, US); Sandoval: Santo Domingo Pueblo, Abbott s.n. (UNM); Socorro: S20, T3S, R4W, Goodrow 522 (UNM); plains east of Datil on hwy 60, Higgins 7737 (NMC); 10 mi. S of Claunch on St. 55., Pase 3566 (NMC); near Cibola. Between Datil and Magdalena on roadside highway 60, Tafoya 131 (UNM); Torrance: Cibola N.F. Gallinas Mtns. Fm Rd. 458, Pase 3621 (NMC); Valencia: El Morro Nat'l Monument, Carlson SC4 (UNM). 10. Monarda maritima (Cory) B.L. Turner, Phytologia. 77(1): 72. 1994. Monarda punctata L. var. maritima Cory, Field & Lab. 17: 52. 1949.--TYPE: U.S.A. Texas: San Patricio Co., 4mi W of Aransas Pass, 14 October 1951, F.B. Jones 648 (holotype: SMU). 150 Habit perennial, herb to 110 cm tall; upright, branching heavily from the base of stem. Stem indumentum densely pubescent; internodes hirsute with 1 - 3 mm long spreading hairs; node with long hairs, nodal plane with long hairs. Penultimate vegetative internode (14) 20 -30 (35) mm long, 0.9 3 mm wide. Leaves 15 - 60 mm long, 4 - 10 mm wide; petiole 0.5 – 2 mm long; serrations to 1 mm wide; leaf shape linear to lanceolate, base acuminate; margins serrate from apex to 75% length of laminar tissue, apex acute; petiole indumentum of long hairs; abaxial surface hirsute with long hairs around the midvein; adaxial surface glabrous or with occasional spreading hairs; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode 5 – 10 mm long, 0.6 - 0.8 mm wide; hirsute. Lower floral bracts 15 - 60 mm long, 3 - 11 mm wide; sessile to minutely petioled; lower bract shape linear to lanceolate, base attenuate, margins distinctly toothed; apex acute; abaxial surface hirsute; adaxial surface glabrous or with occasional spreading hairs; margin ciliate toward the base of the bract; both surfaces green to red to white. Upper floral bracts 12 – 22 mm long, 6 – 9.5 mm wide; upper bract shape lanceolate, base cuneate, margins serrated; apex acute; abaxial surface hirsute; adaxial surface glabrous or with occasional hairs; margins ciliate; abaxial surface green; adaxial surface yellow to white to red. 151 Calyx tube (6) 8 - 10 mm long, 1 - 1.8 mm wide, with minute scattered hairs on outer surface. Calyx lobes 1 – 2.5 mm long, 0.25 - 0.5 mm wide, narrowly triangular, with no hairs along the margins. orifice glabrous or with few hairs. Calyx Lower corolla lip 8 - 10 mm long, 2 – 4 mm wide; mid limb to 2.5 mm long, 1 mm wide, expanding toward the tip; lateral limbs to 1.2 mm long, 1 mm wide. Upper corolla lip 8 - 10 mm long, 3 – 5 mm wide, lobes to 1 mm long and 0.4 mm wide, corolla white with some pink along the edges. Stamens to 15 mm in length, anthers 1.5 – 2.5 mm long, 0.4 – 0.8 mm wide, pollen white. Pistil to 9 mm long. N = 11 (Scora 1967). Phenology. Flowering material has been collected from July through December. Disribution. South eastern coastal Texas. Monarda maritima is a south-eastern Texas species in the narrow-leafed clade (see Phylogeny section) and is most closely related to M. fruticulosa, Monarda n. sp. 2, and M. viridissima. It can easily be distinguished from these by a covering of pilose hairs on almost all the vegetative surfaces. Geographically, M. fruticulosa is the closest neighbor in this clade (150 miles separate them at their closest point), but it has linear leaves and is covered by smaller silvery hairs that gives it a distinct canescence. 152 Figure 42. Herbarium specimen of M. maritima. Johnston 53254.22 (TEX). 153 Representative specimens examined: U.S.A. TEXAS. Aransas: 14 km from Fulton, Churchill 90-930 (MSC); Goose Island State Park, Johnston 53254.27 (TEX); Lamar, situated on the edge of Copano Bay, Palmer 1081 (US); Lamar, Palmer 1081 (US); Copano Bay, Tharp 1553 (US); Kleberg: near Mortila Camphouse, Laureles Division of King Ranch, Johnston 53254.22 (TEX); Laureles Division of King Ranch, near Mortilla Camphouse, Johnston 53254.23 (TEX); eastern Laureles Division of King Ranch, Johnston 53254.24 (TEX); eastern Laureles Division of King Ranch, Johnston 53254.24 (RM); Refugio: Goose Island State park, Scora 2170 (US); Goose Island State park, Scora 2170 (GH, TEX); Austwel, Tharp 5553 (US). 11. Monarda occidentalis ined., comb nov. M. punctata subsp. occidentalis Epling, Madrono 3: 23. 1935. M. punctata var. occidentalis (Epling) Palmer and Steyermark, Ann. Missouri Botan. Gardens 22: 634. 1935.--TYPE: U.S.A. Oklahoma: Woods Co., near Alva, Stevens 3072 (holotype: NY (digital image)!; isotype: GH). Habit annual, herb to 50 cm tall; upright, branching heavily from the middle to upper part of the stem. 154 Stem indumentum of short hairs; internodes puberulent with minute downward-curled hairs; node with short hairs, nodal plane with short hairs. Penultimate vegetative internode 12 – 60 mm long, 1 - 3 mm wide. Leaves 30 - 60 mm long, 6 - 20 mm wide; petiole 5 – 15 mm long; serrations to 0.8 mm wide; leaf shape lanceolate, base acute; margins serrate from apex to 20% - 90% length of laminar tissue, apex acute; petiole indumentum of long hairs; abaxial surface puberulent with short hairs and occasionally longer hairs around the midvein; adaxial surface puberulent; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode 8 – 25 mm long, 0.6 - 0.8 mm wide; puberulent. Lower floral bracts 25 - 70 mm long, 8 – 25 mm wide; sessile, or nearly so; lower bract shape lanceolate, base acute, margins serrate; apex acuminate; abaxial surface puberulent; adaxial surface puberulent; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface green to white (or white with green tips or margins). Upper floral bracts 17.5 - 30 mm long, 6 - 11 mm wide; upper bract shape lanceolate, base cuneate, margins entire; apex acuminate; abaxial surface puberulent; adaxial surface puberulent; margins occasionally ciliate; abaxial surface green; adaxial surface yellow to white. Calyx tube 5 – 7.5 mm long, 1 – 2.5 mm wide, with minute scattered hairs on outer surface. Calyx lobes 0.5 – 1.2 mm long, 0.4 – 1.0 mm wide, broadly deltoid, ciliate. 155 Calyx orifice hirsute. Lower corolla lip 4 – 6.5 mm long, 0.8 - 3 mm wide; mid limb to 2 mm long, 1.5 mm wide, expanding toward the tip; lateral limbs to 0.8 mm long, 0.6 mm wide. Upper corolla lip 3 – 8 mm long, 2 - 4 mm wide, lobes to 0.15 mm long and 0.22 mm wide, corolla white to pink, occasionally with maroon spots. Stamens to 15 mm in length, anthers 0.4 – 0.9 mm long, 0.2 -0.4 mm wide, pollen yellow. Pistil to 17 mm long. N = 11 (Scora 1967). Phenology. Flowering material has been collected from July through December. Distribution. Western Texas, Oklahoma, and southwestern Kansas. M. occidentalis is part of the glabrous calyx clade (see phylogeny). Its closest and most morphologically similar relatives are M. stanfieldii and M. humilis. It can be distinguished from these by its large globose glomerules, short calyx lobe cilia, and white to cream colored corollas. Furthermore, it is separated from M. stanfieldii by its broadly deltoid calyx lobes and from M. humilis by its green (vs. purple) calyces. 156 Figure 43. Herbarium specimen of M. occidentalis. Correll 33017 (TEX). 157 Representative specimens. U.S.A. NEW MEXICO. Valencia: ElMorro National Monument, Vogt 7 (ARIZ); OKLAHOMA. Beaver: 2 mi. S. of Gate along roadside, Laverty 7343 (OKL); Beckham: north fork of Red River at State Highway 30, 6 miles N. of Erick, Goodman 8090 (OKL); Harmon: Hwy 30 N of Salt Fork of Red River, Wagner 3615 (MO); Harper: About 5.5 miles west and 7 miles south of Buffalo, Bellah 76 (OKL); NNE of Buffalo about 12 miles: one mile from Kansas line, Nighswonger 1925 (OKL); Jackson: N boundary, 200 E of active runway.UTM: (0475238382). T3N R20W Sec. 35, McCarthy ALT0280 (OKL); Kingfisher: Kingfisher, Carletan 183 (MSC); Marshall: in vicinity of University of Oklahoma Biological Station, Barclay 851 (US); Roger Mills: Antelope Hils, 4.5 miles northeast of Durham, T. 17N, R 25W, Sect 22., Goodman 8379 (OKL); Woods: north of the Cimarron River ca. 15 miles south of Alva on SH 14 and two miles west on a ranch road, Folley Nr. 580 (OKL); about 5 miles N and 61/2 miles W of Freedom. Roadside of U.S. Hwy. # 64, Nighswonger 3270 (OKL); Woodward: 5 miles E of Mooreland, 36.439136, -99.114304, Springer 374 (OKL); Wolf Creek, 1 mile south-east of Ft. Supply, Waterfall 14926 (US); TEXAS. Andrews: central part of county, Scudday (LL); Childress: on sand hills on south side of Prairie Dog Town fork of the Red River, 10 miles north of Childress, Correll 16859 (TEX); Crane: along Ranch Rd. 1601 about 14 miles south of Penwell. (Ector Co.), Correll 33017 (LL); five to eight miles 158 north of Imperial, Warnock 15444 (LL); Hemphill: Canadian, Palmer 14116 (US); Hutchinson: Maverick Village, Fritch, Drake 108 (TEX); Lynn: O'Donnell, Reed 3437 (US); Stonewall: along U.S. #380, about 1/2 mile west of Swenson, 33.206935, 100.346355, Henderson 63-834 (TEX); Ward: 4 miles northeast of Monahans, Cory 51996 (US); highway 4 miles east of Monahans, Warnock 7878 (LL); Winkler: 3 mi. N.E. of Wink on 115. 100 yds from side of road, Stuessy 157 (TEX). 12. Monarda punctata L., Sp. Pl. 1: 22. 1753.--TYPE: U.S.A. no locality, no date, no collector name (holotype: L, according to Epling (1935)). Monarda punctata L. var. lasiodonta A. Gray, Syn. Fl. N. Amer. 2(1): 375. 1878. Monarda lasiodonta (A. Gray) Small, Fl. S.E. U.S. 1038, 1337. 1903.—TYPE: U.S.A. Texas: no locality, 1834-1935, Drummond s.n. (lectotype: GH (selected by Scora (1967)). Monarda punctata L. subsp. immaculata Pennell, Bull. Torrey Bot. Club 46: 187. 1919. Monarda punctata L. var. 159 immaculata (Pennell) Scora, Univ. Calif. Publ. Bot. 41: 47. 1967.—TYPE: U.S.A. Texas: Victoria Co., Aloe, 8 September 1913, Pennell 5494 (holotype: PH; isotype: NY ( digital image)!) Monarda punctata L. var. intermedia (McClintock & Epling) Waterfall, Rhodora 52: 38. 1950. Monarda punctata L. subsp. intermedia McClintock & Epling, Univ. Calif. Publ. Bot. 20: 184. 1942—TYPE: U.S.A. Texas: Van Zandt Co., near Wills Point, Little 14456 (holotype: UCLA). Monarda punctata L. var. correllii B.L. Turner, Phytologia 77(1): 56. 1994.—TYPE: U.S.A. Texas: Webb Co., red sandy soils along highway 83, 13 mi NW of Webb, 16 July 1957, Correll & Johnston 18109 (holotype: LL!). Habit annual and perennial, herb to 85 cm tall; upright, branching from the base or the middle of the stem. Stem indumentum variable; internodes puberulent with minute downwardcurled hairs and occasionally with longer spreading trichomes; node with short hairs, nodal plane with long hairs. Penultimate 160 vegetative internode 20 – 70 mm long, 0.8 – 3.2 mm wide. Leaves 30 - 100 mm long, 8 – 20 mm wide; petiole 3 – 24 mm long; serrations to 0.4 mm wide; leaf shape lanceolate to ovate, base acute to acuminate; margins serrate from apex to 20% length of laminar tissue, apex acute; petiole ciliate; abaxial surface puberulent or with medium-long hairs around the midvein, occasionally with scattered longer hairs; adaxial surface glabrous; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 5 - 12 mm long, 0.6 - 0.8 mm wide; glabrous or puberulent. Lower floral bracts (25) 30 - 80 mm long, 2 - 13 mm wide; sessile; lower bract shape lanceolate, base attenuate, margins serrate; apex acute; abaxial surface puberulent; adaxial surface glabrous; margin ciliate toward the base; abaxial surface green, adaxial surface green. Upper floral bracts 10 - 36 mm long, 2.8 – 13 mm wide; upper bract shape elliptic, base cuneate, margins entire; apex acuminate; abaxial surface glabrous; adaxial surface glabrous or puberulent; margins glabrous or ciliate; abaxial surface green; adaxial surface purple. Calyx tube 5 – 9.4 mm long, 1 - 3 mm wide, with minute scattered hairs on outer surface. Calyx lobes 0.6 – 1.9 mm long, 0.2 – 0.5 mm wide, narrowly triangular, margins glabrous or ciliate. Calyx orifice glabrous or hirsute. Lower corolla lip (2.5) 6 - 10 mm long, 1.2 – 4.5 mm wide; mid limb to 3.6 mm long, 1.7 mm wide, expanding toward the tip; 161 lateral limbs to 2 mm long, 1.3 mm wide. Upper corolla lip (3.5) 6 - 11 mm long, 2 - 5 mm wide, lobes to 0.4 mm long and 0.5 wide, corolla yellow to cream with deep maroon pigmentation. Stamens to 12 mm in length, anthers to 1.8 mm long, 0.8 wide, pollen yellow. Pistil to 12 mm long. N = 11 (Scora 1967). 162 Figure 44. Herbarium specimen of M. punctata. Marsh 121 (TEX). 163 Phenology. Flowering material has been collected from March through November. Distribution. M. punctata has a wide distribution, extending from northern Mexico southern and eastern United States as far north as Canada. Monarda punctata is probably sister to species in the villous leafed clade(see phylogeny discussion), which, collectively is sister to the remaining species in Monarda section Cheilyctis. It is distinguished from the that clade by the lack of villous indumentum on the adaxial and abaxial (for M. villicaulis) surfaces of the leaves, except for the midveins and petioles, for which it may have longer hairs. Monarda punctata exhibits high levels of intraspecific variation. The most notable differences are in vegetative pubescence and leaf size, and, though much of the variation is continuous, most treatments for this species have included intraspecific rankings. Populations with villous hair on the midveins of the adaxial leaf surfaces are treated as M. punctata var. lasiodonta, and populations where the indumentum is strictly short (0.1 -0.2 mm) appressed hairs (not villous at all) are M. punctata var. intermedia. The previous two varieties are typically found in eastern and central Texas (former) and central and south central Texas (latter). 164 Monarda punctata var. correllii is comprised of populations in south western Texas that exhibit weakly nervate leaves and more widely deltoid calyx lobes, approaching those of M. occidentalis. Populations with narrower (<1.2 cm wide) leaves from south eastern Texas are commonly referred to as M. punctata var. immaculata. Representative specimens examined: Mexico. TAMAULIPAS. Rodriguez 84 (TEX); 10 miles southeast of Nuevo Laredo, U.S.A. ARKANSAS. Garland: Locke 2784 (UARK); access road to Dam #2 on Ark. River, Demaree 410 (UARK); Hot Springs, Demaree 15840 (UARK); Hot Springs National Park, Sugar Loaf Mt., Foreman s.n. (US); Hot Springs, Letterman s.n. (US); Hot Springs, Scora 1056 (US); 1 M west of Three Frks Walnut Creek on 270 north of Crystall Springs, Hempstead: Palmer 12665 (US); McNab, Hot Springs: Roberts 471 (UARK); SW1/4, Sec. 2, T 5S, R 22W, Point Cedar Quadrangle, Jefferson: Locke 26 (UARK); within 1/4 mi. of 3006 West Orlando Drive in SW Pine Bluff, Miller: Buchhohy 335 (UARK); Texarkana, Polk: Moore 400459 (UARK); vicinity of Grannis; 25 mi. S. of Mena, Moore 410-213 (UARK); Bog Springs; Whiskey Peak vicinity, Moore s.n. (UARK); Big Fork, Saline: Moore 400401 (UARK); Bryant, Sebastion: Bigelow s.n. (US); Fort Smith, Union: Moore 68341 (UARK); east of El Dorado (Champagnole Rd.), Thomas 111098 (UARK); beside U.S. 167 south of El Dorado 165 and 3 miles north of Ark. 7 junction; Sec. 10, T18S, R15W, Thomas 111098 (MSC); along US route 167, 5 km N of AR route 7 junction S of town. Sec. 10, T 18S, R 15W, FLORIDA. Bay: Perkins 479 (FLAS); near shores of St. Andrew Sound on e. end of Rafield Peninsula, just s. of 90 degre turn in road to Field Tech Unit, ca. 12 1/2 mile east of the East Bay Bridge, east end of Tyndall AF Base, S17, T6S, R12W, Escambia: Morgan 7296 (RM); highway 125, 5 miles west of Pensacola, Indian River: D'Arcy 2779 (FLAS); roadside hammock, Cardinal Drive, near Humiston Beach, Vero Beach, St. Johns: Herring 75 (FLAS); T6S, R30E, S32; 2.8 miles north of Villano Beach on west side of SR A1A. Along roadside on secondary dunes, TEXAS, Aransas: Correll 18925 (TEX); in open live-oak woods just south of bridge across Copano Bay, Atascosa: Fryxell 1276 (LL); 2 miles S of Poteet, Bexar: Parks Rx 3132 (TEX); east Bexar county, Bowie: Heller 4173 (US); near Texarkana, Brazoria: Fleetwood 9154 (TEX); Brazoria National Wildlife Refuge, Angleton, Cameron: Rudd 3940 (TEX); east of Rio Hondo, Chambers: Tharp s.n. (TEX); Patton, Dimmit: Correll 16025 (LL); 13.5 miles west of Carrizo Springs, Frio: Higdon 53-57 (TEX); northern edge of county, Tharp s.n. (TEX), Goliad: Albers 46040 (TEX); Ander, Harris: Boon 175 (TEX); 6 miles N of Humble, Boon 388 (TEX); three miles north of Humble on north side of San Jacinto River, Traverse 169 (MSC); highway U.S. 59 abou 0.3 miles south of bridge over San Jacinto River, 166 23 miles north of Houston, east side of highway, La Salle: Ramirez 82 (TEX); U.S. hwy. 81, 3 miles south of Cotulla, McMullen: Albers 49035 (TEX); San Miguel River, Webb: Bruni 24 (LL); Farm Road 1472, 15 miles northwest of Laredo, Garcia 25 (TEX); Laredo Junior College, Laredo, York: Kirkman 571 (FLAS); at edge of Patrol Road in old field of Cheatham Annex Naval Supply Center; P.O. Wiliamsburg, WASHINGTONG D.C.: Steele s.n. (MSC). 13. Monarda stanfieldii Small, Fl. S.E. U.S. 1038. 1903. Monarda punctata subsp. stanfieldii (Small) Epling, Madroño 3: 25. 1935. Monarda punctata var. stanfieldii (Small) Cory, Rhodora 38: 407. 1936.--TYPE: U.S.A. Texas: Hayes County, near San Marcos and vicinity, no date, S.W. Stanfield s.n. (holotype: NY (digital image)!). Habit annual, subshrub to 75 cm tall; upright, branching heavily from the middle of stem. Stem indumentum variable; internode with short and long hairs densely packed, node with short hairs, nodal plane with long hairs. Penultimate vegetative internode 18 – 24 mm long, 0.8 - 1.2 mm wide. Leaves 45 - 60 mm long, 15 - 25 mm wide; petiole 2 - 4mm long; serrations 2 mm 167 wide; leaf shape lanceolate to oblanceolate, base attenuate; margins serrate from apex to 25% length of laminar tissue, apex acute to acuminate; petiole indumentum of long hairs; abaxial surface hirsute with medium-long hairs around the midvein; adaxial surface glabrous; margin ciliate toward the base of the leaf; both leaf surfaces green. Basal inflorescence internode 5 - 7 mm long, 0.6 - 0.8 mm wide; glabrous. Lower floral bracts 40 - 75 mm long, 8 - 12 mm wide; sessile; lower bract shape linear to lanceolate, base attenuate, margins entire; apex acuminate; abaxial surface glabrous; adaxial surface glabrous; margin ciliate toward the base of the bract; abaxial surface green, adaxial surface yellow to white. Upper floral bracts 17.5 - 25 mm long, 6 - 11 mm wide; upper bract shape oblanceolate, base cuneate, margins serrated; apex acuminate; abaxial surface glabrous; adaxial surface glabrous; margins glabrous; abaxial surface green; adaxial surface yellow to white. Calyx tube 6.2 - 8.4 mm long, 1.2 - 1.9 mm wide, with minute scattered hairs on outer surface. Calyx lobes 1.2 - 1.6 mm long, 0.3 - 0.5 mm wide, narrowly triangular, with no hairs along the margins. white hairs. Calyx orifice densely hirsute with stiff Lower corolla lip 3.0 - 3.4 mm long, 1.2 - 1.9 mm wide; mid limb up to 2.4 mm long, 0.9 mm wide, not expanding toward the tip; lateral limbs up to 1.4 mm long, 0.8 mm wide. Upper corolla lip 7.0 - 8.0 mm long, 2.8 - 3.6 mm wide, lobes to 168 0.1 mm long and 0.15 mm wide, corolla yellow with deep maroon pigmentation. Stamens to 15 mm in length, anthers to 1.7 mm long, 1.5 mm wide, pollen yellow. Pistil to 15 mm long. N = 11 (Scora 1967). Phenology. Flowering material has been collected from May through July. Disribution. Central Texas in granitic sands. Monarda stanfieldii is limited in distribution to granitic sandy soils in three counties (Burnet, Llano, and Travis) in central Texas. It is probably most closely related, and morphologically most similar, to M. humilis and M. occidentals but can be distinguished easily from them M. stanfieldii’s yellow flowers and the presence of very densely packed white hairs around the opening (orifice) of the calyx tubes. Though other species of Monarda section Cheilyctis do have orifice hairs, the hairs of M. stanfieldii are so distinctive that the feature is obvious even as seen from a short distance. Monarda stanfieldii is found in sympatry with M. punctata, but there are no signs of introgression (at least the author has not seen any specimens that appear intermediate to these two species). 169 Figure 45. Herbarium specimen of Monarda stanfieldii. 9029 (TEX). 170 Lundell Representative specimens examined. U.S.A. TEXAS. Burnet: Inks Dam area: 0.25 mi S of Hwy 29 on 4142. NE side of road., 30.754408, -98.375621, Sammons s.n. (MSC); Llano: Near Inks Dam, 30.729772, -98.381892, Lundell 9029 (MICH); Near Inks Dam, 30.729772, -98.381892, Albers s.n. (TEX); Sandy banks of Llano river, Albers, 49070 (CAS); Granite mountain near Llano, Wolff 3859 (MSC); Granite outcrop on north-west corner of the intersection of U.S. Highway 261 and RM 1431 at Buchanan Dam., 30.761020, -98.460810, Walters 593 (TAMU); Travis: Austin Dam, 30.292920, -97.787504, Tharp s.n. (MICH); Lake Austin Dam, 30.292800, -97.787504, Tharp s.n. (TEX); Austin, Tharp s.n. (MICH); Austin, Tharp s.n. (MICH); Austin, Tharp s.n. (TEX); Austin Dam, 30.292890, -97.787504, Albers s.n. (TEX); Off highway 71, on sand hill bove Perdnales River, 30.389931, 98.085587, Lundell 15109 (MICH); off highway 71, on sand hill bove Perdnales River, 30.389931, -98.085587, Lundell 15109 (US); In Zilker Park near Austin, 30.267804, -97.768057, Barkley 46262 (TEX). 14. Monarda villicaulis ined., comb nov. M. punctata var. villicaulis (Pennell) Shinners, Field and Lab. 21(2): 90. 171 1953. M. punctata ssp. villicaulis Pennell, Bull. Torrey Botan. Club 46: 186. 1919.--TYPE: U.S.A. Indiana: near Clarks, no date, Pennell 6412 (holotype: NY (digital image)!). Habit annual or perennial, subshrub to 60 cm tall; upright, branching heavily from the middle of stem. Stem indumentum variable; internode with long villous hairs densely packed, node with long villous hairs, nodal plane with long villous hairs. Penultimate vegetative internode 20 – 45 mm long, 1.2 – 2.2 mm wide. Leaves 45 – 75 (90) mm long, 8 – 18 mm wide; petiole (4.5) 6.5 – 10 (12) mm long; serrations 0.2 mm wide; leaf shape lanceolate to oblanceolate, base attenuate; margins serrate from apex to 15% length of laminar tissue, apex acute to acuminate; petiole indumentum of long hairs; abaxial surface villous throughout; adaxial surface villous; margin ciliate toward the base of the leaf; both leaf surfaces green. Inflorescence internode 5-18 mm long, 0.6-0.8 mm wide; villous. Lower floral bracts 32 - 56 mm long, 4.5 – 8.5 mm wide; sessile; lower bract shape lanceolate, base attenuate, margins entire; apex acuminate; abaxial surface villous; adaxial surface villous; margin ciliate toward the base of the bract; abaxial surface 172 green, adaxial surface green to white. Upper floral bracts 14 – 25 mm long, 5 – 9.5 mm wide; upper bract shape oblanceolate, base cuneate, margins serrated; apex acuminate; abaxial surface villous; adaxial surface villous; margins glabrous; abaxial surface green; adaxial surface green to white. Calyx tube 5.1 – 6.9 mm long, 1.1 - 2 mm wide, with minute scattered hairs on outer surface. Calyx lobes 0.9 - 1.37 mm long, 0.2 - 0.3 mm wide, lanceolate, with minute hairs along the margins. orifice ciliate with flexible white hairs. Calyx Lower corolla lip 4.8 – 7.9 mm long, 2.2 - 3 mm wide; mid limb to 3.3 mm long, 1.3 mm wide, expanding toward the tip; lateral limbs to 1.1 mm long, 0.85 mm wide. Upper corolla lip 6.2 – 8.7 mm long, 2.2 – 4 mm wide, lobes to 0.3 mm long and 0.3 mm wide, corolla yellow in color with violet pigmentation. Stamens to 22 mm in length, anthers 0.8 mm long, 0.2 wide, pollen yellow. long. Pistil to 21 mm N = 12 (Scora 1967). Phenology. Flowering material has been collected from July through September. Distribution. Northeastern U.S. extending west as far as Indiana and as far south as Missouri. Representative specimens examined: U.S.A. INDIANA. La Porte: Dingler's Pond, just outside (E) of Michigan City, 41.692120, 86.965038, Miller 1828 (MO); Porter: Indiana Dunes State Park, 173 Figure 46. Herbarium specimen of M. villicaulis. Hestbeck 10 (MSC). 174 Miller 542 (MSC); MARYLAND. Worcester: Asateague Island; at roadside in sand at bend in road just NW of Northeach parking lots; North Beach; National Seashore, Hill 15468 (US); MICHIGAN Allegan: Between Mt. Baldhead and Lake Michigan near Saugutuck, Gilly 7172 (MSC); 4 Miles W of Saugatuck, Oval Beach. Secondary Dune., Hestbeck 10 (MSC); Barry: Rutland Twp, sandy slope near Otis Lake, Churchill s.n. (MSC); Bull's Prairie, Drew M264 (MSC); Yankee Springs State Game Area; Sec. 28, R 10 W, T 3 N., Gun Lake Road, opposite ranger station, Eickwort 68 (MSC); T. 3N., R. 9W., Sec. 31., Nielsen s.n. (MSC); Berrien: Chikaming Twp. W1/2 NW Sec. 21-T7S-R20W., Atwood 406 (MSC); Kalkaska: T25N R8W Sec. 19, Ryel L120 (MSC); Kent: Cascade Twp., T6N, R10W, Sec. 22; 2 miles south east of Cascade, Gilly 1078 (MSC); Rogue River State Game Area, Oak Grove, Hoffman 310 (MSC); Plainsfield Twp. T8N, R11W, sec. 26, Parmelee 819 (MSC); Hogadane Creek, Grand Rapids, Shadick s.n. (MSC); Hogadone Creek, Grand rapids, Skeels s.n. (MSC); Mecosta: Austin Twp: T14N, R9W, Sec. 13., Parmelee 1015 (MSC); section 23, Dalton Twsp, Bourdo 306 (MSC); Muskegon: section 23, DaltonTwp, Muskegon (Bourdo Farm), Bourdo 20042 (MSC); Whitehall, CP 302 (MSC); near North Muskegon. T. 10N. R. 17W., Sec. 21.; dunes near Coast Guard Station at end of 175 highway 213, Gillis 5415 (MSC); Newago: Big Prairie Desert, Drew M270 (MSC); Oceana: Upper Silver Lake among jack pines, Bourdo 30340 (MSC); T. 13N., R. 18W., Sec. 6. dunes between Stony Lake and Lake Michigan. Top of Vesper Dune, Gilis 5889 (MSC); Ottawa: T6NR15W, Sec 16, SE-1/4. Lat - 42 deg 54' N on - 86 deg 12' W; campbell site, Field Unit CMP - 12, Goff 1002 (MSC); St. Clair: near Algonac, Dodge 509 (MSC); Van Buren: Drew M171 (MSC); Keeler, Pepoon s.n. (MSC); Washtenaw: on Hogback road, Scora 498 (US). 15. Monarda viridissima Correll, Wrightia 9: 76. 1968.--TYPE: U.S.A. Texas: Bastrop Co., 2 mi E of Bastrop along route 21, 10 September 1968, D.S. Correll 36368 (holotype: TEX!; isotypes: GH, TEX!, UC, US!). Habit perennial herb to 60 cm tall; upright, branching from middle and base of stem. Stem indumentum variable; internodes puberulent with minute downward-curled hairs, occasionally with spreading hairs in the upper parts; node with short hairs, nodal plane occasionally with longer hairs. Penultimate vegetative internode 20 – 50 mm long, 0.9 - 2 mm wide. Leaves 25 - 52 mm long, 4 - 7 mm wide; petiole 1 – 5 mm long; serrations 0.3 mm wide; leaf shape linear-lanceolate, base acuminate; margins 176 serrate from apex to 20% length of laminar tissue, apex acute to acuminate; petiole indumentum of long hairs; abaxial surface puberulent with occasional longer hairs around the midvein; adaxial surface puberulent; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 20 – 40 mm long, 0.6 - 0.8 mm wide; puberulent. Lower floral bracts 20 - 45 mm long, 3 - 7 mm wide; sessile; lower bract shape linearlanceolate, base attenuate, margins serrate, apex acuminate; abaxial surface puberulent; adaxial surface puberulent; margin ciliate toward the base; abaxial surface green, adaxial surface green to purple. Upper floral bracts 13 - 25 mm long, 1.2 – 9 mm wide; upper bract shape lanceolate, base cuneate, margins serrate; apex acuminate to aristate; abaxial surface puberulent; adaxial surface puberulent; margins entire; abaxial surface green; adaxial surface pink to purple. Calyx tube 6.4 – 7.5 mm long, 1.4 – 1.8 mm wide, with minute scattered hairs on outer surface. Calyx lobes 1.1 - 1.6 mm long, 0.3 - 0.5 mm wide, narrowly triangular, rarely ciliate. flexible white hairs. Calyx orifice with short Lower corolla lip 2.6 - 6 mm long, 1.6 - 3 mm wide; mid limb to 3 mm long, 1.8 mm wide, expanding toward the tip; lateral limbs to 1.8 mm long, 1 mm wide. Upper corolla lip 6.5 – 9.8 mm long, 2.7 – 4.2 mm wide, lobes to 0.3 mm long and 0.3 mm wide, corolla white in color with violet pigmentation. Stamens to 24 mm in length, anthers 0.8 mm long, 177 0.2 wide, pollen yellow. Pistil to 23 mm long. N = 11 (Scora 1967). Phenology. Flower material has been collected from June through November. Distribution. Central south Texas in sandy oak woods and roadsides. M. viridissima is in the narrow leaf clade (see phylogeny) and is most closely related (and most morphological similar) to M. fruticulosa, M. maritima, and Monarda n. sp. 2. It has longer and more linear-shaped leaves than Monarda n. sp. 2, and is distinguished from M. fruticulosa and M. maritima by the presence of short spreading hairs at right angles to the stem instead of appressed silvery hairs in the former and spreading pilose hairs in the latter. It also lacks maroon spots on the lower corolla lip. 178 Figure 47. Herbarium specimen of M. viridissima. Correll 19593 (GH). 179 Representative specimens examined: U.S.A. TEXAS. Bastrop: Albers 45062 (TEX); near McDade, Albers 46363 (TEX); along highway east of Elgin, Albers 45-130 (TEX); in woods about 2 miles east of Elgin, Correll 19593 (GH); 2 miles east of Elgin in sandy oak woods, Correll 19593 (TEX); in sandy oak woods near Bastrop, Correll 20400 (TEX); Pine hills east of Edward's saw mill, Duval s.n. (TEX); near the Pleistocene Gravel overlay at the edge of the Lost Pines, Rodgers 46529 (MSC); near Paige in sand east of Edwards' Sawmill in Pine Hills, Small 223 (TEX); adjacent highway 290 near McDade, Sperry s.n. (TAMU); in deep sand of the Carrizo in open woods near the Pleistocene Gravel overlay at the edge of the Lost Pines, Tharp 46529 (TEX); Caldwell: McBryde s.n. (TEX); east of McMahon, Tharp 52-532 (TEX); Gonzales: Waelder, Hopkins s.n. (US); Waelder, Hopkins s.n. (US); Palmetto State Park, Tharp s.n. (TEX); Hays: woods between Colorado and San Marcos, Wright 108 (US); Lavaca: E of Hallettsville on FM 3283 (County road 124) between Vienna and Subline (ca. 5 miles N of Vienna), Fryxell 4965 (MSC, TEX); Milam: ca. 2 mi N. Gause, open sandy area SE Sugarloaf, W. Long Mt. above Beaver Branch, Dubrule 1399 (TAMU); 3 miles north and west of Milano, Wolff 4843 (MSC); Washington: Albers 46127 (TEX); Wilson: Sutherland Springs, Parks 3135 (TEX); Lavernia, Parks 27802 (TEX). 180 16. Monarda n. sp. 2, sp. nov.--TYPE: U.S.A. Florida: Columbia County, Four and 0.5 miles NW of Fort White off of U.S. 27 in Ichetucknee Springs State Park, 10 August 1991, Herring 349 (holotype: FLAS!). Habit perennial, herb to 80 cm tall; upright, branching heavily from the middle of stem. Stem indumentum variable; internodes puberulent with minute downward-curled hairs and spreading longer hairs; node with short hairs, nodal plane with long hairs. Penultimate vegetative internode 27 – 65 mm long, 0.8 - 1.2 mm wide. Leaves 22 - 34 mm long, 5 - 7 mm wide; petiole 3 – 7 mm long; serrations 0.3 mm wide; leaf shape ovate, base acuminate; margins serrate from apex to 10% length of laminar tissue, apex acute; petiole indumentum of long hairs; abaxial surface hirsute with medium-long hairs around the midvein; adaxial surface glabrous; margin ciliate toward the base; both leaf surfaces green. Basal inflorescence internode 15 – 25 mm long, 0.6 - 0.8 mm wide; puberulent with occasional spreading longer hairs. Lower floral bracts 13 - 30 mm long, 3 - 5 mm wide; sessile; lower bract shape ovate to lanceolate, base attenuate, margins serrate; apex acute; abaxial surface hirsute with medium-long hairs around midvein; adaxial surface glabrous; margin ciliate toward the base; abaxial surface green, 181 adaxial surface green to pink. Upper floral bracts 13 – 18 mm long, 3.8 - 4.8 mm wide; upper bract shape elliptic, base cuneate, margins serrate; apex acute; abaxial surface glabrous; adaxial surface glabrous; margins entire; abaxial surface green; adaxial surface purple. Calyx tube 4.5 – 5 mm long, 1.5 – 1.8 mm wide, with minute scattered hairs on outer surface. Calyx lobes 0.9 – 1.3 mm long, 0.2 – 0.4 mm wide, narrowly triangular, with ciliate margins. hairs. Calyx orifice hirsute with flexible white Lower corolla lip 4.8 - 6 mm long, 2.0 – 2.5 mm wide; mid limb to 2.1 mm long, 0.9 mm wide, expanding toward the tip; lateral limbs to 1.2 mm long, 0.7 mm wide. Upper corolla lip 6.8 – 8.2 mm long, 3.1 – 4.1 mm wide, lobes to 2.6 mm long and 0.31 mm wide, corolla cream to yellow with purple pigmentation. Stamens to 28 mm in length, anthers 0.8 mm long, 0.5 mm wide, pollen yellow. Phenology. Pistil to 28 mm long. Flowering material has been collected from August through October. Distribution. Eastern U.S. from Florida to South Carolina. Monarda n. sp. 2 belongs to the narrow-leafed clade and is morphologically most similar to the widespread species, M. punctata. It can be distinguished from other species in the section by its smaller leaves (largest are <3 cm long and 1 cm 182 wide) and narrow stems lending it a spindly stature. It has serrulate upper bracts which are pink to purple in color and pale yellow flowers. Deep maroon spots are present only on the lower corolla lip. 183 Figure 48. Herbarium specimen of Monarda n. sp. 2. Herring 349 (FLAS). 184 Representative specimens examined: U.S.A. FLORIDA. Citrus: along Fla. 490A, Homosasa Springs, Genelle 1621 (FLAS); Columbia: T6S, R15E, S13 four and 0.5 miles NW of Fort White of of US 27 in Ichetucknee Springs State Park. Ca. 0.9 mile NE of South Enrance on SE side of tram road in NE portion of section 1D, Herring 349 (FLAS); Hernando: along FL 476 just w. of the Withlacoochee River, Nobleton, Baltzell 1314 (FLAS); along FL 46 near Hernando-Sumter County Line at Nobleton, Baltzell 5424 (FLAS); McIntosh: near south end of Sapelo Island, Dunun 20462 (US); St. John’s: Fruit Cove Rd., 2 mi. s. of Julington Creek, D'Arcy 1244 (FLAS); SOUTH CAROLINA. Horry: Savanah, 3.8 mi S. of Socaste (3 mi. S. of jct. of 544 & 707 on 544 & 8 mi. E on unnumbered dirt road), Massey 3387 (NCU). 185 CHAPTER 4 OVERVIEW OF ONLINE MONOGRAPH 186 Online Monograph To supplement the growing collection of online resources that expedite plant systematics research such as Index Herbariorum (The New York Botanical Garden 2007), IPNI (The International Plant Names Index 2004), or web-accessible herbaria, a number of other online resources have been developed. Among these are online floras such as eFloras (Brach and Song 2006), The Atlas of Florida Vascular Plants (Wunderlin and Hansen 2008), South Carolina Plant Atlas (Mouseau 2009) and other, more taxon-specific resources such as the Tall Fescue monograph (Aiken, Alderan et al. 2003). Furthermore, large websites, such as Encyclopedia of Life (Wilson 2003) and Discover Life (The Polistes Corporation 2011) are being developed to host such resources. Keeping with this trend, in addition to the traditional monograph presented above, I have also developed content to be hosted by Discover Life (www.discoverlife.org), a website created to document and track natural history, particularly for species in the U.S. It has a large partnership base that includes institutions such as the Academy of Natural Sciences, California Academy of Sciences, the Field Museum of Natural History, Missouri Botanical Garden, Smithsonian Institution, USGS, and the US National Park Service as well as numerous 187 universities and their herbaria. This component of the work will represent an attempt to combine modern modes of information distribution with traditional botanical monography. The online representation of the paper monograph will contain similar content to its printed counterpart, but will be constructed to provide quick access to features in which the user may be interested. It will also provide services unavailable to a print-bound publication and allow future expansion of the monograph to keep the work current and growing. Below I discuss some of these and how the online monograph will improve information access for the reader. Introductory page(s) This contains a formal introduction to Monarda subgenus Cheilyctis as well as an introduction to the use of the website. It also functions as a ―table of contents‖ whereby it provides links to the remaining chapters and services. It functions as the site hub for readers who are interested in working through the monograph in a traditional manner or for the readers that want to navigate elsewhere based on their specific interests. 188 Data As there are abundant amounts of morphological data presented in the monograph, the online monograph organizes these into accessible linked pages where appropriate in the literature. Raw data as well as matrices are made available. For the online monograph I will contribute: CSV file of morphometric data CSV file of phylogenetic data Paup file of character-coded phylogenetic data matrix Specimen Database A feature of considerable utility is the database of Monarda subspecies Cheilyctis specimens used in writing the monograph. From here readers are able to query any information commonly found on herbarium specimen labels, from collection records to phenological patterns. The database will be a work in progress so that with additional collections, the database will continue to expand and display the most current distribution information available. These data are assimilated into the mapping features of Discover Life (see below). For the online monograph I will contribute: 189 27 records of M. austromontana ined 28 records of M. citriodora 30 records of M. clinopodioides 30 records of M. fruticulosa 15 records of M. humilis 20 records of M. maritima 25 records of M. occidentalis ined 28 records of M. pectinata 105 records of M. punctata 20 records of M. stanfieldii 30 records of M. villicaulis ined 25 records of M. viridissima 2 records of Monarda n. sp. 1 7 records of Monarda n. sp. 2 Habitat information Whereas the cost of including abundant images, diagrams, maps, and informal notes on habitats keep them all too infrequent in printed journals, the online framework delivers admission to more content of this type which authors may conservatively exclude from their publications. The online monograph provides additional images of habitats and associated 190 species where available. In addition, it includes notes that are used to broadly characterize the habitats of select taxa. These notes have utility in that they may offer insights into future avenues of research, but are not robust enough for inferring or making deterministic conclusions about how the taxa interact with or have been influenced by their environments. These habitat characterizations include notes on meteorological averages, species compositions, and floral visitor observations collected during standardized field observations made by the author. For the online monograph I will contribute photographic images of live specimens of: M. austromontana ined M. citriora (with supplementary habitat image) M. clinopodioides (with supplementary habitat image) M. fruticulosa (with supplementary habitat image) M. maritima (with supplementary habitat image) M. occidentalis ined (with supplementary habitat image) M. pectinata M. punctata (with supplementary habitat image) M. stanfieldii (with supplementary habitat image) M. villicaulis ined (with supplementary habitat image) 191 M. viridissima (with supplementary habitat image) Habitat characterizations of: M. citriodora M. clinopodioides M. occidentalis M. stanfieldii M. villicaulis M. viridissima Interactive range mapping The online monograph presents a cost effective method of disseminating more habitat images, but it also provides zoomable range maps, a resource that is entirely unavailable in a printed monograph. The data for the maps are supplied from georeferenced herbarium specimens and each mapped location links to the label information of that particular specimen collection point. Specimens were georeferenced using the GEOLocate (Rios and Bart 2010) desktop application. For this process, herbarium locality information (including details such as street intersections and natural landmarks such as lakes and rivers) that is input into the client is used to generate GPS coordinates and polygonal error descriptions. 192 Another valuable feature is that the mapping software used by Discover Life can pool additional locality information from other data sources and add it to the growing database of distribution information for these taxa. Examples of other sources are other herbaria contributing to the website and other biologists that take gpslinked photographs in the field. For the online monograph I will contribute georeferenced data (including estimate of error) for the specimens listed in ―Specimen Database‖ section above. Interactive keys The online monograph provides the traditional published keys in an easily printable format as well as access to an interactive key system which increases the ease of identification to persons not familiar with the keying process as well as for those attempting identification with limited plant material. Interactive keys are increasingly commonplace (Watson and Dalwitz 1992; Sanders and Lee 2003; Weiblen and Deacon 2003), and with the free distribution of Delta system (Description Language for Taxonomy) software, they are becoming easy to construct (Dallwitz 2005; Askevold O'Brien, C). The Delta system is a suite of programs that were designed to aid taxonomic investigation and publication by manipulating large 193 data sets and having a computer present the information in a useful context. Among many of its uses is its ability to facilitate the construction of interactive keys. Instead of relying on only one or a few traits at a time, interactive keys are designed to start with any number of morphological characters which are presented to (and may be recognized by) the user. The choices are usually accompanied by diagrams or images that clearly illustrate the character states. The user simply selects any of the character states presented which match their specimen of interest and species not matching those traits are eliminated from the pool of possibilities. In this way, taxa may be narrowed down until the point at which identification is reached. The user can focus on the traits that are easier to recognize in the beginning and rely on the difficult traits only if necessary. Also, if there is a trait that is unique to the species, and the user identifies it first, then the identification will be complete in only one step, a vast improvement over the traditional, and often lengthy dichotomous keys which require the user to make decisions based on traits in the order determined by the author. At the end of each species ―trail‖ the reader will be able to directly view the species description, specimen images, images of living plants, specimen data, locality data, and range maps. 194 For the online monograph I will contribute: list of plant character states that uniquely identify Monarda subgenus Cheilyctis from all other plant species images of above-mentioned character states that the user can recognize when observing a specimen of Monarda subgenus Cheilyctis list of plant character states that uniquely identify each species of Monarda subgenus Cheilyctis images of above-mentioned character states that the user can recognize when observing a specimen of one of the species of Monarda subgenus Cheilyctis data matrix of character state values of all species in Monarda subgenus Cheilyctis for all previously mentioned characters. These items will be incorporated into the database Discover Life uses for their interactive key project. 195 APPENDICES 196 APPENDIX A Morphometric data for Monarda section Cheilyctis. Character numbers correspond to those presented in the Materials and Methods section 197 Table 3. Morphometric data for Monarda Section Cheilycits ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 Thomas Moore Thomas Locke Moore Scora Moore Roberts Bigelow Palmer Locke Moore Buchhohy Demaree Moore Foreman Letterman Demaree Fryxell Parks Garcia Bruni Higdon Tharp Albers Albers Correll Ramirez Runyon 111098 s.n. 111098 2784 400459 1056 68341 471 s.n. 12665 26 410-213 335 410 400401 s.n. s.n. 15840 1276 Rx 3132 25 24 53-57 s.n. 46040 49035 16025 82 3991 MSC UARK UARK UARK UARK US UARK UARK US US UARK UARK UARK UARK UARK US US UARK LL TEX TEX LL TEX TEX TEX TEX LL TEX RM 198 1 7.5 6.1 6.25 2.5 7.6 3.5 2.9 5.5 6 5.2 5.5 5.7 2.1 5.1 4.6 6.4 7 5.78 6.7 5.2 5.2 4.1 3.4 3.1 4.2 5.9 4.9 6.5 1.69 2 1.2 2.2 2.2 3.2 2.5 2 2.8 2.2 2 1.4 1.8 1.2 1.8 2.1 1.8 2.1 2.8 1.9 1.9 1.8 2.2 2.1 1.5 1.1 1.6 1.9 2.1 1.5 1.2 3 8.62 10.3 9.6 4.6 5.9 5.6 5.2 6.4 8.2 6.5 6.8 5.98 4.8 7.1 6.3 6.6 7.5 7.2 9 5.2 5.6 4.2 5.8 3.2 4.1 5.7 7.5 7.85 2.5 Table 3 (Cont’d) Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 Gamboa Scora Scora Lundell Runyon Runyon Dubrule Lundell Fisher Correll Runyon Pringle Shiller Tharp Carr Lundell O'Brian Garcia Garcia Santiago Corell Correll Runyon Fisher Johnston Runyon Muller Salinas Johnston 158 2224 2223 12792 3991 4333 804 12796 41125 26913 4333 178 42-40 13206 8850 1163 110 110 7785 35452 35452 3991 41125 541835 2628 8053 180 53254.2 TEX US US TEX TEX TEX TAMU US US LL US US US TEX TEX TEX RM TEX TEX TEX GH LL US RM TEX TEX TEX TEX TEX 199 2.9 2.9 3.3 4.7 4.1 2.9 1.35 1.9 2.1 1.28 3.1 2.4 3.1 3.2 1.9 2.3 6.1 2.5 4.2 1.5 1.7 1.25 2.9 2.8 1.3 1.9 2.9 4.8 3.9 1 2.9 1.9 1.5 1.8 1 1 0.8 0.9 1 2 0.8 1.8 1.2 0.9 0.8 1.8 0.9 1.1 1.1 1 0.6 1.2 1.2 1.1 1.1 1.1 1.7 1.2 2.5 4.3 4.2 3.6 3.8 2.8 3.6 1.6 1.9 1.6 4.1 2.5 4.1 1.8 1.4 3.7 4.4 3.74 3.9 1.85 2.4 1.5 3.75 3.1 1.8 2.6 1.9 4.4 3.3 Table 3 (Cont’d) Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 Wooton Wooton Wooton Tafoya Pase Abbott Pase Goodrow DeBruin McIntosh Wooton Wooton Higgins Carlson Brandt Wooton Correll Lundell Warnock Turner Lundell Tyler Carr Gentry Shinners Turner Ruth Scora Orzell s.n. s.n. s.n. 131 3621 s.n. 3566 522 448 1006 s.n. s.n. 7737 SC4 1094 s.n. 19009 14028 46402 95-179 9454 s.n. 13889 51-1784 28599 94-92 978 2011 10573 US US US UNM NMC UNM NMC UNM UNM UNM NMC NMC NMC UNM UNM US TEX TEX TEX TEX TEX TEX TEX TEX TEX MSC US TEX 200 1.55 5.15 3.9 3.7 2.9 4 4.2 1.6 3.5 2.9 4.8 5.4 2.5 3.7 3.5 1.6 8.4 6.1 5.4 4.5 3.4 4.5 7.5 5.4 4.2 7.1 6 6.4 3.4 1.8 2.2 1.6 1.2 1.9 1.1 1.3 1.8 1.8 1.2 2.1 1.6 1.6 0.9 1.8 1.2 3.1 2.9 2.8 2.2 2.2 2.1 2.4 2.5 3.5 3.4 2.1 2.6 1.8 2.8 2.9 2.8 3.4 3.8 3.1 3 2.5 4.1 3 4.8 3.1 3.4 2.55 3.72 2.2 7.5 9.3 6.2 6.3 5.5 4.2 5.1 8.1 6.5 6.8 7.55 6.4 5.4 Table 3 (Cont’d) lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 Orzell Croat Orzell Correll Orzell Crockett Orzell Albers Williges Carr Albers Tharp Strother Cory Runyon Thomas Nixon Correll Orzell Scora Saenz Marsh Scora Scora Charette Orzell Lundell Johnston Folsom 11227 3853 11189 16731 11363 790 10327 s.n. 413 18327 46097 s.n. 175 5799 4322 129407 17314 20809 10947 2219-i 83 121 2133 2133 815 10388 12782 54566 10317 TEX MO TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX MSC US MO TAMU TEX TEX TEX TEX TEX TEX US TEX TEX TEX TEX TEX 201 1 4.8 6.2 5.1 4.8 4.5 6.2 2.9 4.3 9.5 5 5.4 8.2 5.8 5.8 4.5 4.9 5 5.1 5.2 12.8 7.65 4.9 5.3 8.5 5.9 5 6.1 2.2 8.7 2 2.7 2.8 1 3.2 2.4 3.5 1.3 1.8 2.4 2.7 2.1 3.2 2.2 2.5 2.4 1.8 1.1 3.8 1.1 2.9 2.8 1.1 1.8 3.5 2.8 2.8 2.4 3.9 4.5 3 5.8 6.5 4.1 8.6 3.58 8.5 4.6 4.8 6.75 5.4 4.8 6.2 3.8 6.15 5.25 5.7 4.5 7.9 4.7 7.15 6.05 3.75 4.6 10.9 6.2 5.1 6.3 4.25 6.1 Table 3 (Cont’d) lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 Correll Scora Johnston Scora Johnston Johnston Palmer Palmer Johnston Palmer Churchill Johnston Tharp Tharp Scora Vogt Goodman Nighswonger Waterfall Warnock Scudday Correll Carletan Drake Folley Correll Bellah Nighswonger Wagner 26370 2170 53254.2 2170 53254.2 53254.2 1081 1081 53254.3 1081 90-930 53254.2 1553 5553 2170 7 8379 3270 14926 7878 s.n. 33017 183 108 Nr. 580 16859 76 1925 3615 202 TEX TEX TEX US TEX TEX US US TEX US MSC RM US US GH ARIZ OKL OKL US LL LL LL MSC TEX OKL TEX OKL OKL MO 1 5.35 2.3 1.8 2.8 2.25 2.3 2.8 1.8 2.7 2.5 3.1 2.3 2.3 1.45 2.9 3.9 3.6 3.8 2.7 1.2 5.5 6 4.2 3.9 4.5 2.1 4.2 3.6 3.1 2 1.8 1.2 1.8 1.8 1.8 1.8 2 2.2 2.2 2.8 1.8 1.8 2.4 0.9 2.4 1.35 1.9 1.8 2.9 2.1 1.2 2.1 2.05 2.5 1.3 2.1 2 1.8 1.8 3 4.7 4.5 2.8 4.05 3.5 2.95 4.05 4.05 4.6 4.2 4.6 2.9 3.2 4.2 3.6 3.4 6.1 4.1 5.1 4.4 5.7 4.6 3.85 4.4 5.6 4.2 4.8 4.1 3.5 Table 3 (Cont’d) occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 Cory Palmer Reed Barclay Henderson Warnock Stuessy Springer Goodman McCarthy Laverty Boon Correll Massey Dunun Nelson Nelson Bozeman Steele Rudd Boon Fleetwood Credle Traverse Kirkman Tharp Genelle Baltzell Baltzell 51996 14116 3437 851 63-834 15444 157 374 8090 ALT0280 7343 175 18925 3387 20462 685 4243 11643 s.n. 3940 388 9154 2597 169 571 s.n. 1621 5424 1314 US US US US TEX LL TEX OKL OKL OKL OKL TEX TEX NCU US FLAS NCU NCU MSC TEX TEX TEX NCU MSC FLAS TEX FLAS FLAS FLAS 203 1 3.8 4.1 4.4 4.4 3.1 2.6 4.9 3.55 4.6 5 2.25 5.5 6.8 3.3 6.5 5.4 2.6 6.8 4.8 5.4 5.6 5.4 5.3 8.2 5.1 6.5 3.1 3.3 3.2 2 2.2 2 2.8 1.75 1.8 1.8 3.05 1.8 2 1.9 2.5 1.8 2.1 1.2 1.2 1.5 2.2 2.1 1.8 1.8 1.2 0.8 2.4 2.1 1.6 2.1 0.9 1.2 1.2 3 3.8 4.2 5.4 4.6 4 4.2 4.3 3.95 5.2 4.6 3.9 7.4 6.2 3.2 3.4 7.1 6.9 7.2 6.8 4.6 6.4 7.4 8.3 6.1 7 7 3.1 3.2 2.9 Table 3 (Cont’d) punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 D'Arcy Herring Tharp Tharp Wolff Lundell Albers, Sammons Albers Tharp Tharp Tharp Lundell Walters Barkley Albers Lundell Miller Miller Hill Atwood Goff Drew Parmelee Drew Drew Churchill Gilis Eickwort 1244 349 s.n. s.n. 3859 9029 49070 s.n. s.n. s.n. s.n. s.n. 15109 593 46262 s.n. 15109 542 1828 15468 406 1002 M171 819 M264 M270 s.n. 5889 68 FLAS FLAS TEX MICH MSC MICH CAS MSC TEX TEX MICH MICH MICH TAMU TEX TEX US MSC MO US MSC MSC MSC MSC MSC MSC MSC MSC MSC 204 1 2.9 2.7 6.2 5.5 7 7.5 6.8 7.2 5 7 10.3 7.9 5.7 8.1 6.8 7.2 7.9 3.4 6.7 6.5 4.1 3.6 3.3 2.2 3.7 4.2 4 3.8 3.7 2 0.8 1.1 1.6 1.8 2.4 2.2 2.4 2.5 2 2.2 1.8 1.8 1.8 1.9 3.2 2.9 1.5 1.5 2 2 1.7 2.1 2 1.8 2.2 1.2 1.8 1.2 1.5 3 2.7 2.2 4.6 4.8 5.3 7.3 5.2 5.8 6.9 6.2 5.9 4.6 4.4 4.8 10.5 3.8 6.6 6 7.5 7.6 4.5 5.2 4.8 5.3 5.8 5.5 5.7 5.8 4.8 Table 3 (Cont’d) 1 vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 Dodge Bourdo Gilly Hoffman Hestbeck Parmelee Shadick Nielsen Scora Bourdo Skeels Gilly CP Pepoon Gillis Bourdo Ryel Albers Tharp Correll Correll Duval Albers Correll Tharp Hopkins Dubrule Fryxell Fryxell 509 306 1078 310 10 1015 s.n. s.n. 498 20042 s.n. 7172 302 s.n. 5415 30340 L120 40021 46529 19593 20400 s.n. 45-130 19593 52-532 s.n. 1399 4965 4965 MSC MSC MSC MSC MSC MSC MSC MSC US MSC MSC MSC MSC MSC MSC MSC MSC TEX TEX TEX TEX TEX TEX GH TEX US TAMU TEX MSC 205 4 3.5 4.2 4.3 3.65 2.1 3.2 3.85 3.5 3.4 3.5 3.75 3.6 4.2 3.8 3.75 3.6 2.2 2.5 3.15 2 2.3 2.45 2.5 4.9 2.4 2.45 2.4 2.6 2 2.1 1.6 1.7 1.2 2.1 1.9 1.9 2.2 1.8 1.8 2 1.8 1.7 1.9 1.8 2.2 1.7 1.9 1.9 1.8 1.2 1.8 0.9 1.2 1.3 1.9 0.9 1.2 1.8 3 8.4 7.1 6.8 5.4 6.2 5.2 5.1 5.3 4.6 5.7 6.1 5.5 7 8.6 4.7 5.9 5.9 3.1 5 3.9 2.5 3.59 2.92 5 4.45 3.24 3.12 4.1 5.2 Table 3 (Cont’d) Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 Parks Albers McBryde Parks Wright Hopkins Sperry Wolff Albers Rodgers Small Albers Tharp 3135 45062 s.n. 27802 108 s.n. s.n. 4843 46363 46529 223 46127 s.n. TEX TEX TEX TEX US US TAMU MSC TEX MSC TEX TEX TEX 206 1 2.3 2.45 2.5 2.4 2.5 2.3 2.52 2.35 4.2 2.7 2.56 2.27 2.2 2 1.5 1 1.2 1.6 1.7 1.2 1.1 1.6 1.2 1.4 1.2 1.5 1.4 3 3.7 3.1 4.85 3.2 4.6 3.8 3.2 3.5 4.12 4.9 3.82 3.5 2.8 Table 3 (Cont’d) ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 4 1.12 1.4 1.6 0.75 0.9 1.08 0.82 1.2 1.4 0.94 0.9 0.92 1.05 1.1 1.3 1.3 1.1 1.3 1.9 0.7 0.8 0.71 0.84 0.49 0.8 0.8 1.2 0.9 0.2 5 3.45 4.8 4.4 1.8 1.8 1.6 1.6 3 4.2 3.2 2.8 1.7 1.3 2.5 2.2 2.3 2.9 2.7 3.9 2.1 2.8 2.7 3.2 1.6 2.6 2.7 4.7 2.7 0.85 6 16.5 24 18 4 6 6 13 16 15 12 15 7 7.8 14 11 13 12 12.4 17 10 9 7 13 5.1 9.8 12 19 13 0.2 7 4.9 6.3 5.8 2.85 3.4 4.1 2.75 3.4 3.8 3.2 2.4 2.7 2.6 4.6 3.1 3.4 5.1 4.6 5.1 2.7 3.5 2.3 3.1 1.9 2.4 2.9 3.7 4.8 0.69 207 8 4.95 5.6 3.8 2.45 2.2 5.8 2.4 3.2 3.4 2.1 2.65 2.9 2.8 3.5 3.2 3.5 2.1 5.15 7.1 4.1 5.2 3.2 6.1 2.1 3.2 2.6 4.8 3.8 3.3 9 0.85 0.4 0.5 0.15 0.3 0.2 0.18 0.38 0.4 0.3 0.2 0.2 0.3 0.24 0.2 0.25 0.5 0.29 1.1 0.3 0.42 0.46 0.9 0.2 0.46 0.46 0.65 0.45 0.15 10 6.97 7.9 7.8 4.2 5.3 5 3.9 4.8 6.7 5.3 5.3 5.28 4.02 5.7 5.2 5.3 6.3 5.96 7.3 4.2 4.7 3.5 4.5 2.69 3.12 4.5 5.6 6.55 2.48 11 1 3 4 7 4 2 4 3 2 3 4 2 4 6 2 4 3 1 3 2 2 4 3 4 2 3 3 1 5 Table 3 (Cont’d) 4 Frut237 0.2 Frut238 0.3 Frut239 0.22 Frut240 0.36 Frut241 0.19 Frut242 0.11 Frut243 0.91 Frut244 0.21 Frut245 0.21 Frut260 0.19 Frut261 0.27 Frut262 0.12 Frut263 0.24 Frut264 0.22 Frut46 0.13 Frut47 0.9 Frut48 0.21 Frut49 0.36 Frut50 0.19 Frut51 0.2 Frut52 0.22 Frut53 0.15 Frut54 0.3 Frut55 0.35 Frut56 0.18 Frut57 0.26 Frut58 0.19 Frut59 0.88 Frut60 0.21 5 1.7 2.2 2.6 1.8 1.8 1.6 3 0.1 1.6 0.8 2 1.5 2.7 1.2 0.9 1.9 2.4 2.7 1.8 1.5 1.4 0.96 1.95 2.4 1.5 1.2 0.65 3.4 2.21 6 0.23 0.2 0.1 0.1 0.2 0.18 0.18 0.1 0.3 0.18 0.2 0.22 0.1 0.1 0.4 0.6 0.4 0.6 0.2 0.1 0.2 0.1 0.2 0.3 0.3 0.2 0.1 0.3 0.2 7 0.75 2.1 1.4 1.1 1.1 1.2 1.2 0.5 0.55 0.8 1.8 1.1 1.3 0.78 0.41 1.1 0.7 1.1 1.21 0.5 0.72 0.44 1.1 0.9 0.7 0.77 0.57 1.4 1.1 208 8 2 2.65 2.2 5 2.2 2.05 2.05 2 1.9 2.05 2.2 2 2 1.8 1.7 1.8 4.05 2.5 3.2 1.05 1.5 1.9 1.8 2.55 1.8 1.75 1.6 1.45 2 9 0.15 0.25 0.2 0.2 0.2 0.2 0.2 0.2 0.15 0.1 0.18 0.15 0.18 0.2 0.15 0.2 0.2 0.2 0.15 0.15 0.18 0.18 0.2 0.3 0.25 0.2 0.18 0.25 0.18 10 2.477 4.28 4.19 3.59 3.78 2.782 3.582 1.59 1.87 1.582 3.8 2.4 4.09 1.79 1.36 3.64 4.36 3.68 3.88 1.84 2.38 1.49 3.73 3.07 1.77 2.58 1.89 4.37 3.28 11 5 6 3 3 6 4 2 4 4 7 4 6 3 8 4 5 1 4 6 9 4 7 6 3 7 7 6 4 6 Table 3 (Cont’d) Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 4 0.48 0.61 0.51 0.6 0.6 0.6 0.52 0.3 0.59 0.62 0.72 0.61 0.72 0.58 0.78 0.7 0.9 1.2 0.8 1.1 0.8 0.8 0.95 1.3 1.1 1.2 1.2 0.9 0.45 5 1.4 1.6 1.7 1.4 1.7 1 1.3 1.3 2.4 1.6 1.6 1.15 0.86 0.9 1.5 0.9 3.1 4.9 3 2.9 2.3 2.1 1 4.5 2.8 2.6 2.7 2.2 2.1 6 3.8 4 6 6 0.9 4.5 2.5 7 8 3 7 3.9 7 3.5 6 4.2 17 18 8 9 6 4.5 5 10 9 10.2 12 5 6 7 1.6 1.4 1.9 2.1 1.4 1.6 1.35 1.3 1.45 1.6 2.7 1.75 1.9 1.4 2.6 1.4 4.6 5.6 3.9 3.7 3.5 2.7 3.7 4.9 3.8 3.65 4.1 3.6 1.9 209 8 2 1.25 2.75 1.6 4.75 2.7 3.3 5.5 5.5 5.25 2.3 1.15 4.25 2 2.8 0.6 6 5.4 5.4 2.95 4.15 3.15 3 4.85 3.1 4.2 4.8 5.5 6 9 0.2 0.3 0.25 0.2 0.2 0.3 0.25 0.16 0.2 0.2 0.15 0.15 0.15 0.2 0.3 0.15 0.45 1.2 1.1 0.3 0.35 0.4 0.5 1 0.75 0.5 0.7 0.55 0.25 10 2.42 2.5 3.32 2.8 3.71 2.65 2.75 1.8 3.3 2.7 4.1 2.71 2.7 2.2 3.12 1.78 5.8 7.5 5.4 5.4 4.9 3.75 4.6 7.1 5.6 5.78 5.68 5.9 4.8 11 6 6 8 6 4 7 11 6 6 10 7 7 6 8 3 3 1 4 5 2 5 4 2 4 3 5 4 3 3 Table 3 (Cont’d) lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 4 0.5 1.5 0.65 1.4 0.75 1.4 0.56 0.38 1.2 0.65 0.85 0.7 0.34 1.1 0.6 0.8 0.75 0.75 0.7 1.5 0.7 0.7 1 1.1 1.2 0.5 1.2 0.81 0.65 5 2.1 2.6 1 2.5 1.75 3 1.3 1.4 3.5 2.4 1.65 2.1 1.3 2.8 2.4 2.4 1.6 3.6 1.7 3.8 3.4 1.8 1.7 3.9 2.8 2.3 3 1.9 1.8 6 7.5 6.2 5 1.3 6.3 1.7 7 5.5 9 7.5 4.5 8.5 7 12.2 8 8.2 6 15 6 8.5 8.5 6 4.5 9 22 7.2 13 9 9 7 3.2 3.8 2.1 4.5 2.2 5.1 2.3 3.3 4.8 2.8 4.1 3.9 1.8 4.2 3 3.8 2.1 4.55 3.5 5.1 3.4 1.85 3.7 5.3 3.45 2.95 4.1 2.75 4.1 210 8 4.1 2.5 2.6 2.1 3.4 2.8 3.4 1.8 4.6 4.35 2.4 3.6 5.4 4.35 4.3 3.15 2.7 4 3.1 4.65 4.2 3.5 2.2 4.75 2.4 4.2 4.25 2.45 3.5 9 0.3 0.62 0.3 0.42 0.28 1.1 0.32 0.75 1.1 0.21 0.59 0.28 0.3 0.58 2.2 0.3 0.35 0.7 0.4 1.4 0.85 0.3 0.58 0.7 0.6 0.2 0.6 0.45 0.25 10 5.05 5.88 2.9 8.47 2.95 8.33 3.9 4.25 5.85 4.65 4.35 5.35 3.1 4.93 4.45 4.88 3.1 6.4 3.15 6.3 5.2 3.15 4.33 10 4 4.38 5 3.35 5.2 11 1 6 4 2 2 2 3 4 4 3 4 1 4 3 4 4 3 5 4 1 1 2 1 3 2 6 4 6 3 Table 3 (Cont’d) lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 4 1.1 0.7 0.8 0.75 0.45 0.4 0.8 0.72 0.79 0.68 0.6 0.41 0.48 0.6 0.7 0.6 1.3 0.9 0.6 0.91 1.3 1.1 0.9 1.1 1.2 0.9 1.3 0.75 1.1 5 1.75 1.6 0.7 0.5 0.7 2 1.4 1.4 1.5 1.5 1.4 1.2 1.3 1.4 1.4 1.8 2.7 1.3 3.6 1.8 2.6 2.1 2.2 2.55 2.5 1.6 1.4 1.4 1.4 6 4 1.9 0.5 1.6 1.3 0.8 1.2 1.2 2 1.5 2.1 0.8 1 1.3 1.2 8.6 7.2 5.2 6.5 7 11 10 12.5 7.8 10.5 6.4 7 5.1 8 7 3.9 2.1 1.6 2.9 1.9 1.2 2.9 2.9 2.2 2.3 2.2 1.6 1.9 2.15 2 0.16 3.9 3.5 2 2.5 3.05 2.9 1.6 3.1 3.1 2.4 3 3.3 2 211 8 2.35 5.8 6.25 3.75 2 2.85 3.5 3.5 4.65 4.1 4.25 2.8 3.5 4.62 4.25 2.45 1.65 2.2 3.25 3.05 2.8 4.2 2.35 3 2.7 3 4.1 2.4 2.4 9 0.6 0.45 0.2 1 0.25 0.3 0.6 0.6 0.35 0.32 0.42 0.25 0.3 0.42 0.3 0.2 0.55 0.3 0.3 0.5 0.35 0.55 0.25 0.6 0.34 0.42 0.3 0.32 0.3 10 4.3 4.31 2.75 3.89 3.37 2.87 3.93 3.93 4.4 4.05 4.39 2.82 3.1 4.07 3.48 2.54 4.7 3.58 4.45 3.7 4.6 3.6 2.6 3.62 4.55 3.56 4.1 3.59 2.7 11 4 1 2 3 4 1 1 1 3 2 4 3 2 1 2 9 4 13 10 14 6 7 6 1 5 4 10 5 3 Table 3 (Cont’d) occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 4 1.2 1 1.1 0.75 1 1.2 1.6 0.75 1.1 0.8 0.7 1.4 1.3 0.59 0.69 0.92 1.35 1.3 0.9 0.98 1.3 1.2 0.95 1.1 0.9 1.53 0.6 0.7 0.59 5 1.4 1.5 2.3 1.7 2.4 1.7 1.8 1.9 2.85 2.2 1.9 2.3 0.8 1.35 1.5 2.1 1.7 2.1 1.9 1.7 1.6 1.9 2.8 1.8 2 2.3 1.4 1.2 0.7 6 8 0.54 9.3 7 7 6.5 6 6 11 7 5 12 14 5.5 7 11.4 12.2 12.5 9.5 11 10.5 9 15 9 9.8 14.5 7 6 5 7 2.6 3.2 2.8 2.6 3.2 1.9 2.1 2.3 2.55 3.5 2 5.2 4.6 2.1 1.9 5.1 4.2 4.8 4.1 3.8 3.8 3.6 4.75 4.5 4.5 4.75 2.2 1.4 1.9 212 8 2.5 2.2 3.1 2.25 2.8 3.2 3.1 3.3 1.85 1.35 2.1 6 5.5 2.1 2 3.4 3.5 6 5.4 2.33 4.8 4.2 5.6 5.7 3.7 6.15 2 3.5 2.2 9 0.35 0.3 0.4 0.25 0.3 0.43 0.45 0.3 0.25 0.3 0.25 0.8 0.5 0.2 0.2 0.48 0.8 0.7 0.65 0.6 0.5 0.48 0.65 0.58 0.4 0.76 0.25 0.2 0.19 10 3.25 3.6 4.47 3.9 3.3 3.5 3.7 3.35 4.1 3.9 3.4 6.2 4.8 2.9 2.8 6.06 5.68 5.95 5.85 4.36 5.35 6.3 6.8 5.2 5.95 5.55 3.4 2.6 2.4 11 2 3 7 6 1 7 11 6 6 5 11 2 7 4 2 4 5 6 3 4 3 1 2 2 5 2 5 5 3 Table 3 (Cont’d) 4 punc304 0.6 punc305 0.68 Stan271 1.3 Stan272 1.5 Stan273 1.6 Stan274 2.3 Stan275 1.75 Stan276 1.8 Stan91 2.2 Stan92 2.3 Stan93 2.1 Stan94 1.55 Stan95 1.9 Stan96 1.6 Stan97 2.1 Stan98 1.4 Stan99 1.8 vill156 1.2 vill157 1.5 vill162 1.7 vill207 0.95 vill208 0.98 vill209 0.95 vill210 1.2 vill211 1.1 vill212 0.98 vill213 1.1 vill214 1.1 vill215 0.8 5 0.7 1.2 2.1 2.2 2 2.3 2.4 2.6 1.9 2.7 1.7 2.1 1.4 2.4 5 2 2.6 2.9 3.7 4.1 1.35 0.17 1.5 2.1 1.6 2.8 2.2 2.4 1.8 6 3 5 10.2 11.2 14.6 19 12.5 13.2 15.4 17 16 10.5 7.5 14 24.5 7 14 9 13 9 9.4 4.5 8.3 7.8 7.5 10.2 10.5 11 9.2 7 1.4 1.2 2.6 2.8 3.2 3.42 2.8 3.5 5 3.8 3.4 2.8 3.65 3.2 6.1 2.2 4.55 2.9 5 5.4 2.3 3.2 2.6 3.3 3.2 2.2 2.8 3.3 2.3 213 8 1.8 0.9 3.16 3.27 3.14 3.88 2.91 3.5 2 4.32 3.51 3 3.3 2.9 5.25 2 3 3 6.25 6.6 4.3 5.8 3.2 4.8 2.8 3.2 3.5 4.8 4.85 9 0.18 0.3 0.82 0.76 0.86 0.98 0.83 0.85 0.7 1.1 1.45 0.55 0.95 0.85 1.4 0.55 0.8 0.4 0.75 0.78 0.32 0.31 0.25 0.35 0.18 0.22 0.3 0.4 0.26 10 2.4 1.7 3.58 3.68 3.84 4.6 3.95 4.48 5.36 4.5 4.3 3.55 3.65 3.4 8.05 3.1 5.2 5.1 6.2 6.28 3.56 4.75 3.8 4.7 5.05 4.48 4.65 4.7 3.88 11 4 5 4 3 5 4 2 1 2 5 4 6 5 4 5 4 4 1 1 5 2 2 6 4 6 4 5 6 4 Table 3 (Cont’d) vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 4 1.5 1.3 1.2 1.08 1.5 1.1 0.98 0.98 0.8 1.2 1.4 1.2 1.3 1.7 0.8 1.3 1.3 0.41 0.45 0.45 0.25 0.55 0.5 0.32 0.42 0.41 0.5 0.41 0.47 5 3 3.85 2.4 3.1 1.8 1.8 1.6 1.78 1.4 2.2 2.8 1.6 4.3 3.8 1.6 2.3 2.7 0.9 2.2 1.3 0.85 0.6 0.7 1.6 1.2 0.85 0.7 1.1 2.3 6 6.8 8.5 9.2 11 8 8.1 7 5 6 8.4 9 8 8.2 9 5.6 12 8 0.1 4.9 3.5 1.2 2.8 4.5 4.2 4 1.2 4.2 4.2 4.8 7 4.2 4.8 3.5 2.4 3.3 2.8 3 3.5 2.2 3.2 3.3 3.6 6 4.1 2.1 3.5 3.2 2.6 2.7 1.3 1.45 0 1.32 3.2 2.9 2.5 1.25 3.15 2.65 214 8 4 4.8 3.6 3.1 2.6 4.5 3.8 6.25 4.2 5 3.5 5 6 4.05 4.95 5.2 3.7 1.45 5.9 1.8 1.15 0 4.75 7.5 3.75 1.45 4.5 7.45 6.2 9 0.28 0.38 0.24 0.2 0.15 0.31 0.28 0.3 0.4 0.5 0.4 0.35 0.4 0.3 0.25 0.6 0.4 0.25 0.24 0.23 0.18 0 0.18 0.2 0.28 0.25 0.19 0.18 0.22 10 7.72 6.25 5.88 4.3 5.4 4.5 4.25 4.8 4 4.86 5.2 4.7 6.18 7.7 4.14 4.7 5.1 3.09 4.51 3.55 2.38 3.31 2.47 4.58 4.05 3.12 2.7 3.68 4.72 11 5 3 2 5 3 5 2 3 4 2 3 4 2 4 6 5 2 2 3 4 8 3 4 4 9 2 5 3 2 Table 3 (Cont’d) Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 4 0.52 0.51 0.42 0.44 0.45 0.42 0.53 0.4 0.42 0.48 0.5 0.46 0.36 5 0.7 0.8 1.2 1.4 2 1.4 0.7 0.82 1.2 1.3 0.89 0.72 0.72 6 3.3 2.9 3.4 3.6 4.5 4.2 4.1 1.4 3.8 4.3 4.2 2.5 1.4 7 1.3 1.5 3.1 1.4 2.5 3.2 1.5 2.4 2.8 3.1 1.8 2.2 1.65 215 8 2.2 4.5 4.3 1.6 6.3 6.4 3.8 2.2 4.2 6.5 4.1 2.58 1.15 9 0.19 0.18 0.19 0.22 0.2 0.2 0.22 0.18 0.24 0.23 0.19 0.23 0.21 10 3.37 2.81 4.51 2.84 4.15 3.38 2.79 3.36 3.74 4.47 3.4 3.25 2.66 11 2 4 5 2 3 2 4 3 7 4 5 2 5 Table 3 (Cont’d) ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 12 6.3 5.8 5.2 3 5.6 4.65 3.8 4.5 5.2 4.8 4.5 3.8 4.1 3.7 4.8 5.7 4.6 5.08 4.9 3.4 5.7 3.3 4.1 3.3 4.2 4.9 6.8 5.3 1.45 13 8.2 12.2 8 6.8 8 9.2 6.6 7.2 7.8 6.4 5.2 8.6 6 6 8.5 10 8.2 7.8 6.8 4 10 6 6.9 3.2 7.2 4.4 9 4 3 14 2.6 2.6 2.1 1.2 2.1 1.55 1.6 2.18 2.4 2.2 2.1 1.5 1.4 1.1 2.4 1.8 1.5 1.8 3.1 1.6 2.5 1.1 1.7 1.4 1.8 2.4 4.2 2.2 0.3 15 2.55 2.95 2.6 1.3 2.08 1.9 1.4 2 2.7 2.7 1.5 2.6 1.5 2.1 2.1 2.4 2.2 2.3 2.5 2.4 2.1 1.3 1.9 1.6 2 2.1 2.8 2.6 0.8 216 16 9.5 11.5 8 6.5 9.1 13 5.8 6 11 10 8.1 8.2 8.5 9.1 8.2 8 8.5 9.8 8.2 7.5 7 6.1 6.5 5.1 7.8 7.8 9 9.8 4.5 17 1.15 1.05 1.1 0.7 0.85 0.7 0.82 0.9 1.2 1 0.82 1.2 0.5 1 1 1.1 0.9 1.1 1.2 0.75 0.7 0.6 0.9 0.65 0.7 0.75 1.9 0.8 0.3 18 9.24 8.16 7.82 6.29 8.43 8.59 6.62 7.1 7.66 7.54 6.85 7.52 9 6.8 7.56 7.59 7.61 6.82 6.26 6.73 7.29 5.82 6.26 5.28 6.02 5.36 5.9 5.7 5.34 19 1.53 1.87 1.52 1.36 1.42 1.52 1.42 1.58 1.62 1.55 1.59 1.64 1.5 1.45 1.38 1.58 1.56 1.54 2.43 2.68 2.23 2.05 1.87 1.78 1.75 1.94 1.8 1.85 1.52 Table 3 (Cont’d) Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 12 3.7 2.4 3.7 2.1 3.2 2.8 2.8 1.6 1.8 1.9 2.8 1.45 3.5 1.4 1.4 2.6 3.15 5.6 2.15 1.6 1.6 0.89 2.9 1.65 1.49 1.3 1.2 3.1 1.2 13 3.2 4.5 5 6.5 3.1 3.5 2.4 2.5 7 3 4.4 3.5 4.7 3.1 3.1 3.4 4.8 5.1 3 2.2 4.5 2.8 2.8 4 4.5 4 2.2 3.8 2.9 14 1.7 0.7 0.7 0.4 1.1 0.45 0.64 0.5 1.4 0.3 0.6 0.3 0.68 0.3 0.3 0.6 1.6 3.2 0.25 0.4 0.6 0.42 1.1 0.6 0.55 0.35 0.3 0.6 0.5 15 1.35 1.35 1.2 1.9 1.5 1.55 1.7 1.2 1.6 1.1 1.27 1 1.18 1.1 0.9 1.32 1.9 1.9 1.05 1.1 1.35 0.84 1.5 1.15 1.92 0.9 1.2 1.2 1.6 217 16 7 7 8 8.5 9 6 7.1 7 5 3.8 7.1 4.8 6.8 4.75 5.2 7.1 7.1 7 3.5 5.1 7.5 4.1 4.5 6 5 12 6 5 7.1 17 0.6 0.45 0.6 0.55 0.7 0.71 0.6 0.61 1.3 0.6 0.5 0.4 0.58 0.6 0.72 0.65 1.09 0.9 0.5 0.7 0.75 0.5 0.8 0.65 0.7 0.06 0.7 0.2 0.8 18 6.23 6.22 6.02 5.26 5.34 4.66 6.13 4.86 5.36 5.21 6.1 5.48 5.14 6.15 6.15 5.58 5.96 5.22 5.09 5.54 6.51 5.14 5.68 5.27 5.28 6.34 5.4 5.29 5.41 19 1.37 1.6 1.39 1.31 1.38 1.6 1.55 1.25 1.59 1.5 1.49 1.41 1.36 1.52 1.52 1.42 1.43 1.35 1.31 1.31 1.35 1.31 1.2 1.49 1.56 1.68 1.33 1.29 1.21 Table 3 (Cont’d) Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 12 2.3 2.6 3.4 2.9 2.9 3.5 2.7 2.2 3.7 3.6 3.3 3.1 3.8 2.4 3.6 2.4 4.2 6.9 4.6 5.4 4.9 4.7 4.7 4.6 5.5 5 5.4 6 4.1 13 4 6 5.1 6 3.5 5 7 3 7.3 3.5 8 6 4 2.2 7.2 4 9.5 10.5 7 8 6 7 8 7.8 6.5 8.2 7.4 9.5 4 14 1.3 1.7 1.5 1.3 1.1 1.4 1.9 0.8 2.4 2.1 2.1 1.2 1.1 1.2 1.7 1.1 1.3 2.5 1.1 3 1.7 1.6 1.3 1.3 2.2 2.2 1.8 2.1 1.6 15 1.7 1.6 1.8 1.25 1.3 1.4 2.2 1.5 1.1 1.4 0.19 1.25 1.4 0.98 1.3 1.1 2.7 2.8 2.25 2.4 2.1 2.15 2.2 2.1 1.8 2.2 2 3.15 1.6 218 16 6.5 0.8 6 8 4.8 4 5.1 5.1 7.3 7.1 9.2 6 5 5.6 4.5 5.1 10.5 12 7 12 9 11 13 9 8.5 9.5 9 11 5.9 17 0.9 1 1.2 0.3 0.36 0.21 1.2 0.66 0.5 0.9 0.89 0.5 0.85 0.45 0.3 0.49 1.1 0.9 7 0.8 1.1 0.9 0.7 0.9 0.75 0.85 9 1.4 0.7 18 6.24 5.25 5.26 6.54 5.53 7.23 5.25 5.65 7.33 5.45 5.23 5.09 5.64 5.72 5.24 6.23 6.43 7.01 6.65 6.54 6.52 6.54 6.95 7.1 6.04 7.16 7.55 7.02 6.1 19 1.84 1.24 1.04 2.39 1.59 1.41 1.56 1.62 1.8 1.78 1.72 1.5 1.74 1.52 1.48 1.69 2.04 1.74 1.19 1.47 1.44 1.06 1.44 1.39 1.52 1.4 1.5 1.41 1.51 Table 3 (Cont’d) lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 12 3.56 5.4 5.4 6.5 5.6 7.2 5.5 4.3 6.65 3.24 3.5 5.15 3.2 4.12 3.2 2.9 5.5 7.12 5.6 6.8 7.5 5.7 3.4 7.9 5.55 3.15 4.05 4.4 5.2 13 4.8 8.3 7.8 11.5 7.85 8 7.8 3.2 7.15 4.8 8.5 6 2.2 7.2 4.8 4.5 8.3 5.5 8.8 13 6.5 8 9 7.5 8.2 4.5 7.3 4.1 6.2 14 0.85 2.1 2.1 1.4 2 3.8 2.2 2.2 2.2 1.2 1.4 1.8 1.6 1.4 1.1 0.45 2 2.6 2.2 1.9 4.5 2.1 0.8 3.2 2.2 0.8 1.1 2.6 0.8 15 2 1.8 2.1 1.9 2.1 2.8 2.2 1.9 1.9 2 1.8 2.2 1.8 2.2 1.8 1.5 1 2.2 2.1 2.3 2.1 2.1 1.7 2.6 2.1 1.8 2.2 2.1 2.1 219 16 7.4 7.8 6.2 11.2 6.5 9.5 5.5 6.6 8.8 8.2 7.2 6.4 5.5 7.6 7.2 6 6.5 7.2 6.6 10 8.3 2.8 6.4 8 8 7 7.5 9 5.2 17 0.95 0.92 1.1 0.89 0.98 0.8 1.04 0.72 1.15 1.04 0.82 0.98 1.4 1.08 1.05 0.6 1 0.7 1.4 1.2 0.65 1.05 0.73 0.6 0.95 1 1.5 0.7 0.9 18 6.61 7.17 7.7 8.32 7.48 7.18 7.1 6.92 7.52 7.12 7.2 7.06 6.98 7.42 7.15 8.12 7.69 5.8 8.62 6.84 7.91 7.26 7.16 7.92 7.12 7.13 7.03 7.16 7.01 19 1.52 1.84 1.92 1.51 1.68 1.35 1.48 1.51 1.49 1.36 1.42 1.53 1.55 1.42 1.41 1.38 1.41 1.55 1.51 1.39 1.32 1.35 1.38 1.78 1.45 1.38 1.42 1.36 1.37 Table 3 (Cont’d) lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 12 3.45 4.9 3.5 2.6 2.2 2.55 4.5 4.5 3.7 3.1 3.2 2.7 3.1 3.45 5.2 2.9 3.15 3.6 3.9 3.2 4.05 5.5 3.2 2.9 4 3.4 3.8 3.7 3.5 13 9.5 6.4 5 11 4.8 7 8.6 4.8 6.2 4.85 6.8 3.8 7 4.3 5.4 5 6 5.5 6.2 8 7 10 8.5 10 6.5 7.2 6 5.8 6.9 14 0.73 2.1 1.3 1.1 0.9 0.3 1.9 1.9 0.9 1.3 1.8 1.1 0.9 1.5 1.6 1.3 1.7 1.8 2.4 1.8 2.3 2.4 1.3 1.5 2.2 2 2.2 1.8 1.8 15 1.6 1.55 2 1.7 1.2 1.45 1.55 1.55 1.5 1.42 1.5 1.2 1.3 1.45 1.9 1.9 1.9 1.9 1.7 1.85 2.4 2.8 2.1 2.5 2.1 1.75 1.8 2.1 2.2 220 16 6.5 8 6 8 8.5 9.5 6 6 9.3 7.2 7.6 7.4 7.9 8.2 8.5 7 8 8.5 8 7.5 9 8.5 10.5 9 9.4 7.7 8.2 8.4 8.5 17 0.72 0.9 1 0.9 0.6 0.65 0.75 0.75 0.6 0.72 0.75 0.6 0.55 0.9 1 1.1 0.9 0.75 0.9 0.9 1 1.6 1.1 0.9 1.2 1 1.1 0.84 0.7 18 7.04 6.83 5.79 7.54 5.92 5.81 6.12 5.86 5.92 6.24 6.11 6.13 6.21 6.72 7.69 5.71 7.29 6.26 5.9 6 6.05 5.23 6.68 6.8 5.7 6.35 7.15 6.2 6.1 19 1.35 1.57 1.47 1.65 1.44 1.5 1.48 1.3 1.42 1.42 1.34 1.5 1.51 1.47 1.65 1.32 2.12 2.04 1.41 1.77 1.82 1.59 1.93 1.6 1.3 1.86 2.05 1.82 1.45 Table 3 (Cont’d) occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 12 3.7 3.5 4 3.4 3.6 2.9 3.9 3.3 4.7 4.7 3.4 6.2 5.2 2.3 2.6 5.8 5.7 5.5 6.1 3.5 5.2 4.8 6.1 5.4 5.9 6.2 2.2 2.6 1.8 13 7.6 5.9 8.4 7 8.5 5 12 7.9 7.5 7.1 4.5 11 9.5 3.5 3.9 9.8 10.5 10.2 11 9.4 9.4 8.8 9.4 9.8 10 12.4 3.2 4.8 4.1 14 2.1 2.1 2.2 1.7 1.6 1.6 2 1.2 2.45 2.2 1.2 2.1 1.6 1.2 1.2 1.4 1.8 1.9 2.2 0.9 1.6 1.3 2.3 1.8 1.7 2.4 1.3 1.2 1.1 15 2.1 2 2.1 1.9 2.1 2 2.1 1.8 2.05 2.3 2 3.2 3.2 1.7 1.8 3.5 3.2 3.1 3.3 1.65 2.9 2.8 3.5 3.1 3.4 3.2 1.6 1.6 1.4 221 16 8.2 8.1 8.5 8 7 7 8 7 5.1 8 7.5 7.2 5.6 4.1 4.1 7.6 7.4 6.8 7.2 6.9 6.1 6.3 6.75 5.8 7.3 7.4 3.8 4.8 3.8 17 0.9 0.78 0.9 0.6 0.6 0.8 0.7 0.4 0.75 0.85 0.5 0.9 0.5 0.75 0.8 0.65 0.58 0.45 0.57 0.72 0.42 0.35 0.7 0.55 0.58 0.6 0.7 0.82 0.8 18 6.4 6.5 6.8 6.1 6.12 6.12 5.95 6.1 5.2 5.75 6.3 6.18 5.43 4.85 4.85 8.07 8.23 6.48 6.95 6.9 7.09 5.67 6.1 6.86 7.5 7.01 4.95 4.85 4.75 19 1.51 1.9 1.52 1.83 1.77 1.56 1.83 1.92 1.31 1.43 1.75 1.43 1.23 1.69 1.63 1.77 1.63 1.46 1.54 1.5 1.47 1.43 1.52 1.47 1.62 1.32 1.69 1.74 1.63 Table 3 (Cont’d) punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 12 1.7 1.3 4.3 6.2 4.8 3.7 3.5 4.1 5 4.4 4.4 3.9 3.2 3.8 5.3 7.8 5.3 3.9 6.3 6.5 3.8 4.1 3.7 4.2 3.6 3.4 3.8 4.4 4.2 13 4.2 3.2 8 11 12 9 10 11 11 1.3 11 12 8 8.9 11 9 9 9 13 13.4 6.2 6.3 6.2 7.2 5.4 6.35 5.8 6.1 4.7 14 0.8 0.4 1.5 2.6 1.8 1.5 1.3 1.5 1.8 1.2 1.4 1.6 0.09 1.6 1.9 3.6 1.1 1.4 2.5 2.4 2.2 2.1 1.8 2.1 1.8 1.6 2.1 1.8 1.8 15 1.8 1.3 2.5 2.2 2.6 2.1 1.8 1.75 1.9 2.4 2 1.9 1.7 2.35 2.8 2.7 2.6 1.8 3.4 3.4 1.8 2.2 1.7 2 1.6 1.5 1.4 1.8 2 222 16 4.1 4.2 5.9 9.8 10.4 8.2 6.5 8.6 10 11.5 7.5 9 6.3 9 11 10 6.5 6.9 7.5 7.6 7.5 8.2 6.8 8 6.4 7 6.2 5.5 7.2 17 0.7 0.38 0.8 0.8 0.85 0.95 0.63 0.82 0.7 0.7 0.55 0.82 0.52 1 0.9 0.9 0.8 1.3 0.6 0.63 0.9 1.2 0.9 1.1 1.1 0.8 0.95 0.98 0.6 18 4.9 4.8 7.52 7.34 8.64 7.12 8.31 7.68 8.22 7.87 7.43 7.52 8.41 6.83 8.82 7.32 7.36 6.5 6.4 6.73 5.87 6.87 6.52 6.79 5.13 5.91 6.72 6.53 6.68 19 1.65 1.62 1.95 1.72 1.86 1.72 1.72 1.81 1.23 1.45 1.64 1.86 1.67 1.72 1.84 1.66 2.4 1.35 1.52 1.29 1.14 1.96 1.2 1.36 1.56 1.22 1.72 1.22 1.38 Table 3 (Cont’d) vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 12 5.45 0.5 3.5 3.2 3.4 3.8 3.7 4.2 3.8 3.5 3.6 4.3 5.6 5.4 4.1 4.2 3.4 2.9 4.2 2.45 2.35 2.35 2.6 2.65 2.7 2.8 2.6 2.65 3.2 13 7.8 7.5 6.2 6.5 5.5 6.8 6.6 8.5 6.2 7 7 7.5 8 8 4.5 6 7.1 3.4 4.2 4.5 3.9 6.5 6 4.8 6.5 3.7 5.1 4.5 4.6 14 2.5 2.6 1.9 1.55 1.9 2 1.9 2.1 1.2 2.3 1.7 2 2.8 2.8 2 1.7 1.5 0.51 0.75 0.65 0.6 0.75 0.6 0.98 1.2 0.61 0.6 0.98 0.76 15 2.2 2.3 1.8 1.7 1.7 1.9 1.7 2.4 2.2 1.9 1.75 2.2 2.2 2.5 1.9 1.8 1.8 1.6 2.3 1.7 1.3 1.6 1.82 2.35 1.7 1.6 1.62 2.15 2.2 223 16 5.4 7 6.2 6.5 6 7.5 7.3 8 8.5 6 7 7 8 5 7 5 6.2 7.2 7 9.2 8.6 7.2 7.2 6.9 5.6 8.1 6.9 7 6.8 17 1.2 1.1 0.9 0.7 1 0.95 0.93 1.3 0.85 1.2 1.25 1.4 0.9 1.4 0.4 0.9 1.3 1.2 0.7 0.8 0.52 0.75 0.65 1.1 0.75 1.2 0.75 1.2 0.82 18 5.28 5.74 6.72 5.92 5.23 6.81 6.72 6.49 5.96 5.72 6.43 6.85 5.92 5.24 6.52 6.35 6.58 7.38 7.28 7.04 7.12 6.59 6.82 6.72 6.63 6.56 7.2 6.62 7.45 19 1.58 1.21 1.82 1.21 1.42 1.35 1.32 1.82 1.21 1.48 1.22 1.81 1.22 1.46 1.35 1.32 1.54 1.62 1.55 1.59 1.64 1.62 1.63 1.57 1.58 1.63 1.68 1.59 1.56 Table 3 (Cont’d) Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 12 2.42 2.55 2.64 2.4 3.1 2.75 2.5 2.7 2.6 2.54 2.55 2.2 2.3 13 6.2 5.8 4.2 4.4 4.5 4.4 5.5 4.2 6.4 4.6 5.8 6.2 3.8 14 0.8 0.62 0.87 0.6 0.74 0.85 0.62 0.61 0.83 0.84 0.64 0.72 0.64 15 1.7 1.83 2.15 1.5 2.3 2.14 1.54 1.7 1.8 2.2 1.7 1.5 1.4 224 16 7.3 6.3 6.2 8.5 6.7 6.8 6.5 7.2 5.8 6.9 7.14 1.16 7.6 17 0.74 0.68 1 0.8 0.82 0.9 0.76 0.8 0.78 0.8 0.6 0.72 0.5 18 6.95 6.59 6.48 7.15 7.18 6.98 7.26 7.32 6.74 6.89 7.14 7.26 7.18 19 1.53 1.57 1.62 1.58 1.63 1.64 1.65 1.62 1.63 1.57 1.59 1.58 1.62 Table 3 (Cont’d) ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 20 0.63 0.65 0.89 0.68 0.79 0.53 0.64 0.72 0.78 0.71 0.65 0.75 0.62 0.65 0.69 0.75 0.69 0.63 0.9 0.93 0.87 0.91 0.92 0.9 0.87 1.1 0.84 0.95 0.84 21 1.62 1.58 1.37 1.34 1.21 1.42 1.1 1.17 1.18 1.25 1.1 1.23 1.52 1.26 1.32 1.19 1.23 1.12 1.22 1.21 1.49 1.59 1.66 1.03 1.06 1.08 1.7 1.4 1.18 22 0.37 0.3 0.29 0.25 0.27 0.26 0.25 0.22 0.27 0.29 0.26 0.25 0.32 0.26 0.2 0.26 0.28 0.27 0.29 0.26 0.57 0.42 0.54 0.4 0.37 0.48 0.3 0.46 0.28 23 0.42 0.46 0.9 0.5 0.49 0.3 0.37 0.42 0.45 0.47 0.46 0.46 0.38 0.39 0.41 0.46 0.45 0.45 0 0 0 0 0 0 0 0.8 0 0 0.6 225 24 5.92 6.05 6.12 4.35 4.21 6.05 4.15 5.89 5.82 6.05 4.5 5.9 5.86 4.2 5.91 5.86 5.9 4.8 5.64 7.11 8.45 4.16 3.37 3.68 2.88 6.52 4.61 4.5 4.13 25 3.26 3.42 3.35 3.31 2.94 3.4 3.9 3.15 3.26 3.3 3.1 3.4 3.31 3.85 3.2 3.22 3.31 3.2 3.04 3.7 3.75 2.27 1.57 1.82 2.51 2.31 1.85 2.1 2.96 26 3.18 3.3 3.35 3.26 2.58 3.42 2.67 2.98 3.35 3.3 2.7 3.4 3.12 2.75 2.86 3.25 3.1 2.8 2.58 2 1.84 1.43 1.04 1.37 1.66 1.94 1.19 1.2 1.89 27 1.17 1.09 1.26 1.25 1.29 1.3 1.25 1.26 1.25 1.23 1.31 1.32 1.15 1.1 1.25 1.26 1.21 1.29 1.28 1.09 1.22 0.87 0.68 0.82 0.88 0.98 0.9 0.8 1.07 Table 3 (Cont’d) Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 20 0.72 0.7 0.6 0.76 0.69 0.65 1.26 0.56 0.67 0.61 0.61 0.71 0.52 0.55 0.67 0.69 0.57 0.78 0.45 0.67 0.5 0.78 0.56 0.43 0.73 0.54 0.43 0.47 0.5 21 1.23 1.29 1.9 1.75 1.15 1.91 1.58 1.17 1.14 1.2 1.23 1.2 2.15 1.37 1.37 2.01 1.72 1.38 1.29 1.48 1.23 2.16 1.28 2.1 1.03 1.36 1.42 1.39 1.28 22 0.22 0.38 0.27 0.36 0.39 0.39 0.4 0.39 0.28 0.3 0.31 0.25 0.31 0.44 0.44 0.29 0.37 0.26 0.25 0.24 0.23 0.16 0.14 0.2 0.23 0.26 0.44 0.45 0.25 23 0.41 0.56 0.53 0.49 0.61 0.87 1.13 0.57 0.51 0.53 0.52 0.57 0.61 0.55 0.52 0.48 0.47 0.58 0.61 0.57 0.55 0.49 0.59 0.57 0.51 0.48 0.56 0.54 0.62 226 24 4.32 4.25 3.62 3.32 3.18 4.46 5.4 4.82 3.68 3.72 4.32 4.05 3.84 2.76 2.76 3.03 2.97 3.47 3.01 2.4 2.38 3.79 4.82 3.84 4.17 3.24 2.75 3.92 3.67 25 2.78 2.3 2.01 2.54 2.64 2.64 2.85 2.52 2.32 2.19 2.5 2.46 2.05 1.75 1.75 1.78 2.23 2.42 2.23 1.85 1.92 1.84 2.35 2.29 2.09 2.07 1.82 2.04 1.98 26 1.4 1.59 1.61 1.56 1.95 2.37 2.34 2.32 1.4 2.18 1.69 1.5 1.84 1.76 1.76 1.86 1.44 1.93 2.05 1.41 1.3 1.47 2.52 2.38 1.36 1.48 1.33 1.85 1.6 27 1.31 0.83 1.09 1.16 1.15 1.49 1.33 1.24 0.8 1.1 1.1 1.1 0.82 0.99 0.99 1.47 1.02 1.15 1.03 0.91 0.8 0.68 1.33 1.11 0.81 0.73 0.64 1.2 0.88 Table 3 (Cont’d) Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 20 0.39 0.31 0.33 0.64 0.36 0.51 0.45 0.4 0.46 0.51 0.45 0.42 0.53 0.34 0.46 0.58 0 0 0.63 0.52 0.62 0 0 0.55 0.62 0.71 0.63 0 0 21 1.06 0.39 0.61 0.71 0.81 0.59 0.72 0.64 0.72 0.66 0.71 0.81 0.68 0.82 0.76 0.74 1.37 1.21 0.77 1.16 1.16 1.01 1.24 1.63 1.06 1.29 1.35 1.25 1.23 22 0.34 0.24 0.57 0.47 0.44 0.38 0.4 0.48 0.48 0.54 0.52 0.5 0.46 0.52 0.39 0.42 0.4 0.32 0.26 0.45 0.26 0.24 0.39 0.3 0.23 0.2 0.33 0.25 0.24 23 0.4 0.42 0.41 0.43 0.41 0.41 0.42 0.41 0.41 0.42 0.43 0.44 0.41 0.41 0.42 0.41 0.85 0.71 0.68 0.69 0.75 0.68 0.66 0.65 0.64 0.96 1.1 0.46 0.48 227 24 3.23 2.08 2.59 3.25 2.88 2.86 3.74 2.78 2.84 3.77 3.78 2.85 2.9 3.1 3.63 3.7 6.96 6.61 5.8 6.65 4.8 7.08 6.72 6.72 6.38 6.59 6.98 6.61 6.42 25 1.6 1.44 1.92 2.41 1.9 1.76 2.4 2.6 2.89 3.21 3.13 2.75 2.84 3.12 3.11 3.13 2.63 2.43 2.42 2.5 2.46 2.38 2.18 2.69 2.51 2.31 2.8 2.42 2.53 26 1.27 1.2 0.82 1.82 2.1 1.91 3.42 1.5 1.7 2.12 1.5 1.42 1.5 1.63 1.38 1.49 2.63 2.23 1.43 1.88 1.57 2.14 1.47 1.92 1.76 2.88 2.1 2.73 1.82 27 0.71 1.1 0.81 2.19 1.25 1.36 1.16 1.12 1.15 1.27 1.3 1.02 1.11 1.19 1.21 1.06 1.04 0.85 0.76 0.81 1.01 1.1 0.86 0.85 1.17 0.83 0.98 0.84 1.12 Table 3 (Cont’d) 20 lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 0 0.59 0.58 0.64 0.68 0.71 0 0 0 0 0.58 0.71 0.63 0.62 0 0.7 0.67 0.69 0.68 0.67 0.63 0.58 0.65 0.58 0 0 0.64 0 0.63 21 1.44 1.36 1.14 1.63 1.59 1.19 1.65 1.32 1.33 1.31 1.24 1.55 1.33 1.29 1.36 1.41 1.36 1.28 1.36 1.34 1.32 1.31 1.3 1.61 1.3 1.29 1.42 1.3 1.32 22 0.32 0.31 0.31 0.47 0.45 0.28 0.36 0.37 0.32 0.31 0.31 0.37 0.4 0.35 0.33 0.29 0.31 0.29 0.33 0.34 0.33 0.32 0.33 0.42 0.31 0.32 0.3 0.29 0.31 23 0.71 0.68 0.65 0.61 0.71 0.6 0.73 0.64 0.69 0.63 0.65 0.7 0.75 0.74 0.67 0.74 0.6 0.63 0.68 0.74 0.78 0.75 0.7 0.63 0.65 0.81 0.68 0.71 0.65 228 24 6.12 7.2 5.8 5.73 8.36 6.35 4.18 6.11 4.66 6.42 6.23 4.42 6.05 5.23 6.08 8.02 6.3 3.33 6.5 7.9 6.9 5.02 6.5 8.49 4.95 5.24 5.44 4.54 5.75 25 2.21 3.21 2.9 2.99 3.92 3.38 2.14 2.96 3.12 2.57 3.24 2.38 3.1 3.05 2.62 4.32 3.56 1.86 3.5 3.24 3.1 2.54 3.35 4.29 2.34 3.1 2.48 2.54 2.6 26 2.8 2.77 1.3 3.56 2.89 1.79 1.71 2.23 1.7 2.84 1.82 2.15 2.19 1.92 2.63 2.8 1.5 1.7 1.7 2.52 2.46 1.54 1.3 3.05 1.54 2.1 1.3 1.13 1.49 27 1.1 1.2 1.01 0.81 1.65 0.85 0.81 1.07 0.96 0.92 0.87 0.92 1.05 0.95 0.96 0.96 1.2 0.84 1.13 1.26 0.98 0.76 1.1 1.68 0.92 1.1 0.85 0.68 0.86 Table 3 (Cont’d) lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 20 0.68 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 21 1.33 1.31 1.24 1.69 2.32 0.76 1.8 1.24 1.62 1.23 1.13 1.2 0.9 1.29 1.6 0.93 0.7 0.92 0.75 1.09 0.66 0.74 0.61 0.8 0.76 0.93 0.82 0.96 0.69 22 0.33 0.42 0.45 0.4 0.41 0.43 0.39 0.41 0.4 0.42 0.39 0.31 0.41 0.4 0.28 0.27 0.27 0.51 0.39 0.34 0.31 0.5 0.34 0.34 0.38 0.42 0.31 0.33 0.34 23 0.72 0 0 0 1.2 1.15 0 0 0 0 0 0.7 0 0 0 0.61 0.61 0.55 0.57 0.56 0.55 0.58 0.56 0.54 0.55 0.58 0.59 0.57 0.57 229 24 6.78 4.86 3.53 6.3 3.77 4.21 4.12 4.52 3.64 4.61 4.57 3.89 4.21 4.72 6.3 4.8 5.65 5.5 4.6 4.2 6.52 4.3 4.3 4.2 4.8 5.6 5.5 4.3 5.1 25 3.52 3.01 2.98 2.98 2.95 2.87 3.01 2.94 2.92 2.89 3.14 3.12 2.85 2.95 3.02 0.84 1.91 1.93 2.3 2.4 2.18 1.58 2.86 2.2 2.1 2.1 2.07 2.32 2.19 26 1.26 1.85 1.5 2.05 1.48 1.54 1.52 1.64 1.45 1.56 1.4 1.6 1.65 1.76 2.12 1.24 1.51 1.3 1.15 1.26 2 1.15 1.32 1.28 1.23 1.29 1.36 1.25 1.3 27 1.13 1.03 0.79 1.13 0.74 0.81 0.82 0.86 0.76 0.87 0.85 0.8 0.83 0.98 1.16 0.59 0.59 0.54 0.61 0.71 1.07 0.49 0.78 0.73 0.79 0.61 0.62 0.69 0.65 Table 3 (Cont’d) 20 occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 0 0 0 0 0 0 0 0 0 0 0 0.52 0 0.66 0.59 0.49 0.57 0.62 0.58 0.4 0.46 0.52 0.49 0.56 0.52 0.63 0.67 0.63 0.58 21 0.85 0.86 0.92 0.72 1.05 0.82 0.98 0.72 0.92 0.76 0.96 1.58 0.66 1.1 0.98 1.14 1.22 1.29 0.96 1.25 1.14 1.41 1.35 1.2 1.08 0.85 1.09 0.96 1.08 22 0.35 0.48 0.39 0.3 0.31 0.32 0.33 0.29 0.31 0.35 0.32 0.25 0.16 0.22 0.35 0.44 0.31 0.28 0.3 0.22 0.27 0.24 0.26 0.3 0.28 0.25 0.2 0.19 0.2 23 0.59 0.55 0.61 0.64 0.67 0.55 0.58 0.61 0.62 0.58 0.62 0 0 0 0 0 0.42 0 0 0 0 0.3 0 0 0 0 0.98 0.95 0.97 230 24 5.62 5.6 5.8 5.8 4.5 4.1 4.3 5.45 4.2 4.8 4.6 4.89 6.3 5.4 5.6 10.24 8.65 7.4 7.47 6.23 5.64 7.52 7.32 6.1 9.5 5.96 5.2 4.8 5.4 25 2.2 2 2.3 2.2 2.35 2.19 2.26 2.32 2.2 2.15 2.19 2.42 2.28 2.14 2.3 2.98 2.65 2.96 2.62 3.24 2.74 2.36 2.29 2.65 2.81 2.68 2.1 2 2.08 26 1.4 1.35 1.42 1.38 1.2 1.27 1.22 1.67 1.42 1.28 1.32 2.24 2.24 2 1.81 2.24 2.81 2.27 2.42 1.28 1.85 2.22 2.15 2.04 2.1 2.06 1.9 1.8 1.9 27 0.78 0.58 0.84 1.1 0.82 0.71 0.72 0.74 0.75 0.69 0.84 1.2 1.2 0.85 0.8 1.44 1.55 0.72 1.22 1.08 1.3 1.35 1.26 1.25 1.2 1.14 0.82 0.73 0.78 Table 3 (Cont’d) punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 20 0.56 0.51 0.78 0.81 0.79 0.82 0.81 0.83 0.84 0.82 0.85 0.75 0.68 0.93 0.82 0.89 0.76 0.5 0.5 0.48 0.48 0.51 0.51 0.48 0.52 0.47 0.53 0.47 0.49 21 1.06 1.25 1.55 1.51 1.41 1.56 1.42 1.31 1.14 1.27 1.47 1.27 1.41 1.56 1.35 1.53 1.69 1.2 1.14 1.12 0.91 1.19 1.04 1.26 1.16 0.94 1.12 1.1 1.14 22 0.21 0.19 0.45 0.34 0.31 0.37 0.35 0.41 0.39 0.38 0.38 0.4 0.31 0.36 0.28 0.37 0.54 0.27 0.2 0.22 0.2 0.27 0.25 0.25 0.25 0.21 0.26 0.25 0.24 23 0.95 0.96 0.71 0.69 0.68 0.73 0.71 0.69 0.72 0.71 0.68 0.71 0.65 0.62 0.84 0.6 0.73 0.38 0 0.41 0.48 0.52 0.45 0.46 0.42 0.38 0.41 0.42 0.48 231 24 5.3 5.5 3.1 3.15 3.2 3.15 3.1 3.12 3.39 3.3 2.96 3.12 2.9 3.11 3.25 3.16 3.11 7.24 6.5 5.58 5.84 6.93 7.81 6.4 4.97 5.64 7.1 7.23 5.9 25 2.2 2.12 1.68 1.59 1.74 1.52 1.49 1.35 1.8 1.69 1.72 1.25 1.54 1.53 1.86 1.62 1.72 2.8 2.67 2.75 2.76 2.67 2.82 2.31 2.24 2.81 2.72 2.91 2.38 26 2.1 2.05 2.1 2.12 2.22 1.95 1.9 2.1 2.35 2.11 2.15 1.96 1.8 1.85 2.28 2.16 1.88 2.55 2.95 1.74 2.31 2.4 3.38 2.46 2.36 2.29 2.31 3.14 2.52 27 0.8 0.8 0.84 0.81 0.78 0.75 0.73 0.75 0.64 0.73 0.68 0.74 0.75 0.76 0.79 0.8 0.85 1.56 2.19 1.04 0.89 1.26 1.33 1.22 1.1 1.14 1.28 1.27 1.19 Table 3 (Cont’d) vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 20 0.48 0.53 0.51 0.48 0.44 0.54 0.49 0.54 0 0.52 0.48 0.49 0.51 0.52 0.48 0.51 0.46 0.39 0.4 0.43 0.41 0.39 0.4 0.4 0.42 0.4 0.39 0.4 0.39 21 1.15 0.98 1.13 1.05 1.15 1.24 1.08 1.2 1.17 1.15 1.09 1.18 1.08 1.12 1.22 1.14 1.16 1.15 1.32 1.35 1.19 1.42 1.22 1.18 1.16 1.23 1.42 1.1 1.22 22 0.23 0.21 0.25 0.22 0.25 0.24 0.26 0.25 0.22 0.25 0.25 0.26 0.21 0.27 0.26 0.24 0.25 0.39 0.4 0.41 0.38 0.39 0.42 0.39 0.41 0.42 0.41 0.43 0.44 23 0.47 0.42 0.45 0.42 0.51 0.62 0.37 0 0.29 0.51 0.38 0.38 0.37 0.38 0.39 0.51 0.38 0.63 0.71 0.64 0.59 0.63 0.61 0.58 0.53 0.52 0.7 0.61 0.58 232 24 5.2 5.56 6.82 6.12 5.22 6.15 7.62 6.82 6.22 5.12 7.72 7.1 6.45 5.12 6.35 7.42 7.1 2.9 3.4 2.87 5.72 3.19 3.6 3.3 3.51 5.89 3.4 3.34 5.79 25 2.34 2.71 2.64 2.52 2.31 2.43 2.72 2.62 2.5 2.32 2.67 2.59 2.56 2.31 2.43 2.72 2.71 1.85 1.96 1.87 2.07 2.1 1.94 1.85 2.25 2.74 1.95 1.78 2.85 26 2.33 2.42 2.39 2.32 2.41 2.39 2.24 2.38 2.31 2.35 2.82 2.52 2.42 2.46 2.41 3.05 2.58 1.5 2.82 1.75 1.95 1.8 1.76 1.78 2.57 1.8 2.63 1.87 1.94 27 1.12 1.08 1.23 0.92 1.15 1.23 1.28 1.15 0.91 1.15 1.36 1.27 0.95 1.17 1.26 1.34 1.22 1.15 1.15 1.11 1.14 1.12 1.24 0.98 1.34 1.2 1.19 1.19 1.41 Table 3 (Cont’d) Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 20 0.42 0.4 0.41 0.38 0.4 0.37 0.4 0.41 0.4 0.39 0.39 0.38 0.41 21 1.18 1.55 1.55 1.26 1.18 1.19 1.36 1.47 1.5 1.39 1.27 1.19 1.18 22 0.42 0.39 0.41 0.38 0.42 0.43 0.44 0.39 0.38 0.44 0.42 0.41 0.39 23 0.62 0.5 0.72 0.64 0.63 0.62 0.65 0.63 0.6 0.68 0.71 0.59 0.65 233 24 3.49 3.43 3.5 3.12 3.5 3.32 3.47 3.64 3.24 3.62 3.48 3.5 4.1 25 1.82 1.8 1.75 1.83 2.36 1.95 2.75 1.9 1.92 2.25 2.1 2.86 2.12 26 1.81 2.16 2 1.43 1.65 1.75 1.96 2.52 1.56 1.74 1.83 2.05 2.1 27 1.64 1.28 0.95 1.04 1.09 1.23 1.27 1.36 1.1 1.14 1.32 1.48 1.42 Table 3 (Cont’d) ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 28 1.02 1.32 1.02 1.04 1.02 1.03 1.22 1.15 1.09 1.1 1.19 1.1 1.1 1.2 1.17 1.1 1.12 1.21 0.97 0.92 0.65 0.55 0.61 0.68 0.7 1.04 0.92 0.91 1.19 29 0.76 1.01 0.87 0.8 0.76 0.88 0.96 0.99 0.96 0.91 0.85 0.95 0.75 0.92 0.95 0.94 0.89 0.86 0.75 0.75 0.99 0.58 0.57 0.52 0.82 0.75 0.75 0.6 0.85 30 10.87 11.18 11.22 11.23 8.24 10.87 8.12 8.25 9.52 10.18 7.62 9.48 10.7 8.5 8.35 9.51 9.8 7.8 7.95 7.08 9.31 8.5 6.59 4.42 3.45 5.8 5.4 5.7 6.76 31 4.05 4.23 4.09 4.05 3.84 4.12 3.85 3.76 4.05 4.15 3.82 4.08 4.02 4 3.9 4.01 4.2 3.81 3.6 4.15 4.89 3.44 2.7 2.58 2.04 2.8 2.24 2.43 2.81 234 32 0.41 0.39 0.41 0.39 0.14 0.38 0.31 0.33 0.41 0.42 0.28 0.39 0.4 0.35 0.34 0.39 0.41 0.31 0.12 0.11 0.11 0.21 0.14 0.18 0.18 0.17 0.14 0.15 0.25 33 0.22 0.23 0.27 0.26 0.21 0.23 0.24 0.25 0.24 0.26 0.25 0.2 0.23 0.25 0.26 0.25 0.27 0.27 0.26 0.24 0.25 0.23 0.23 0.22 0.19 0.21 0.16 0.2 0.27 34 0.65 0.66 0.65 0.65 0.69 0.66 0.69 0.7 0.65 0.66 0.64 0.22 0.66 0.67 0.69 0.66 0.65 0.63 1.1 1.1 1.1 1.2 0.4 1.2 1.2 1.1 1.1 1.2 0.99 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 28 1.49 1.49 1.39 1.37 1.17 1.67 1.36 1.38 1.23 1.2 1.39 0.98 1.02 0.8 0.8 0.77 1.35 0.94 1.17 1.03 0.78 0.88 1.21 1.25 0.43 0.83 0.83 0.53 0.89 29 0.98 0.72 0.76 0.84 1.25 1.57 1.51 1.05 0.82 0.6 0.85 0.84 0.62 1.2 1.2 0.96 0.89 0.99 0.86 0.66 0.48 0.6 0.69 0.66 0.68 0.63 0.47 0.76 0.79 30 7.1 6.8 11.92 7.29 5.85 7.1 7.07 6.8 6 7.25 7.1 6.58 7.2 6.72 6.75 6.27 6.98 7.01 7.09 6.61 5.72 5.33 6.72 7.37 6.47 6.81 5.78 5.7 5.79 31 3.8 3.3 3.01 2.04 2.24 3.65 4.15 3.8 3.2 2.8 3.45 2.72 2.85 3.15 3.2 2.7 2.52 3.52 3.94 3.23 2.42 2.65 3.46 2.92 3.19 3.21 2.51 2.07 2.48 235 32 0.25 0.26 0.46 0.28 0.6 0.2 0.18 0.18 0.13 0.13 0.2 0.2 0.14 0.16 0.16 0.32 0.63 0.14 0.37 0.18 0.17 0.12 0.16 0.14 0.25 0.1 0.08 0.11 0.08 33 0.28 0.25 0.36 0.39 0.8 0.51 0.52 0.49 0.24 0.12 0.26 0.27 0.15 0.27 0.27 0.37 0.55 0.08 0.33 0.11 0.08 0.16 0.05 0.09 0.2 0.05 0.12 0.26 0.18 34 0.57 0.77 0.33 0.77 0.88 1.1 0.56 0.39 0.28 0.45 0.62 0.99 0.5 0.62 0.62 0.72 0.55 0.81 0.7 0.9 0.85 0.8 0.52 0.39 0.73 0.4 0.2 0.52 0.49 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 28 0.47 0.82 0.43 1.5 1.3 1.12 1.01 1.2 0.82 0.97 0.88 0.86 0.96 1.05 1.19 1.3 1.2 1.97 1.01 1.32 1.2 0.78 0.7 1.14 1.24 1.09 1.25 1.1 1.22 29 0.61 0.87 0.48 1.46 0.98 0.91 0.61 0.89 1.9 0.87 0.79 0.77 0.9 0.86 0.97 0.72 0.8 0.75 0.64 0.45 0.64 0.8 0.8 0.7 0.76 0.52 0.89 0.59 0.72 30 6.5 4.18 4.86 5.67 4.8 4.95 5.24 4.88 5.3 4.62 5.1 4.12 4.65 4.98 5.12 5.23 7.5 7.04 7.12 5.98 6.83 6.92 7.98 5.75 6.95 8.12 7.2 7.98 7.12 31 2.55 2.22 2.51 2.75 2.5 2.84 2.68 2.99 2.8 2.73 2.89 2.48 2.56 2.97 2.86 2.81 3.25 3.9 3.2 2.4 3.2 2.25 3.12 2.2 3.3 3.17 2.75 3.38 3.24 236 32 0.13 0.18 0.14 0.11 0.12 0.22 0.22 0.18 0.11 0.12 0.14 0.09 0.08 0.1 0.13 0.14 0.18 0.21 0.22 0.21 0.22 0.19 0.18 0.22 0.22 0.19 0.21 0.21 0.21 33 0.21 0.19 0.21 0.25 0.22 0.26 0.28 0.18 0.17 0.15 0.21 0.12 0.11 0.18 0.21 0.19 0.21 0.24 0.23 0.21 0.24 0.21 0.2 0.2 0.24 0.21 0.24 0.22 0.24 34 0.47 0.35 0.4 0.41 0.39 0.19 0.2 0.38 0.36 0.4 0.41 0.42 0.41 0.42 0.4 0.42 0.98 1 0.82 0.78 1.01 1.08 1.53 0.8 1.3 0.92 0.93 0.94 1.2 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 28 1.12 1.27 0.8 1.3 1.39 1.45 1.21 1.16 0.71 1.25 1.25 1.31 1.22 0.86 1.12 1.18 0.96 0.88 1.39 1.46 0.77 0.88 0.81 1.7 1.09 0.85 0.55 0.66 0.76 29 0.95 0.91 0.62 0.87 1.06 0.85 0.84 0.62 0.77 0.86 0.86 0.83 0.72 0.81 0.85 0.96 0.88 0.56 0.92 0.94 0.82 0.57 0.91 1.23 0.85 0.84 0.76 0.53 0.87 30 7.56 10.12 7.5 10.22 10.35 7.61 7.13 8.09 5.66 7.68 8.12 7.43 7.95 6.2 7.75 8.32 7.7 6.23 7.85 8.35 7.59 7.61 7.8 9.95 7.98 7.65 8.21 7.48 8.32 31 3.28 4.6 3.2 3.74 4.14 4.87 4.78 4.04 3.34 3.41 4.82 4.68 4.09 3.32 3.46 3.7 3.8 2.44 4 2.85 3.28 3.3 4.02 5.05 3.94 3.52 4.01 3.62 3.89 237 32 0.3 0.31 0.28 0.3 0.37 0 0.25 0.3 0.29 0.35 0.26 0.26 0.29 0.31 0.34 0.25 0.31 0.3 0.31 0.27 0.21 0.24 0.28 0.28 0.31 0.3 0.34 0.28 0.27 33 0.45 0.38 0.48 0.45 0.56 0 0.54 0.47 0.5 0.43 0.45 0.54 0.48 0.51 0.42 0.51 0.47 0.2 0.5 0.58 0.39 0.52 0.46 0.53 0.39 0.43 0.51 0.5 0.52 34 1.05 1.09 1.12 1.45 1.05 1.13 1.04 0.98 1 1.05 1.12 1.1 1.14 0.98 0.99 1.11 1.16 1.15 1.14 1.12 1.13 1.05 1.13 1.06 1.06 1.12 1.03 1.05 1.11 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 28 0.83 0.85 0.76 0.96 0.72 0.78 0.75 0.82 0.69 0.8 0.72 0.75 0.81 0.79 0.94 0.69 0.38 0.4 0.62 0.69 0.85 0.45 0.74 0.76 0.71 0.52 0.55 0.72 0.68 29 0.93 0.72 0.66 0.79 0.6 0.68 0.68 0.65 0.7 0.62 0.67 0.68 0.69 0.72 0.86 0.58 0.52 0.53 0.58 0.57 0.64 0.51 0.54 0.58 0.55 0.55 0.56 0.58 0.56 30 7.74 7.6 6.85 8.54 6.73 6.9 6.92 6.93 6.76 6.76 6.72 6.92 7.01 7.4 8.78 3.6 4.64 7.2 5.5 6 6.3 5.7 7.92 4.2 5.5 6.3 5.5 5.85 5.5 31 3.81 4.2 3.94 4.21 3.89 3.89 4.12 3.97 4.02 3.94 3.85 3.76 3.79 3.95 4.03 1.3 1.6 2.2 2.5 2.26 3.96 2.45 4.5 2.8 2.2 2.25 1.98 2.3 2.2 238 32 0.31 0.19 0.11 0.19 0.12 1.13 0.1 0.13 0.99 0.11 0.11 0.14 0.12 0.16 0.22 0.14 0.13 0.14 0.14 0.15 0.12 0.13 0.14 0.13 0.13 0.13 0.14 0.15 0.14 33 0.45 0.28 0.14 0.35 0.13 0.17 0.15 0.17 0.2 0.14 0.13 0.12 0.14 0.27 0.39 0.21 0.18 0.2 0.21 0.21 0.17 0.16 0.14 0.2 0.18 0.19 0.19 0.21 0.18 34 1.15 0.32 0.15 0.37 0.17 0.2 0.24 0.26 0.19 0.35 0.26 0.16 0.21 0.21 0.38 0.55 0.6 0.61 0.34 0.59 0.52 0.59 0.98 0.49 0.52 0.54 0.52 0.53 0.49 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 28 0.73 0.55 0.78 0.72 0.68 0.74 0.72 0.81 0.73 0.74 0.66 1.06 0.62 0.96 0.91 0.72 1.18 0.92 0.72 0.85 0.66 0.59 0.79 0.78 0.68 0.84 0.95 0.96 0.89 29 0.58 0.53 0.6 0.63 0.61 0.61 0.58 0.63 0.53 0.54 0.58 0.73 0.8 0.67 0.67 0.74 1.03 0.69 0.71 0.78 0.68 0.7 0.69 0.69 0.72 0.86 0.64 0.68 0.67 30 6.2 6.5 6.5 6.2 6.5 3.89 5.8 5.7 4.1 5.7 6.2 7.3 6.2 8.2 7.6 8.4 9.2 7.58 10.25 7.85 6.32 7.01 7.54 6.43 9.4 8.86 7.75 7.15 7.62 31 2.4 2.3 2.8 3.19 2.56 1.03 2.3 3.3 1.8 2.42 2.36 3.46 2.66 3.45 4.1 3.3 4.6 3.2 4.26 4.1 3.06 2.28 3.14 3.12 3.45 3.89 3.5 3.26 3.1 239 32 0.14 0.14 0.15 0.13 0.14 0.14 0.14 0.14 0.13 0.13 0.14 0.22 0.18 0.2 0.22 0.19 0.24 0.21 0.17 0.28 0.24 0.21 0.24 0.24 0.19 0.23 0.22 0.25 0.19 33 0.2 0.18 0.2 0.17 0.19 0.22 0.2 0.18 0.2 0.21 0.2 0.29 0.24 0.27 0.3 0.13 0.35 0.24 0.21 0.44 0.31 0.38 0.35 0.32 0.19 0.31 0.26 0.28 0.28 34 0.5 0.53 0.55 0.49 0.6 0.54 0.55 0.5 0.51 0.42 0.55 1.12 0.88 0.99 1.1 1.17 1.19 0.85 1.1 1.14 0.99 1.66 1.42 1.12 1.1 1.16 1.1 1.1 1.16 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 28 1.2 0.91 1.05 1.15 1.19 0.72 0.88 1.2 1.27 0.96 0.82 1.34 0.6 0.66 1.18 1.31 1.19 1.22 1.1 1.1 0.95 1.07 1.13 1.16 1.03 1.02 1.04 1.12 1.15 29 0.7 0.63 0.82 1.1 0.82 0.61 0.75 0.72 0.7 0.8 0.72 0.73 0.71 0.58 0.76 1.26 0.55 0.9 0.72 0.73 0.56 0.63 0.83 0.77 0.79 0.59 0.65 0.82 0.76 30 7.15 6.8 7.13 7.41 7.55 7.38 7.41 7.19 7.63 7.56 7.89 7.22 7.45 7.3 7.56 7.43 7.12 7.9 8.1 8.3 6.91 8.1 6.8 7.9 6.21 7.2 7.85 7.72 6.53 31 3.3 3.2 3.12 3.49 3.24 3.58 3.51 3.61 3.26 3.32 3.95 3.57 3.48 3.54 3.18 3.52 2.95 2.1 3.89 3.85 3.55 3.9 3.35 3.2 2.72 3.4 3.8 3.45 3.3 240 32 0.21 0.26 0.09 0.085 0.09 0.09 0.08 0.09 0.09 0.09 0.09 0.09 0.085 0.09 0.09 0.08 0.09 0.2 0.21 0.22 0.11 0.27 0.19 0.21 0.24 0.18 0.25 0.21 0.22 33 0.29 0.31 0.12 0.13 0.13 0.12 0.12 0.12 0.12 0.13 0.13 0.13 0.11 0.13 0.14 0.12 0.13 0.32 3.1 0.31 0.18 0.54 0.3 0.24 0.26 0.22 0.45 0.32 0.26 34 1.12 1.21 1.14 1.1 1.1 1.18 1.1 1.12 1.54 1.45 1.43 1.08 1.05 1.21 0.98 0.89 1.11 1.4 1.2 1.14 1.12 1.17 1.15 1.17 1.15 1.19 1.27 1.36 1.12 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 28 1.1 1.02 1.05 1.01 0.98 1.12 1.13 1.06 1.01 1 1.11 1.08 0.98 1.02 1.13 1.12 1.09 1.2 1.55 1.22 1.77 1.7 1.05 0.69 1.6 1.65 1.77 1.06 1.27 29 0.75 0.58 0.68 0.55 0.77 0.78 0.82 0.62 0.58 0.76 0.84 0.61 0.62 0.81 0.75 0.82 0.64 1.04 1.36 0.95 1.14 0.78 0.81 0.82 1.16 0.96 1.16 0.78 1.16 30 6.42 7.28 7.86 6.8 6.62 7.5 7.3 8.2 7.4 6.81 8.67 7.98 7.2 7.34 6.45 8.3 7.62 8.5 9.24 6.85 7.56 7.73 7.45 7.85 8.65 9.6 7.54 7.82 9.54 31 2.9 3.6 3.75 3.61 2.92 3.1 3.28 3.86 3.45 2.7 2.2 3.8 3.45 2.89 3.15 3.25 3.85 3.6 3.98 3.24 4.1 3.72 3.58 3.49 3.86 4.2 3.65 3.71 4.13 241 32 0.23 0.19 0.26 0.21 0.25 0.24 0.22 0.26 0.19 0.25 0.22 0.28 0.19 0.25 0.22 0.23 0.25 0.25 0.28 0.26 0.27 0.34 0.29 0.3 0.31 0.28 0.26 0.38 0.33 33 0.28 0.22 0.48 0.27 0.28 0.25 0.32 0.41 0.33 0.32 0.31 0.48 0.29 0.31 0.32 0.33 0.48 0.31 0.33 0.31 0.33 0.34 0.3 0.33 0.32 0.31 0.28 0.32 0.31 34 1.18 1.19 1.23 1.15 1.18 1.16 1.15 1.16 1.13 1.23 1.36 1.28 1.16 1.17 1.16 1.32 1.21 0.86 0.89 0.91 0.95 0.97 0.92 1.1 0.98 0.85 0.88 1 0.95 35 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 28 1.36 1.3 1.27 0.93 1.76 1.24 1.62 1.75 1.12 1.64 1.32 1.58 1.62 29 1.23 1.21 1.18 0.99 0.91 1.1 1.13 0.86 0.98 0.95 0.99 1.05 0.92 30 7.56 7.72 6.52 6.52 7.86 7.43 8.41 7.51 6.6 7.7 7.52 8.35 7.74 31 3.61 3.6 3.52 2.81 3.63 3.49 4.24 3.58 3.2 3.72 3.55 4.3 3.62 242 32 0.25 0.22 0.25 0.26 0.27 0.29 0.32 0.26 0.25 0.26 0.28 0.29 0.3 33 0.3 0.28 0.31 0.32 0.33 0.31 0.3 0.29 0.31 0.32 0.32 0.31 0.28 34 0.98 0.76 0.92 0.94 0.96 0.88 0.91 0.95 0.95 0.95 0.91 0.9 0.94 35 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 37 1 0 1 0 0 0 0 1 0 1 0 0 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 0 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 1 1 1 1 1 1 0 1 0 1 0 0 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 243 41 1 0 1 1 1 0 0 1 0 1 0 0 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 244 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 245 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 246 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 37 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 247 41 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 37 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 248 41 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 0 0 0 0 1 0 0 0 1 1 1 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 1 1 0 0 1 1 1 1 0 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 37 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 249 41 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 37 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 39 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 250 41 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 36 Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 37 0 0 0 0 0 0 0 0 0 0 0 0 0 38 1 1 1 1 1 1 1 1 1 1 1 1 1 39 0 0 0 0 0 0 0 0 0 0 0 0 0 40 0 0 0 1 1 1 1 1 1 1 1 1 1 251 41 0 0 0 0 0 0 0 0 0 0 0 0 0 42 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 44 ark76 ark77 ark81 ark83 ark84 ark85 ark287 ark288 ark289 ark290 ark291 ark408 ark409 ark410 ark411 ark413 ark414 ark415 corr460 corr461 corr462 corr463 corr464 corr465 corr466 corr468 corr470 corr471 Frut236 45 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 252 49 1 1 1 1 1 0 1 1 1 1 0 1 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 50 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 Frut237 Frut238 Frut239 Frut240 Frut241 Frut242 Frut243 Frut244 Frut245 Frut260 Frut261 Frut262 Frut263 Frut264 Frut46 Frut47 Frut48 Frut49 Frut50 Frut51 Frut52 Frut53 Frut54 Frut55 Frut56 Frut57 Frut58 Frut59 Frut60 45 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 253 49 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 50 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 Hum16 Hum17 Hum18 Hum19 Hum20 Hum21 Hum22 Hum23 Hum24 Hum25 Hum26 Hum27 Hum28 Hum29 Hum30 Hum31 inter106 inter107 inter108 inter110 inter111 inter112 inter113 inter116 inter118 inter119 inter120 inter225 inter226 45 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 0 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 254 49 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 50 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 lasio250 lasio251 lasio252 lasio253 lasio254 lasio255 lasio256 lasio257 lasio258 lasio259 lasio292 lasio293 lasio294 lasio295 lasio296 lasio61 lasio62 lasio63 lasio64 lasio65 lasio66 lasio67 lasio68 lasio69 lasio70 lasio71 lasio72 lasio73 lasio74 45 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 1 1 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 255 49 0 1 1 0 0 0 1 0 0 0 0 0 0 0 0 1 1 0 1 0 0 1 1 1 0 0 0 0 0 50 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 lasio75 Marit121 Marit122 Marit123 Marit124 Marit125 Marit126 Marit126 Marit127 Marit128 Marit129 Marit130 Marit131 Marit132 Marit133 occ1 occ10 occ11 occ12 occ13 occ14 occ15 occ2 occ3 occ309 occ310 occ311 occ312 occ313 45 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 46 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 256 49 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 50 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 occ314 occ315 occ316 occ317 occ318 occ4 occ5 occ6 occ7 occ8 occ9 punc151 punc152 punc154 punc155 punc158 punc159 punc160 punc161 punc163 punc164 punc165 punc246 punc247 punc248 punc249 punc301 punc302 punc303 45 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 46 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 1 1 1 48 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 257 49 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 0 0 1 1 0 1 0 1 1 1 50 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 punc304 punc305 Stan271 Stan272 Stan273 Stan274 Stan275 Stan276 Stan91 Stan92 Stan93 Stan94 Stan95 Stan96 Stan97 Stan98 Stan99 vill156 vill157 vill162 vill207 vill208 vill209 vill210 vill211 vill212 vill213 vill214 vill215 45 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 258 49 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 50 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 Table 3 (Cont’d) 44 vill216 vill217 vill218 vill31 vill32 vill33 vill34 vill35 vill36 vill37 vill39 vill40 vill41 vill42 vill43 vill44 vill45 Virid136 Virid137 Virid138 Virid139 Virid140 Virid141 Virid142 Virid143 Virid144 Virid145 Virid146 Virid147 45 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 0 1 1 1 1 1 1 1 1 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 259 49 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 50 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 Table 3 (Cont’d) 44 Virid148 Virid149 Virid150 Virid219 Virid220 Virid221 Virid222 Virid223 Virid282 Virid283 Virid284 Virid285 Virid286 45 1 1 1 1 1 1 1 1 1 1 1 1 1 46 1 1 1 1 1 1 1 1 1 1 1 1 1 47 1 1 0 0 1 1 1 1 1 1 1 1 1 48 1 1 1 1 1 1 1 1 1 1 1 1 1 260 49 1 1 0 0 1 1 1 1 1 1 1 1 1 50 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 APPENDIX B Morphometric data for Monarda section Aristatae. Character numbers correspond to those presented in the Materials and Methods section 261 Table 4. Morphometric data for Monarda Section Aristatae austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 Maysilles Laferriere Van Devender LeSueur Worthington Correll Neff Wilson Martin Scora Bye Jenkins Bye Spellenberg Scora Jenkins Bye Bolanos Neff Benitez Vega Tenorio Granfelt Turner Lane Estrada Dorr Vanderpool Nighswonger 8272 2017 96-378 887 8910 20146 91-13 8457 s.n. 2716 2684 89-308 8725 9243 2716 89-297 1795 1448 91-13 1984 2541 1773 6-224 2098 2767 2681 2381 759 3143 262 TEX TEX ARIZ TEX TEX LL TEX TEX ARIZ TEX MEXU TEX MEXU MEXU TEX ARIZ MEXU MEXU MEXU MEXU MEXU MEXU ARIZ ARIZ TEX CAS TEX OKL OKL 1 5.1 4.7 4.2 3.1 4.2 4.1 6.5 4.5 4.4 3.2 4.7 3.8 5.2 4.7 4.3 3.6 4.1 4.1 5.8 4.7 4.2 3.1 3.8 3.5 3.3 2.2 2.4 4.85 6.4 2 1 1.2 1.8 1.1 1 1.3 1.1 1.5 2.5 0.9 1.1 1.2 0.9 1.4 1.8 1 1.1 1.1 1.3 1.2 1.7 1.2 1.4 1.3 1.2 3.2 1.2 1.8 2.8 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 Johnson Ettner Williams First Massey Semple Folley Goodman Goodman Albers Carr Thalis Tharp Howell Correll Correll Correll Correll Correll Correll Henderson Lundell Crook Crook Nighswonger Nighswonger Pearce Sanders Lawson 254 77-1 331 7 2070 566 564 7849 8384 35013 13595 30 54996 2 25295 19007 32917 18997 16806 19448 63-938 15083 953 1009 672 1221 57 132 40 263 OKL OKL OKL OKL OKL OKL OKL OKL OKL TEX TEX TEX TEX LL LL LL LL LL TEX TEX TEX LL OKL OKL OKL OKL OKL OKL OKL 1 5.8 5.8 5.7 6.2 4.9 5.2 5.6 6.3 4.9 5.3 2.6 4.2 5.95 5.86 6.32 6.2 4.92 6.21 5.9 5.85 6.3 6.1 4.7 4.6 2.6 4.2 4.4 3.6 6.3 2 2.2 2.1 2.2 2.4 2.7 2.5 2.2 2.6 2.4 1.8 1.4 1.8 2.2 2 1.8 2.5 2.4 2.6 2.5 1.8 2.4 2.4 1.6 1.6 1.8 1.6 1.5 1.8 1.4 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 Pearce Proctor Thompson Thompson Thompson Huff Correll Correll Correll Correll Kutac Albers Albers Tharp Correll Landon Simpson Carter Ajilvshi Starbuck Dubrule Mahler Carter Maysilles Garcia Scora Semple Semple Segers 57 KEG0328 S0520 S0527 S0486 1391 27489 16407 23400 32847 s.n. 48019 45Ph025 51-535 16741 172 139 91 8400 1926 307 1135 13788 7771 399 2222 599 578 s.n. 264 OKL OKL OKL OKL OKL OKL LL LL LL LL TEX TEX TEX TEX LL RM TAMU TAMU TAMU TAMU TAMU TEX TEX TEX MICH TEX MO MO RM 1 5.3 7.2 4.3 4.7 2.3 5.2 4.1 3.4 4.5 4.6 5.2 6.2 5.7 3.8 4.5 4.3 2.8 4.7 3.8 5.2 4.5 4.3 4.1 3.3 3.5 6 5.3 4.5 5.8 2 1.6 2.6 1.2 2.5 1.4 1.8 1.7 1.3 1.8 1.3 1.8 2 1.5 1.7 1.6 1.5 1.7 1.5 1.7 1.5 1.8 1.7 1.6 1.8 2 3.2 1.9 1.9 2.2 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 Webster Correll Sanders Correll Lundell Correll Rachaner Warnock Hutchins Shinners Ballinger Sanders LeSassier Lawsen Daniels Tracy Higgins Rosson Demaree Churchil Theiret Haynes Wooton Wooton Wooton Neely Porter Long Kugel 7075 18391 75084 33768 14224 22048 132 5672 1133 30061 s.n. 75084 225 384 13 7991 6948 956 7554 7482138 59653 9692 s.n. s.n. s.n. 3566 9356 s.n. 2098 265 TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX TEX MSC MSC MSC MSC MSC TEX MSC KNK KNK NMC NMC NMC RM RM RM MSC 1 5.5 6.9 6 5.9 6.9 4.5 5.2 5.6 5.4 4.5 2.7 8.6 3.2 3.9 3.7 4.1 3.8 3.8 3.7 3.3 4.2 3.5 2.5 4.2 5.5 5.4 4.2 5.4 5.6 2 2.1 2.5 3.1 2.4 2.9 2.2 1.5 3.2 2.3 2.2 2.9 2.5 2.4 2.6 1.9 1.2 1.8 2.1 2.4 2.5 2.3 2.5 2.1 2.4 2.8 1.9 1.7 2.4 2.6 Table 4 (Cont’d) austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 3 5.1 5.3 3.2 3.6 3.7 5.8 4.9 5.3 3.5 3.2 3.5 6.6 5.3 5.2 4.1 3.6 3.7 7.2 5.1 5.4 4.1 3.3 3.2 5.7 4.2 3.4 3.6 7.2 7.8 4 0.44 0.9 0.6 0.51 0.63 0.61 0.42 1 0.5 0.42 0.7 0.6 0.41 1.2 0.52 0.43 0.62 0.57 0.45 0.8 0.68 0.53 0.62 0.51 0.51 0.6 0.3 0.85 1.2 5 1.8 1.8 1.7 1.5 1.8 2.9 2.9 3.1 1.5 1.3 1.4 4.6 2.7 2.7 1.6 1.2 1.7 2.7 2.5 2.8 1.7 1.2 1.6 3.2 2.1 1.3 2.2 5 4.8 266 6 1.1 2.1 1.8 1 1.3 1.5 1 9.2 4.2 1.2 1.4 1.6 0.8 1.5 1.9 2.2 1.6 1.8 2.1 1.5 1.6 1.8 1.4 1.8 1.8 0.8 3 11 11.5 7 3.1 2.8 2.3 1.5 2.1 3.2 3.4 2.9 2.1 1.6 2.2 4.2 3.3 3.1 2.4 1.7 2.1 3.3 3.6 3.1 2.2 2.1 2.4 2.9 2.6 1.5 2.3 3.9 4.1 8 2.9 3.9 3.5 3.2 3.5 3.8 3.3 3.2 4.1 3.5 3.4 4.1 4.3 4.2 3.2 3.3 4.1 3.6 3.4 3.5 3.7 3.3 3.3 4.2 3.8 4.1 3.8 3.7 3.9 9 0.24 0.3 0.28 0.24 0.32 0.28 0.23 0.24 0.31 0.3 0.24 0.28 0.34 0.31 0.26 0.29 0.3 0.31 0.32 0.3 0.31 0.27 0.28 0.29 0.31 0.18 0.25 0.26 0.28 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 3 6.9 6.8 4.9 6.6 6.8 7.2 7.1 7.4 6.5 6.6 4.8 5.9 5.5 6.4 7.2 6.2 6.5 6.4 7.4 7.2 6.8 7.5 4.5 4.2 4.6 4.7 4.55 5.1 4.7 4 1.1 0.86 0.92 0.85 0.76 0.98 0.95 1.1 0.86 0.8 0.52 0.63 0.68 0.72 0.86 0.73 0.75 8.6 1.2 0.98 0.98 1.1 0.84 0.75 0.9 0.93 0.86 0.95 0.62 5 4.2 4.7 3.8 4.5 4.4 4.6 4.8 4.7 4.4 4.3 3.5 3.9 4.2 4.5 4.8 4.5 4.5 4.6 4.9 4.7 4.6 4.8 0.9 0.85 1 1.2 1.1 2.2 1.3 267 6 10.5 9.5 10 11 8.5 8 10.5 11 8.5 9.5 7.5 10.5 10 11.5 11 10.5 11 9.5 8.5 9.5 11 10.5 2.5 2.4 3.1 3.2 2.8 6.1 2.6 7 3.6 3.8 3 3.6 3.7 4.1 4 3.9 3.8 3.8 2.5 4.1 3.6 3.7 3.8 3.6 4.1 4 3.5 3.8 3.7 3.8 2.8 2.85 3 3.2 3.05 3.3 3.1 8 4.1 3.7 3.9 3.8 2.9 3.9 3.8 3.9 3.7 3.8 3.8 3.8 3.9 3.5 3.6 3.5 3.7 3.8 3.9 4.1 4.1 3.9 4.1 3.6 4.2 4.1 4.05 4.5 2.9 9 0.3 0.27 0.26 0.25 0.26 0.27 0.28 0.29 0.3 0.28 0.28 0.27 0.29 0.26 0.28 0.26 0.25 0.28 0.27 0.26 0.28 0.27 0.26 0.23 0.25 0.45 0.25 0.36 0.26 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 3 5.15 3.8 4.3 4.6 4.4 4.65 4.5 4.3 4.45 4.35 5.2 3.9 4.5 5.2 4.6 4.6 4.5 4.3 5.2 4.6 4.4 4.65 4.6 5.6 5.7 7.3 5.2 6.3 6.5 4 1.05 0.6 0.72 0.9 0.75 1.2 1.05 0.81 0.88 0.8 0.98 0.52 0.74 0.92 0.9 0.89 0.85 0.7 0.96 0.65 0.87 0.88 0.92 1.2 1.1 1 0.69 0.78 0.84 5 2.8 0.9 1.1 1.1 1.2 1.2 1.1 1.2 1.2 0.9 1.4 0.8 1.1 1.8 0.9 1.1 0.98 0.86 1.4 1.1 0.85 0.9 0.95 1 1.2 7.8 2.7 4.4 4.4 268 6 2.4 1.8 1.9 2.6 2.4 3 2.5 2.5 2.6 2.2 2.2 2 2.4 5.2 3.1 3.1 3.2 2.6 4 2.5 2.4 2.7 3.2 0.7 0.6 8.5 9.8 10.5 13 7 3.2 2.6 2.9 3.2 3.1 3.15 3.2 3.2 3.1 2.8 3.1 2.6 3.2 3.2 3.1 3.1 2.9 2.9 3.2 3.1 2.85 3.1 3.1 4.9 5.1 3.4 3.4 4.3 3.5 8 4.2 3.8 3.75 4.1 4 4.2 4.1 4.1 4 3.8 4.1 3.75 3.2 4.35 4.08 4.05 4.15 3.5 4.2 3.2 4.1 4.1 4.1 0 0 3.8 3.9 3.5 3.6 9 0.34 0.22 0.19 0.26 0.25 0.3 0.29 0.26 0.27 0.23 0.32 0.24 0.27 0.31 0.24 0.32 0.26 0.25 0.35 0.3 0.26 0.26 0.4 0 0 0.25 0.26 0.28 0.27 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 3 6.4 3.8 4.8 5.2 4.2 3.8 4.5 4.2 4 4.2 4.1 6.5 4.6 3.7 4.3 3.8 3.6 4.1 3.8 4.2 5.8 4.5 3.6 3.9 4.1 4.3 3.5 4.1 5.8 4 0.75 0.6 0.9 1.1 0.72 0.6 0.8 0.8 0.65 0.7 0.71 0.7 0.78 0.62 0.73 0.62 0.58 0.62 0.63 0.81 0.75 0.72 0.56 0.6 0.72 0.76 0.68 0.65 0.72 5 4.2 1.3 2.3 1.6 1.8 1.5 1.7 1.6 1.6 1.7 1.7 2.7 1.9 1.5 1.7 1.5 1.6 1.5 1.7 1.8 2.4 2.1 1.4 1.5 1.7 1.5 1.3 1.7 1.8 269 6 9.2 3.5 3.8 4.1 4 3.6 3.6 3.8 3.7 3.8 4.1 7.2 4.23 3.5 4.2 3.2 3.4 4.1 3.7 4.2 5.2 4.5 3.2 2.5 2.6 2.4 2.8 3.8 4 7 3.3 2.3 3.1 3.2 2.55 2.3 2.4 2.6 2.1 2.6 2.6 4.3 2.6 2.53 2.6 2.1 1.9 2.5 2.6 2.5 3.8 3.1 2.4 2.3 2.4 2.6 2.4 2.6 2.1 8 3.5 3.5 5 4.5 3.95 3.5 3.7 4.2 3.4 3.78 4 3.8 3.8 3.6 4.1 3 3.2 3.9 3.7 3.6 3.7 3.8 3.2 3.5 4.1 4 3.6 3.8 4.2 9 0.28 0.25 0.33 0.46 0.3 0.28 0.28 0.33 0.28 0.3 0.32 0.5 0.26 0.27 0.34 0.25 0.28 0.31 0.29 0.27 0.3 0.33 0.26 0.29 0.31 0.27 0.24 0.27 0.32 Table 4 (Cont’d) austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 10 4.99 5.09 3.02 3.5 3.57 5.65 4.8 4.38 3.08 3.08 3.36 6.44 5.22 5.05 3.91 3.38 3.54 7.02 4.89 5.25 3.94 3.12 3.06 5.52 4.02 3.2 3.3 6.1 6.65 11 12 3 3 4 2 3 5 5 1 5 2 4 4 4 2 6 1 8 5 4 3 5 2 7 6 2 5 5 2 3 3 3.5 2.7 2 2.8 2.2 3.2 3.8 2.5 1.7 3.4 2 3.3 3.5 2.7 2.1 3.2 3.1 2.9 2.7 2.5 2.3 3.3 2.7 2.9 1.9 3.3 3 3.35 270 13 4.1 4.7 4.2 3.8 5.2 1.8 5.2 5.2 4.5 3.2 5.5 1.2 4.6 4.8 5.2 3.3 4.7 2.5 4.6 5.1 4.2 3.6 5.1 2.7 4.8 1.9 2.2 4.2 4.5 14 1.1 0.9 0.8 1 1.2 1.1 1.2 1 1 0.6 1.4 0.7 0.9 1.1 1.2 0.8 1.2 0.9 0.8 1 1.1 0.8 1.2 1.1 0.9 0.8 1.3 1.5 1.2 15 0.72 0.85 1.1 0.9 0.9 1.2 0.85 0.9 1.2 0.9 1.1 1.2 0.1 0.95 1.1 0.9 1 1.1 1 1.1 1.2 1.2 1.3 1.2 1.2 0.55 1.75 1.55 1.7 16 0.9 0.9 1.1 1.1 0.9 0.9 1.1 0.9 1.1 1.1 1.2 0.8 0.9 0.8 1.1 1.2 1.1 0.9 1.3 1.1 1.2 1.3 1.2 0.8 0.9 0.9 4.5 4.5 6 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 10 5.85 5.85 3.9 5.5 5.95 6.4 6.05 6.3 5.65 5.65 4.05 4.85 4.5 5.25 6.1 5.15 5.4 5.45 6.55 6.25 5.7 6.45 4.25 3.96 4.29 4.38 4.27 4.49 4.44 11 12 2 4 3 5 2 4 2 5 3 2 2 2 4 5 4 3 4 2 2 2 3 2 2 1 1 2 3 2 3 3 3.4 3.35 3.37 3.29 3.31 3.3 3.25 3.27 3.3 3 3.1 3.15 3.25 3.25 3.15 3.14 3.19 3.07 3.16 3.2 3.15 2.2 3.7 3.6 2.6 3.5 3.9 3.2 271 13 4.2 3.9 4.1 4.2 4.4 4.3 4.4 4.2 4 3.8 2.6 3.9 4.1 4.1 4.3 4.2 4.5 3.8 3.9 4.1 4.1 4.3 4.5 5.2 6.2 4.3 6 6.8 5.4 14 1.2 1.4 1.2 1.3 1.3 1.2 1.4 1.4 1.3 1.2 1.3 1.3 1.4 1.5 1.4 1.4 1.3 1.5 1.2 1.3 1.4 1.4 0.6 0.49 0.55 0.54 0.65 0.7 0.63 15 2.1 1.8 1.76 1.65 1.64 1.74 1.85 1.92 1.83 1.74 1.52 1.65 1.74 1.76 1.62 1.83 1.76 1.92 1.59 1.74 1.76 1.9 1.5 1.6 1.6 1.3 1.7 1.9 1.8 16 6 5 4.5 5.5 6 6 5.5 5 5 4.5 4.4 4.5 5 5.5 6 5.5 5 5.5 4.5 5 5.5 5 4.5 4.5 4.7 4.2 4.8 4.2 4.6 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 10 4.91 3.62 4.11 4.34 4.16 4.35 4.25 4.05 4.19 4.13 4.98 3.7 4.26 4.68 4.29 4.29 4.18 4.04 4.8 4.35 4.16 4.38 4.28 4.9 5.1 6.4 4.3 5.3 5.5 11 1 2 2 2 1 4 1 1 1 1 1 1 1 4 7 1 6 2 1 5 1 1 1 1 1 1 1 1 1 12 4.2 3.32 3.2 3.6 3.4 3.4 3.3 3.3 3.5 3.3 3.8 3.4 3.4 3.5 3.4 3.2 3.5 3.6 3.5 3.3 3.1 3.4 3.1 4 4.3 3.7 3.1 3.2 3.2 272 13 7.3 4.8 4.8 5.8 5.2 4.8 4.7 4.9 5.6 5.1 6.2 4.9 5.8 5.8 6.1 4.5 5.5 5.3 5.4 5.2 4.8 5.8 4.6 17 19 4.3 4 3.9 4.4 14 3.2 0.56 0.6 0.7 0.58 0.6 0.58 0.6 0.65 0.62 0.51 0.59 0.61 0.68 0.66 0.57 0.58 0.52 0.66 0.61 0.58 0.64 0.57 1.4 1.6 2 1.2 1.3 1.2 15 1.5 1.5 1.5 1.9 1.5 1.8 1.7 1.6 1.8 1.6 1.4 1.6 1.9 1.8 1.7 1.5 1.5 1.6 1.8 1.8 1.7 1.6 1.5 1.2 1.1 1.6 1.73 1.65 1.74 16 4.2 4.55 4.3 4.2 4.3 4.9 4.7 4.2 4.3 4.6 4.4 4.4 4.5 4.4 4.7 4.3 4.5 4.6 4.4 4.5 4.6 4.7 4.3 4.1 4 4.8 4.8 5.4 5.3 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 10 5.5 3.45 4.42 4.79 3.8 3.4 4.14 3.82 3.63 3.8 3.69 5.78 4.177 3.35 3.88 3.48 3.26 3.69 3.43 3.78 5.28 4.05 3.28 3.65 3.84 4.06 3.22 3.72 5.4 11 5 14 4 8 8 6 4 5 3 14 9 4 1 11 5 1 3 2 8 9 5 7 10 9 6 4 5 6 7 12 3.25 3.6 5.6 3.8 4 3.8 2.1 4.6 3.5 3.6 3.2 5.8 4.1 3.7 4.2 3.6 3.7 4.1 3.8 4 4.8 3.5 2.9 3.4 4.5 2.9 2.4 3.8 3.7 273 13 4.1 5.2 6.2 7 7.2 6.4 3.5 6 5.4 5.8 6.8 6.3 6.8 6.6 7 7.1 6.9 6.8 6.7 6.7 6.4 6.5 3.8 5.1 5.8 4.1 3.5 5.8 6.2 14 1.5 1.6 1.7 2.6 2.1 2.3 1.3 2.6 1.8 2 1.9 2.2 2.1 1.9 2.3 3.9 3.2 2.4 2.1 1.9 2.5 2.3 1.7 1.85 2.5 1.5 1.4 2.3 2.2 15 1.65 1.2 1.7 1.9 1.6 1.4 1 1.7 1.4 1.4 1.3 2.3 1.7 1.3 1.8 1.4 1.3 1.5 1.6 1.5 1.9 1.5 1.2 1.3 1.5 1.4 1.6 1.6 1.4 16 5.7 3.6 3 3.4 3.2 3.1 2.6 2.9 2.7 2.9 2.9 3.8 2.9 2.7 3.45 2.7 2.6 3.4 2.8 2.7 3.5 3.1 3.1 2.8 2.7 2.7 2.9 3.1 3.2 Table 4 (Cont’d) austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 17 0.19 0.24 0.21 0.25 0.28 0.21 0.18 0.25 0.24 0.26 0.35 0.2 0.2 0.26 0.23 0.25 0.28 0.18 0.19 0.21 0.19 0.24 0.27 0.25 0.26 0.3 0.5 0.5 0.6 18 7.22 6.99 7.04 6.43 6.65 6.98 6.53 7.18 7.08 6.86 6.92 6.59 7.07 6.49 6.63 6.87 6.75 6.61 7.12 7.05 6.72 6.65 6.83 6.88 6.79 3.02 7.6 7.2 7.32 19 1.71 2.2 2.65 1.94 2.21 2.53 2.19 2.43 2.51 2.41 2.46 2.25 2.63 2.15 2.21 2.31 2.29 2.1 2.24 2.15 2.13 2.22 1.98 2.06 2.18 1.36 1.5 1.62 1.49 274 20 0.58 0.48 0.6 0.5 0.5 0.72 0.7 0.55 0.35 0.39 0.49 0.53 0.58 0.49 0.61 0.64 0.54 0.57 0.56 0.51 0.53 0.51 0.52 0.62 0.63 0.38 0.56 0.59 0.55 21 3.61 3.59 5.59 4.41 3.25 2.02 4.2 3.66 3.67 3.56 3.77 4.56 4.23 3.71 3.95 3.74 3.52 3.29 3.61 3.74 4.05 3.2 3.68 3.6 3.9 2.36 4.25 4.36 4.15 22 0.07 0.07 0.06 0.06 0.06 0.06 0.07 0.06 0.07 0.07 0.07 0.07 0.07 0.07 0.06 0.07 0.06 0.07 0.06 0.06 0.06 0.07 0.07 0.06 0.07 0.06 0.13 0.13 0.16 23 0.98 0.62 1.15 1.07 0.97 0 0 0 1.05 1.04 1.03 0.86 0.92 1.04 1.03 1.05 0.97 0.96 1.09 1.1 1.08 0.97 0.75 0.86 1.04 0.75 0.98 0.88 0.95 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 17 0.7 0.5 0.5 0.5 0.6 0.6 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.5 0.6 0.6 0.5 0.6 0.5 0.5 0.6 0.5 0.47 0.39 0.43 0.39 0.45 0.43 0.46 18 7.85 7.76 7.52 8.12 7.76 8.15 7.52 7.68 8.42 7.58 7.86 7.68 8.21 7.3 7.5 7.72 7.5 7.55 8.26 7.6 7.72 7.59 7.72 8.51 7.04 6.16 7.43 7.5 8.32 19 1.55 1.56 1.54 1.63 1.69 1.54 1.62 1.55 1.6 1.57 1.59 1.73 1.48 1.64 1.52 1.48 1.58 1.56 1.57 1.58 1.6 1.68 1.61 1.69 1.51 1.68 1.59 1.68 1.72 275 20 0.56 0.62 0.64 0.61 0.58 0.57 0.55 0.56 0.56 0.62 0.63 0.58 0.63 0.57 0.52 0.54 0.64 0.62 0.56 0.55 0.58 0.55 0.64 0.58 0.7 0.68 0.64 0.71 0.77 21 4.18 4.36 4.28 4.39 4.42 4.16 3.98 3.85 4.19 1.2 4.6 4.2 3.89 4.2 4.12 4.19 4.25 3.95 3.98 2.85 4.2 4.15 3.55 4.01 3.33 3.6 4.73 3.62 3.95 22 0.12 0.13 0.17 0.16 0.18 0.2 0.15 0.14 0.14 0.18 0.15 0.14 0.13 0.14 0.15 0.13 0.14 0.15 0.15 0.17 0.14 0.13 0.15 0.13 0.16 0.11 0.13 0.13 0.15 23 0.92 0.94 0.86 0.86 0.89 0.84 0.81 0.89 0.95 0.94 0.92 0.92 0.95 0.99 0.98 1.02 0.96 0.95 0.94 0.95 0.98 0.96 0.66 0.66 0.65 0.62 0.64 0.65 0.66 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 17 0.39 0.42 0.42 0.46 0.44 0.36 0.35 0.42 0.45 0.43 0.41 0.43 0.42 0.44 0.46 0.42 0.42 0.39 0.43 0.47 0.44 0.45 0.41 3.5 3.5 0.5 0.5 0.55 0.54 18 7.18 7.62 8.32 7.12 6.52 7.52 7.52 6.88 7.35 8.21 7.14 7.82 8.43 7.69 7.53 6.88 7.15 8.42 8.12 7.26 7.68 7.64 7.25 6 6.1 7.68 7.73 7.49 7.5 19 1.53 1.59 1.58 1.48 1.67 1.6 1.58 1.59 1.55 1.68 1.55 1.62 1.68 1.59 1.58 1.72 1.55 1.63 1.68 1.53 1.32 1.64 1.57 2 1.9 2.19 2.07 1.86 2.05 276 20 0.7 0.5 0.66 0.61 0.53 0.75 0.63 0.67 0.65 0.68 0.69 0.61 0.66 0.65 0.64 0.66 0.65 0.66 0.65 0.67 0.62 0.68 0.7 0.8 0.8 0.58 0.63 0.62 0.63 21 3.45 3.66 3.85 3.42 3.52 3.86 4.12 3.69 3.66 4.02 3.72 3.81 3.62 3.71 3.86 3.72 3.42 4 3.98 3.55 3.62 3.62 3.42 1.5 1.4 3.5 1.7 2.67 2.39 22 0.16 0.14 0.12 0.15 0.12 0.12 0.12 0.13 0.15 0.12 0.12 0.14 0.14 0.13 0.15 0.12 0.15 0.12 0.14 0.13 0.12 0.15 0.16 0.21 0.2 0.15 0.12 0.14 0.13 23 0.6 0.49 0.71 0.7 0.51 0.52 0.68 0.65 0.62 0.64 0.64 0.71 0.68 0.63 0.67 0.64 0.65 0.68 0.64 0.65 0.69 0.6 0.72 0.38 0.4 0.94 0.94 0.92 0.85 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 17 0.5 0.5 0.6 0.53 0.45 0.45 0.22 0.5 0.43 0.41 0.38 0.51 0.41 0.39 0.48 0.4 0.41 0.47 0.41 0.38 0.48 0.37 0.34 0.41 0.49 0.32 0.28 0.44 0.51 18 7.42 6.21 8.56 9.58 9.9 7.6 8.5 9.4 6.8 7.78 6.82 8.21 6.68 7.6 8.12 7.1 6.92 8.89 8.45 6.59 8.47 6.42 6.32 7.12 9.12 8.3 6.72 9.5 9.4 19 1.74 1.58 1.84 2.41 1.88 1.76 1.83 1.95 1.72 1.8 1.62 1.79 1.85 1.82 1.86 1.8 1.75 1.89 1.85 1.83 1.82 1.62 1.63 1.71 1.86 1.85 1.83 1.92 2.02 277 20 0.63 0.63 0.65 0.68 0.64 0.65 0.65 0.66 0.68 0.66 0.65 0.67 0.66 0.64 0.62 0.53 0.63 0.64 0.62 0.7 0.68 0.67 0.66 0.65 0.65 0.65 0.64 0.66 0.65 21 1.92 2.6 2.94 2.84 3.5 2.72 2.8 3.1 2.65 2.8 2.67 2.89 2.74 2.81 2.96 2.68 2.76 2.88 2.82 2.8 2.9 2.59 2.61 2.79 3.12 2.81 2.82 3.45 2.92 22 0.12 0.09 0.05 0.07 0.05 0.06 0.06 0.07 0.05 0.06 0.08 0.05 0.06 0.07 0.06 0.04 0.04 0.05 0.06 0.06 0.07 0.08 0.09 0.07 0.06 0.05 0.05 0.08 0.09 23 0.93 0.62 0.73 0.74 0.71 0.66 0.69 0.68 0.75 0.65 0.64 0.62 0.6 0.65 0.64 0.62 0.64 0.75 0.63 0.65 0.62 0.64 0.65 0.66 0.68 0.67 0.6 0.62 0.65 Table 4 (Cont’d) austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 24 5.52 8.6 9.53 7.19 7.42 6.59 9.27 6.26 7.33 7.37 8.82 10.02 6.85 7.12 6.62 7.19 7.24 7.32 7.19 7.24 7.42 6.82 6.94 6.83 6.47 4.8 7.62 5.81 6.4 25 2.17 2.6 2.52 2.7 2.48 2.5 2.47 2.49 2.12 2.54 3.14 2.53 2.47 2.53 2.64 2.49 2.56 2.51 2.42 2.46 2.45 2.51 2.45 2.43 2.56 2.41 2 2.1 2.1 26 2 2.6 4.61 2.5 3.55 2.42 3.82 2.5 2.78 3.07 3.4 3.38 2.56 3.72 3.64 3.42 2.86 2.64 2.38 3.74 3.68 2.55 2.57 3.64 2.39 1.4 2.21 2.16 2.21 278 27 1.47 2.01 2.43 1.29 2.37 0.95 2.67 2.07 2.48 1.88 1.69 1.75 1.86 2.13 2.42 1.68 1.75 2.38 2.08 2.19 2.27 1.86 2.38 2.47 2.62 1.2 1.28 1.26 1.32 28 0.69 1.55 1.12 1.21 1.45 1.2 1.65 1.6 1.39 1.26 1.44 1.03 1.25 1.27 1.39 1.53 1.08 1.09 1.24 1.57 1.61 1.45 1.38 1.46 1.47 0.8 1 1.12 1.13 29 0.9 1.61 1.18 1.04 1.04 0.9 1.5 1.26 1.45 1.04 1.48 1.28 1.18 1.24 1.46 1.57 1.38 1.35 1.27 1.09 1.07 1.28 1.26 1.34 1.28 0.76 0.94 0.88 0.94 30 4.73 7.6 7.2 6.47 5.5 6.27 5.26 7.1 6.84 6.55 6.92 6.4 6.92 5.32 5.71 6.48 6.28 5.96 7.42 7.13 6.37 6.58 6.78 6.92 6.82 5 8.2 8.8 8.6 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 24 6.33 5.52 6.1 5.92 5.43 7.45 5.3 5.8 5.84 5.62 7.2 6.25 6.34 5.6 6.3 3.25 5.62 5.7 5.74 5.63 7.14 6.3 5.45 5.94 6 5.5 7.98 5.5 5.86 25 2.15 1.96 2.13 2.21 1.92 1.9 2.1 2.1 2.14 1.98 1.95 1.96 2.1 2.13 2.14 2.13 1.98 2.01 2.04 1.75 2.02 1.98 1.84 2.32 1.98 2.03 1.85 1.86 2.22 26 2.2 2.15 2.14 2.15 2.09 2.22 2.12 2.22 2.19 1.99 2.4 2.2 2.19 2.12 2.14 2.1 2.13 2.11 2.14 2.1 2.2 2.3 1.91 2.17 2.23 2.03 3.18 2.05 2.15 279 27 1.28 1.19 1.56 1.36 1.19 1.25 1.19 1.82 1.32 1.22 1.62 1.29 1.24 1.25 1.28 1.26 1.24 1.75 1.26 1.15 1.48 1.3 1.17 1.42 2 1.2 2.36 1.22 1.43 28 1.05 1.19 1.09 1.12 1.14 1.13 1.15 1.2 1.08 1.15 1.08 1.15 1.08 1.11 1.09 1.11 1.15 1.19 1.1 1.12 1.11 1.08 1.31 1.03 1.13 1.22 0.92 1.19 1.1 29 0.88 0.82 0.92 0.87 0.75 0.92 0.88 0.93 0.86 0.72 0.93 0.91 0.94 0.9 0.92 0.87 0.86 0.92 0.79 0.82 0.87 0.89 0.61 0.83 0.97 0.87 1.14 0.68 0.86 30 8.56 8.3 8.53 8.74 8.46 9.21 8.52 8.59 7.89 7.85 8.23 7.65 8.47 8.7 8.8 8.5 8.4 8.6 7.9 7.8 8.3 8.1 5.4 5.89 5.58 5.49 6.1 5.45 5.72 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 24 6.12 5.42 6.22 6.15 5.72 7.83 7.14 5.78 6.47 6.17 6.52 5.68 5.81 5.53 7.85 5.84 6.05 5.96 6.14 6.28 5.53 5.52 6.21 8 7.2 9.65 6.19 5.2 6.5 25 2.14 1.94 2.31 1.99 2.01 1.92 1.96 2.01 2.03 2.24 1.97 1.96 2.22 1.76 1.82 1.99 2.13 2.29 2.23 2.1 1.96 1.92 1.97 4.2 4.1 3.5 3.2 3.02 3.2 26 2.22 2.04 2.19 2.22 2.14 2.3 2.72 2.1 2.22 2.34 2.3 1.86 2.14 2.08 3.15 2.1 2.19 2.15 2.15 2.21 1.98 2.01 2.15 2 2.1 2.14 3.71 2.4 3 280 27 1.82 1.21 1.43 1.52 1.25 1.32 2.26 1.3 1.9 1.5 1.95 1.26 1.43 1.27 2.14 1.42 1.65 1.52 1.43 1.96 1.19 1.21 2.01 0.8 0.9 2.05 1.89 1.68 1.7 28 1.14 1.29 1.09 1.11 1.13 0.98 1 1.18 1.16 1.01 1.14 1.27 1.13 1.25 1.02 1.18 1.12 1.1 1.01 1.12 1.28 1.29 1.12 0.9 0.9 1.81 0.89 0.77 1.01 29 0.94 0.72 0.87 0.95 0.86 0.95 1.02 0.89 0.95 0.84 0.95 0.71 0.82 0.72 1.01 0.89 0.94 0.93 0.85 0.89 0.75 0.72 0.95 0.78 0.88 1.36 0.93 0.86 1.2 30 5.49 5.3 5.78 5.56 5.5 5.9 5.8 5.5 5.47 5.72 5.62 5.34 5.76 5.39 6.15 5.52 5.74 5.92 5.72 5.59 5.38 5.44 5.62 7.4 9 7.8 7.89 7.92 8.15 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 24 5.3 6.44 7.8 9.23 7.93 7.06 7.3 8.2 6.5 7.4 6.52 7.8 8.2 7.4 8.1 6.4 6.6 8.1 7.1 8.2 7.2 6.72 6.48 6.65 8.42 7.45 8.3 8.1 8.4 25 3.12 2.22 2.74 2.69 2.87 2.35 2.4 2.75 2.19 2.5 2.45 2.8 3.2 2.52 2.58 2.3 2.2 2.56 2.45 2.8 2.75 2.31 2.29 2.21 2.64 2.61 2.52 2.92 2.86 26 1.92 2.15 3.6 3.53 2.11 2.27 2.2 2.4 2.14 2.4 2.24 2.42 2.52 2.3 2.19 2.21 2.19 2.41 2.35 2.67 2.54 2.19 2.23 2.21 2.28 2.31 2.75 3.21 3.48 281 27 1.42 0.99 1.12 1.75 1.31 1.13 1.28 1.35 1.12 1.18 1.1 1.09 1.12 1.29 1.28 1.15 1.11 1.22 1.23 1.1 1.13 0.98 1.06 1.09 1.25 1.26 1.12 1.33 1.41 28 0.79 0.86 0.72 0.61 0.9 0.77 0.75 0.82 0.65 0.78 0.84 0.79 0.82 0.76 0.81 0.72 0.66 0.83 0.76 0.87 0.73 0.85 0.81 0.71 0.83 0.76 0.87 0.91 0.82 29 0.9 0.8 0.93 0.78 1.01 0.85 0.85 0.9 0.79 0.84 0.86 0.92 0.83 0.84 0.88 0.8 0.78 0.91 0.84 0.86 0.92 0.81 0.82 0.82 0.89 0.84 0.83 0.97 0.94 30 8.4 6.59 9.8 10.2 11.54 6.2 6.24 7.2 5.4 7.1 6.6 8.2 6.45 6.3 6.9 5.5 5.9 7.1 6.3 6.35 8.75 6.68 6.62 5.57 7.12 6.35 5.98 8.56 8.23 Table 4 (Cont’d) austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 31 2.59 3.7 3.5 2.58 2.6 2.8 2.73 3.4 3.1 2.9 2.87 2.62 2.91 2.63 2.58 3.24 3.1 2.67 3.5 3.42 3.08 2.92 3.04 2.88 2.79 2.93 4.1 3.8 4.31 32 0.6 0.58 0.65 0.49 0.52 0.61 0.51 0.64 0.61 0.59 0.62 0.58 0.57 0.51 0.53 0.57 0.55 0.52 0.64 0.6 0.57 0.56 0.6 0.61 0.63 0.08 0.96 0.86 0.91 33 0.83 0.58 0.66 0.66 0.59 0.62 0.63 0.59 0.61 0.6 0.61 0.59 0.57 0.61 0.63 0.4 0.64 0.61 0.59 0.6 0.63 0.65 0.64 0.61 0.62 0.16 0.72 0.81 0.77 282 34 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1.1 1.2 0.98 35 36 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 31 4.82 3.75 3.9 4.5 4.32 4.87 4.63 3.89 3.85 3.65 4.12 3.95 4.25 4 4.2 4.7 3.8 3.9 3.86 3.7 7.1 4.1 2.5 2.8 3.3 2.69 3.2 2.7 2.65 32 0.85 0.84 0.8 0.82 0.82 0.95 0.81 0.84 0.81 0.85 0.9 0.86 0.82 0.85 0.92 0.86 0.83 0.82 0.82 0.87 0.89 0.9 0.87 0.76 0.9 0.83 0.96 0.83 0.78 33 0.63 0.68 0.72 0.61 0.64 0.72 0.73 0.71 0.62 0.65 0.71 0.82 0.73 0.82 0.75 0.63 0.69 0.73 0.67 0.66 0.75 0.79 0.67 0.59 0.72 0.84 0.69 0.65 0.58 283 34 1.1 0.97 1.1 1.12 0.97 1.03 1.1 0.99 1.01 1.02 0.98 0.99 1.03 1.1 1.01 1.04 0.98 1.02 0.97 0.98 0.97 0.92 0.12 0.15 0.17 0.13 0.12 0.12 0.16 35 36 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 31 2.9 2.75 2.82 3.21 2.74 3.15 3.12 2.86 3.4 2.8 2.4 2.7 2.73 2.46 3.15 2.72 3.2 2.9 2.7 3.4 2.7 2.4 3.1 2.1 2.1 3.89 3.8 3.47 3.2 32 0.91 0.85 0.82 0.92 0.85 0.95 0.91 0.84 0.92 0.81 0.89 0.84 0.77 0.88 0.84 0.85 0.86 0.77 0.77 0.92 0.86 0.83 0.91 0.2 0.2 0.43 0.41 0.36 0.39 33 0.73 0.68 0.61 0.75 0.82 0.71 0.72 0.83 0.71 0.58 0.73 0.65 0.55 0.65 0.72 0.85 0.76 0.61 0.61 0.73 0.68 0.66 0.68 0.19 0.18 0.3 0.38 0.31 0.32 284 34 0.15 0.12 0.16 0.17 0.12 0.11 0.12 0.12 0.16 0.16 0.15 0.13 0.12 0.13 0.14 0.15 0.16 0.15 0.13 0.16 0.13 0.15 0.17 0 0 0 0 0 0 35 36 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 31 2.9 2.05 3.6 2.9 2.6 2.45 2.7 2.8 2.5 2.6 2.3 3.1 2.8 2.9 2.7 2.6 2.4 2.75 2.6 2.65 3.1 2.2 2.3 2.6 2.7 2.63 2.72 2.85 2.9 32 0.37 0.5 0.52 0.54 0.5 0.5 0.52 0.51 0.5 0.51 0.51 0.51 0.53 0.51 0.5 0.52 0.52 0.51 0.51 0.5 0.51 0.52 0.52 0.5 0.53 0.51 0.52 0.5 0.52 33 0.29 0.29 0.3 0.32 0.31 0.31 0.31 0.32 0.28 0.3 0.28 0.32 0.32 0.31 0.3 0.29 0.39 0.31 0.31 0.3 0.29 0.3 0.28 0.29 0.32 0.3 0.3 0.32 0.33 285 34 0 0.89 0.9 0.85 0.79 0.8 0.79 0.8 0.82 0.86 0.84 0.85 0.78 0.78 0.82 0.81 0.81 0.84 0.78 0.79 0.92 0.91 0.9 0.85 0.83 0.81 0.82 0.83 0.85 35 36 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 37 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) 38 austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 286 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 44 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 Table 4 (Cont’d) 38 citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 287 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 44 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) 38 clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 288 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 44 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 Table 4 (Cont’d) 38 parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 39 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 40 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 41 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 289 42 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 43 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 44 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 4 (Cont’d) 45 austro383 austro384 austro385 austro386 austro387 austro388 austro389 austro390 austro391 austro392 austro393 austro394 austro395 austro396 austro397 austro398 austro399 austro400 austro401 austro402 austro403 austro404 austro405 austro406 austro407 n.sp.1_440 citriodora358 citriodora359 citriodora360 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 290 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 4 (Cont’d) 45 citriodora361 citriodora362 citriodora363 citriodora364 citriodora365 citriodora366 citriodora367 citriodora368 citriodora369 citriodora370 citriodora371 citriodora372 citriodora373 citriodora374 citriodora375 citriodora376 citriodora377 citriodora378 citriodora379 citriodora380 citriodora381 citriodora382 clino166 clino167 clino168 clino169 clino170 clino171 clino172 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 291 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 4 (Cont’d) 45 clino173 clino174 clino175 clino176 clino177 clino178 clino328 clino329 clino330 clino331 clino332 clino333 clino334 clino335 clino336 clino337 clino338 clino339 clino340 clino341 clino342 clino343 clino344 mexicana475 mexicana476 parva450 parva451 parva452 parva453 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 292 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Table 4 (Cont’d) 45 parva454 pectinata181 pectinata182 pectinata183 pectinata184 pectinata185 pectinata186 pectinata187 pectinata188 pectinata189 pectinata190 pectinata191 pectinata192 pectinata193 pectinata194 pectinata195 pectinata345 pectinata346 pectinata347 pectinata348 pectinata349 pectinata350 pectinata351 pectinata352 pectinata353 pectinata354 pectinata355 pectinata356 pectinata357 46 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 47 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 293 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 LITERATURE CITED 294 Aiken, G., S. 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