Abstract a Taxonomic Reevaluation of Trifolium Variegatum Nutt

ABSTRACT A TAXONOMIC REEVALUATION OF TRIFOLIUM VARIEGATUM NUTT. BASED ON MORPHOLOGY by Benjamin George Foster Trifolium variegatum Nutt. is a species of clover (Fabaceae) found in western North America. As a result of its geographic, habitat, and phenotypic diversity, the species has a complex taxonomic history. Hundreds of herbarium specimens of this and related species were studied in order to reevaluate the taxonomic status of the numerous morphotypes found within this species complex. Five segregates, T. appendiculatum, T. geminiflorum, T. polyodon, T. subsalinum, and T. ultramontanum, were considered to be potentially distinct from T. variegatum. Based on close investigation and the results of multivariate analyses, two segragates were accepted at the species level. The species recognized are Trifolium polyodon, a Monterey County, CA, endemic, T. appendiculatum, a large-flowered species found in cismontane California and southwestern Oregon, and T. variegatum, the most widespread and phenotypically diverse taxon. A TAXONOMIC REEVALUATION OF TRIFOLIUM VARIEGATUM NUTT. BASED ON MORPHOLOGY

A Thesis

Submitted to the

Faculty of Miami University

in partial fulfillment of

the requirements for the degree of

Master of Science

Department of Biology

Benjamin G. Foster

Miami University

Oxford, Ohio

2015.

Advisor______(Michael A. Vincent)

Reader______(R. James Hickey)

Reader______(Richard C. Moore) Table of Contents Introduction ...... 1 Materials and Methods ...... 6 Results and Discussion ...... 10 “Trifolium geminiflorum” ...... 20 “Trifolium subsalinum” and “Trifolium ultramontanum” ...... 28 Trifolium polyodon ...... 37 Trifolium appendiculatum ...... 45 Further Discussion ...... 54 Taxonomic Treatment ...... 55 References ...... 69 Appendices ...... 72

List of Tables Table 1. Published taxa in the Trifolium variegatum species complex (herbarium acronyms from Thiers, [continuously updated]) ...... 3 Table 2. List of taxa and synonyms accepted by McDermott in An Illustrated Key to the North American Species of Trifolium (1908) ...... 4 Table 3. List of taxa and synonyms accepted by Zohary and Heller in The Genus Trifolium (1984) ...... 5 Table 4. List of taxa and synonyms accepted by Isely in The Jepson Manual (1993) ...... 6 Table 5. List of taxa and synonyms accepted by Vincent in The Jepson Manual (2012) ...... 6 Table 6. Characters and measurements used in this study for multivariate analyses ...... 8 Table 7. Character loadings on PC’s ...... 12 Table 8. Proportion of specimens collected by month ...... 46

iii

List of Figures Figure 1. Drawings of laciniate organs ...... 10 Figure 2. Principle components analysis projection of all T. variegatum s.l. OTU’s, colored according to potential groups...... 13 Figure 3. Principle components analysis projection of all T. variegatum s.l. OTU’s, colored according to accepted groups ...... 14 Figure 4. Principle components analysis 3-dimensional projection of all T. variegatum s.l. OTU’s, colored according to accepted groups ...... 14 Figure 5. Cluster analysis using average linkage of all OTU’s, colored according to potential groups ...... 15 Figure 6. Cluster analysis using average linkage of all OTU’s, colored according to accepted groups ...... 16 Figure 7. Cluster analysis using Ward’s method of all OTU’s, colored according to potential groups ...... 17 Figure 8. Discriminant analysis of hypothetical groups ...... 18 Figure 9. Discriminant analysis of accepted groups ...... 19 Figure 10. Discriminant analysis of recognized groups, with restricted characters ...... 20 Figure 11. Distribution map of “T. geminiflorum” specimens ...... 22 Figure 12. Beanplot of banner petal length ...... 22 Figure 13. Beanplot of stipule length ...... 23 Figure 14. Beanplot of leaflet length ...... 23 Figure 15. Beanplot of flower number ...... 24 Figure 16. Specimen with two plants, one exhibiting “T. geminiflorum” morphology and the other “T. variegatum” morphology (Botti 154 (CAS))...... 25 Figure 17. Specimen exhibiting “T. geminiflorum” morphology and “T. variegatum” morphology (Keck 1237 (CAS))...... 26 Figure 18. “Trifolium geminiflorum” morph...... 27 Figure 19. “Trifolium geminiflorum” morph...... 27 Figure 20. “Trifolium subsalinum” morph (Williams 83-84-6 (RENO))...... 29 Figure 21. “Trifolium subsalinum” morph...... 30

Figure 22. Distribution map of “T. subsalinum” specimens...... 31 Figure 23. “Trifolium subsalinum” morphs compared...... 31 Figure 24. Beanplot of the difference between banner petal length and calyx length ...... 32 Figure 25. Beanplot of banner petal length to calyx length ratio ...... 32 Figure 26. Beanplot of stem diameter ...... 33 Figure 27. Specimen from Arizona exhibiting intermediate morphology between “T. subsalinum” and T. variegatum s.s. (Buegge 873 (ASU)) ...... 34 Figure 28. Nevada specimen exhibiting intermediate morphology between “T. subsalinum” and T. variegatum s.s. (Kennedy 3056 (CAS)) ...... 35 Figure 29. “Trifolium ultramontanum” morph ...... 36 Figure 30. “Trifolium ultramontanum” morph ...... 37 Figure 31. Map showing the distribution of specimens with pigmented or unpigmented calyx lobes within the Trifolium variegatum complex ...... 37 Figure 32. Distribution map of T. polyodon specimens ...... 38 Figure 33. Drawing of dissected calyx of a specimen of T. polyodon (Heller 6759 (ISC)) ...... 39 Figure 34. Beanplot of leaflet width ...... 39 Figure 35. Distribution map of specimens with more than 5 calyx lobe teeth per calyx, all belonging to T. variegatum ...... 40 Figure 36. Beanplot of tooth number per calyx ...... 40 Figure 37. Beanplot of tooth length ...... 41 Figure 38. Specimen of T. variegatum with many calyx lobe teeth (Neese 9097 (BRY)) ...... 42 Figure 39. Specimen of T. variegatum with many calyx lobe teeth (Neese 9097 (BRY)) ...... 43 Figure 40. Cultivated specimen of T. polyodon (Yadon s.n. (MU)) ...... 44 Figure 41. PCA projection of all T. variegatum s.l. OTU’s, colored according to accepted groups ...... 45 Figure 42. Distribution map of T. appendiculatum specimens ...... 47 Figure 43. Beanplot of banner petal length ...... 47 Figure 44. Beanplot of wing petal length ...... 48 Figure 45. Beanplot of keel petal length ...... 48

Figure 46. Beanplot of connate filaments length ...... 49 Figure 47. Beanplot of peduncle length...... 49 Figure 48. Beanplot of seed length ...... 50 Figure 49. Comparisons of floral organs and seeds between T. appendiculatum and T. variegatum...... 50 Figure 50. Beak of T. appendiculatum keel petals ...... 51 Figure 51. Trifolium appendiculatum specimen with beaked keels, from Tulare County, CA (Straw 2620 (MU)) ...... 52 Figure 52. Trifolium appendiculatum specimen without beaked keels, from Tulare County, CA (Thorne 36989 (ISC)) ...... 53 Figure 53. Distribution map of T. polyodon specimens ...... 58 Figure 54. Distribution map of T. appendiculatum specimens ...... 61 Figure 55. Specimen of T. variegatum (Gillett and Crompton 12951 (MU)) ...... 66 Figure 56. Distribution map of T. variegatum specimens ...... 67

This work is dedicated to Our Lady, sedes sapientiae, rosa mystica, and causa nostrae laetitiae.

vii

Acknowledgements I owe thanks to all herbaria that supplied specimens on loan and to the curator of the Notre Dame Greene Herbarium, Barbara Hellenthal, for letting me look through the type specimens there. She, and the director of that herbarium, Dick Jensen of St. Mary’s College, on several occasions gave me some important insights on various aspects of this project. I would also like to thank Michael Vincent and MU for requesting and processing aforementioned loans for me. Gratitude ought also to be supplied to Kelsey Cook and Megan Ruffley for their comradery and company in 75 Upham Hall. Megan unselfishly spent her time watering my clovers while I was away, and Kel was always there to listen to whatever strange idea popped into my head. The debt I owe to my parents, Bill and Jane Foster, is significant and their support and encouragement for the past two years is pretty far down on the list. Likewise, thank you, Kathleen Arens, for agreeing to marry me. Lastly, I proffer thanks to my advisor, Michael Vincent, who broke the record for time spent consulting with graduate students during my stay at Miami, and to my committee members, Jim Hickey and Rich Moore. They have helped me come closer to understanding what it means to be a plant taxonomist.

viii

Introduction The flora of California (especially the California Floristic Province) is well studied because of its remarkable array of biodiversity. Its rate of endemism has been estimated to range from approximately 50% (2125 endemic species / 4452 native species; Raven and Axelrod, 1978) to 35% (2265 endemic minimum-ranked taxa / 6500 native minimum-ranked taxa; Baldwin, 2012). The landscape heterogeneity of this area, combined with a mediterranean-type climate (Axelrod, 1973), are likely important causes of this diversity. The entire western floristic region is also notable in its plant diversity (Takhtajan, 1986). The genus Trifolium L. is an exemplar of this pattern. According to Zohary and Heller (1984), 65% of native North American Trifolium species are found exclusively in the region west of the Rocky Mountains, while only 8% are located exclusively in the east. The genus Trifolium (the clovers) is a member of the Fabaceae and is characterized by its trifoliate leaves (with a few exceptions), papilionaceous flowers found in a head to head-like inflorescence, and herbaceous habit. It is further differentiated by its diadelphous stamens and persistent corollas. There are approximately 240 species of Trifolium, which can be found worldwide but is most speciose in North and South America, Eurasia, and Africa (Zohary and Heller, 1984). Its centers of diversity are the Mediterranean region and the parts of North and South America with mediterranean-type climates. As noted by Peter Raven (1963), on these two continents, the genus has an amphitropical disjunct distribution. Several species, such as Trifolium repens and T. pratense, are cultivated as forage crops (Zohary and Heller, 1984), and thereby have become introduced and naturalized worldwide. A phylogenetic study of the genus based on nuclear ribosomal DNA ITS and chloroplast trnL intron sequences by Ellison et al. (2006) confirmed its monophyly. This study was unable to determine if the New World clovers originated in either North or South America. Ellison et al. lumped all North American clovers into section Involucrarium, claiming that the supposedly well-apomorphied Involucrarium s.s. (Zohary and Heller, 1984) is not monophyletic. Involucrarium s.s, with the exception of one member species, is depicted in their phylogeny as separated into two monophyletic groups emerging from a large polytomy. It is arguable that this is evidence against recognizing a smaller Involucrarium. Trifolium variegatum Nutt. is a member of sect. Involucrarium (both sensu Ellison et al., 2006 and sensu Zohary and Heller, 1984). This species is distinguished by its annual life cycle, a wheel-like involucral bracts, lack of pubescence, uniform calyx lobes, and white-tipped corollas. It is found throughout western North America, ranging from British Columbia in Canada south through the western U.S. (Arizona, California, Idaho, Montana, Nevada, Oregon, Utah, Washington, and Wyoming) and into Baja California, Mexico. It is found throughout both Takhtajan’s Rocky Mountain Floristic Region (including the Vancouverian and Rocky Mountain Provinces) and his Madrean Floristic Region (including the Great Basin, Californian, and Sonoran Provinces; Takhtajan, 1986). Within these regions, it can be found from sea-level up to 10,000 feet in elevation, and can grow along streams, in vernal pools, in gravel, in open meadows, and

1 in forests. Trifolium variegatum displays a wide range of morphological diversity, which has resulted in a complicated taxonomic history. Originally described by Thomas Nuttall in 1838, T. variegatum s.l. encompasses approximately 20 names (Table 1). Only one of these names, T. polyodon, is currently recognized as a species separate from the rest of the complex, with the others relegated to varietal status or synonymy within T. variegatum. In addition to all of these published species and varieties, various authors have introduced new combinations. Studies by McDermott (1908), Zohary and Heller (1984), Isely (1993), and Vincent (2012) have all used different names to describe the morphological diversity in this species complex. These important contributions will be discussed in further detail.

The first taxonomic review of this species complex was provided by Laura McDermott (1910) in An Illustrated Key to the North American Species of Trifolium. She accepted T. variegatum, T. appendiculatum, and T. polyodon as species, having grouped many other names into three varieties of T. variegatum (Table 2). Into T. variegatum var. pauciflorum (Nutt.) McDermott she subsumed T. dianthum Greene, T. pusillum Greene, T. geminiflorum Greene, T. ultramontanum Greene, T. phaeocephalum Greene, and T. subsalinum Greene. She placed T. calophyllum Greene into T. variegatum var. trilobatum (Jeps.) McDermott, and placed T. morleyanum Greene and T. variegatum var. major Lojac into T. variegatum var. melananthum (Greene) McDermott. Trifolium splendens Heller and T. rostratum Greene were considered synonymous with T. appendiculatum Lojac. Some of these synonymized names were given the rank forma, but here this is ignored for clarity. McDermott’s forms are listed in synonymy in the taxonomic section of this thesis. It is in McDermott’s monograph that the confusion caused by the name Trifolium pauciflorum is made manifest. In short, the name was first published by d’Urville in 1822 for a species located in Asia Minor, but was also used illegitimately for a North American species by Nuttall in 1838. Thus, there are two type specimens for the name T. pauciflorum. In 1841, Steudel noted this problem and called Nuttall’s species T. oliganthum as a replacement name for that of Nuttall. Currently, Trifolium oliganthum is not considered a part of the T. variegatum species complex (Zohary and Heller, 1984, among others). However, McDermott used T. pauciflorum Nutt. as the basionym for one of her new varieties of T. variegatum, in addition to having treated T. oliganthum as a separate species. A number of floras have also used this varietal nomenclature, which is allowed under the Code [Art. 11.2]. However, this means that T. variegatum var. pauciflorum and T. oliganthum share the same type specimen, and therefore T. variegatum var. pauciflorum (Nutt.) McDermott is an obligate synonym of T. oliganthum. Additionally, in 1883 Lojacono mentions a variant of T. pauciflorum Nutt., but does not give it a new name. It is clear that the T. pauciflorum Lojacono discusses is not the T. pauciflorum of Nuttall, and is instead the smaller morph of T. variegatum. Lojacono’s discussion is noted by Greene, who in 1897 gives Lojacono’s concept the name T. geminiflorum. Thus, T. pauciflorum Nutt. and all combinations based on it can only be referred to T. oliganthum Steud. The few- flowered specimens of T. variegatum can be called T. geminiflorum.

A world-wide monograph of Trifolium by Zohary and Heller (1984) also revised Trifolium variegatum (Table 3). Trifolium polyodon was recognized as a separate species. Trifolium variegatum was divided into two varieties: T. variegatum var. variegatum (including as synonyms T. melananthum Hook. & Arn., T. pauciflorum sensu Lojac., T. trilobatum Jeps., T. morleyanum Greene, T. calophyllum Greene, T. dianthum Greene, T. geminiflorum Greene, T. phaeocephalum Greene, T. pusillum Greene, T. subsalinum Greene, and T. ultramontanum Greene) and T. variegatum var. rostratum (Greene) J. Martin emend. D. Heller (nom. inval.; art. 30.8; including T. appendiculatum Lojac., T. rostratum Greene, and T. splendens Heller). Duane Isely also worked on this species complex, though he never revised it formally, having chosen to call his divisions “phases” instead of “varieties”. His opinions were presented in The Jepson Manual (Hickman, 1993) and in the posthumously published Native and Naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and Hawaii) (Isely, 1998). He tentatively lumped T. polyodon in with the rest of T. variegatum, which he divided into several phases (Table 4). His phases differ between the two publications (the former with 5 and the latter 7). Only the Jepson Manual phases are reported here. Phase 1 consists of T. appendiculatum Lojac. var. rostratum (Greene) Jeps., phase 2 of T. variegatum var. melananthum (Hook. & Arn.) Greene and T. appendiculatum Lojac. var. appendiculatum, phase 3 of T. variegatum var. trilobatum (Jeps.) Jeps. and part of T. appendiculatum Lojac., phase 4 of T. polyodon Greene, and phase 5 of T. geminiflorum Greene. Michael Vincent’s contributions can be found in the The Jepson Manual, ed. 2 (Baldwin et al., 2012). He recognized T. polyodon as a species separate from T. variegatum and its 3 varieties (Table 5). Trifolium variegatum var. geminiflorum (Greene) Vincent is roughly equivalent to Isely’s phase 5 and what was commonly called T. variegatum var. pauciflorum (Nutt.) McDermott. The second recognized variety is T. variegatum var. major Lojac., which corresponds to T. appendiculatum Lojac. var. appendiculatum, T. variegatum var. melananthum (Hook. & Arn.) Greene, and Isely’s phase 2. Finally, T. variegatum Nutt. var. variegatum, is equivalent to T. appendiculatum Lojac. var. rostratum (Greene) Jeps. and Isely’s phases 1 and 3. These treatments have done much to advance our knowledge of this group. However, the diversity of opinion, increasing through time, suggests that further work is necessary. Through the examination of hundreds of herbarium specimens, this project has aimed to reevaluate the taxonomy of this species complex by identifying taxon limits as supported by morphological data. It is hoped that this study will stabilize the group’s nomenclature, provide a resource for non-taxonomic work based on this species, such as biogeography or population genetics, and be a foundation for any future phylogenetic analysis.

Table 1. Published taxa in the Trifolium variegatum species complex (herbarium acronyms from Thiers, [continuously updated]).

Name Publication Type Herbarium

Trifolium appendiculatum Lojacono Nuovo Giorn. Bot. Ital. 15: 181, Lemmon s.n. PAL? 1883 Trifolium calophyllum Greene Pittonia 3: 213, 1897 E.L. Greene s.n. NDG

Trifolium dianthum Greene Pittonia 3: 217, 1897 J. Macoun 94 NDG

Trifolium melananthum Hooker & Arnott Bot.Beechey Voy. 331, 1838 Douglas s.n. K

Trifolium morleyanum Greene Erythea 3: 47, 1895 Baker & Nutting NDG s.n. Trifolium geminiflorum Greene Pittonia 3: 216, 1897 R.M. Austin s.n. NDG

Trifolium phaeocephalum Greene Pittonia 3: 216, 1897 R.M. Austin s.n. NDG

Trifolium polyodon Greene Pittonia 3: 215, 1897 E.L. Greene s.n. NDG

Trifolium pusillum Greene Pittonia 3: 217, 1897 C.C. Parry 45. NDG

Trifolium rostratum Greene Proc.Acad.Nat.Sci.Philad. 47: 54, Chestnut s.n. NDG 1895 Trifolium splendens A.Heller Muhlenbergia 1: 115, 1905 A. Heller 6691 BKL

Trifolium spinulosum var. triste Torrey and A. Fl. N. Amer. 1: 318, 1838 T. Nuttall s.n. BM Gray Trifolium subsalinum Greene Pittonia 3: 219, 1897 E.L. Greene s.n. NDG

Trifolium trilobatum Jepson Bull. Torrey Bot. Club 18: 322, 1891 W.L. Jepson 14854 JEPS

Trifolium ultramontanum Greene Pittonia 3: 218, 1897 E.L. Greene s.n. NDG

Trifolium variegatum Nuttall Fl. N. Amer. 1: 317, 1838 T. Nuttall s.n. NY

Trifolium variegatum var. major Lojacono Nuovo Giorn. Bot. Ital. 15: 183, Lemmon s.n. PAL? 1883 Trifolium variegatum var. parunuweapensis S.L. Utah Flora (ed. 3) 425, 2003 S.L. Welsh & K.H. BRY Welsh Thorne 25120

Table 2. List of taxa and synonyms accepted by McDermott in An Illustrated Key to the North American Species of Trifolium (1908). McDermott’s Recognized Taxa Basionyms and Synonyms T. variegatum var. pauciflorum (Nutt.) McDermott T. pauciflorum Nutt. T. dianthum Greene T. pusillum Greene T. geminiflorum Greene T. ultramontanum Greene T. phaeocephalum Greene T. subsalinum Greene

T. variegatum var. trilobatum (Jeps.) Jeps. T. trilobatum Jeps. T. calophyllum Greene

Table 2. (continued)

T. variegatum var. melananthum (Hooker & Arnott) Greene T. melananthum Hooker & Arnott T. morleyanum Greene T. variegatum var. major Lojac

T. appendiculatum Lojac. T. splendens Heller T. rostratum Greene

T. polyodon Greene

Table 3. List of taxa and synonyms accepted by Zohary and Heller in The Genus Trifolium (1984). Zohary and Heller’s Recognized Taxa Basionyms and Synonyms T. variegatum Nutt. var. variegatum T. melananthum Hook. & Arn. T. pauciflorum sensu Lojac. T. trilobatum Jeps. T. morleyanum Greene T. calophyllum Greene T. dianthum Greene T. geminiflorum Greene T. phaeocephalum Greene T. pusillum Greene T. subsalinum Greene T. ultramontanum Greene T. variegatum var. pauciflorum (Nutt.) McDermott T. variegatum var. pauciflorum (Nutt.) McDermott fo. phaeocephalum (Greene) McDermott T. variegatum var. trilobatum (Jeps.) Jeps. T. variegatum var. melananthum (Greene) McDermott T. variegatum var. melananthum (Greene) McDermott fo. morleyanum (Greene) McDermott T. variegatum var. melananthum (Greene) McDermott fo. major (Lojac.) McDermott

T. variegatum var. rostratum (Greene) J. T. appendiculatum Lojac. Martin emend. D. Heller (nom. inval.; art. T. rostratum Greene 30.8)* T. splendens Heller T. appendiculatum Lojac. fo. rostratum (Greene) McDermott T. appendiculatum Lojac. var. rostratum (Greene) Jeps.

T. polyodon Greene

*Trifolium variegatum var. rostratum (Greene) J. Martin is not a valid name, since it was published in a Ph.D. thesis (art. 30.8). Heller’s emendavit does not rectify the situation, since this act is taxonomic, not nomenclatural (art. 47.1). Even if this were to be viewed as valid publication, it occurred in 1984, later than Trifolium variegatum var. rostratum (Greene) C.L.

Hitch. (Vasc. Pl. Pacif. N.W. 3: 370. 1961), and thus is a superfluous name, and therefore illegitimate (art. 52.1).

Table 4. List of taxa and synonyms accepted by Isely in The Jepson Manual (1993). Isely’s Recognized Taxa Synonyms Phase 1 T. variegatum Nutt. var. variegatum T. appendiculatum Lojac. var. rostratum (Greene) Jeps.

Phase 2 T. variegatum var. melananthum (Hook. & Arn.) Greene T. appendiculatum Lojac. var. appendiculatum

Phase 3 T. variegatum var. trilobatum (Jeps.) Jeps. T. appendiculatum Lojac.

Phase 4 T. polyodon Greene

Phase 5 T. geminiflorum Greene

Table 5. List of taxa and synonyms accepted by Vincent in The Jepson Manual, ed. 2 (2012). Vincent’s Recognized Taxa Basionyms and Synonyms T. variegatum var. geminiflorum T. geminiflorum (Greene) (Greene) Vincent T. variegatum var. pauciflorum sensu McDermott Isely phase 5

T. variegatum var. major Lojac. T. appendiculatum Lojac. var. appendiculatum T. variegatum var. melananthum (Hook. & Arn.) Greene Isely phase 2

T. variegatum Nutt. var. variegatum T. appendiculatum Lojac. var. rostratum (Greene) Jeps. Isely phases 1 & 3.

T. polyodon Greene

Materials and Methods Specimens from the following herbaria were utilized for this study: ASU, BM, BRY, CAS, ISC, MU, NY, MO, RENO, SD, SRP, and WIS (for herbarium acronyms, see Thiers, [continuously updated]). Dried specimens were examined under a dissecting microscope. Macroscopic linear measurements and angle measurements were made using a metric ruler and protractor, respectively. Microscopic characters were measured using a Wild MC3 dissecting microscope and a calibrated ocular micrometer. Flowers were rehydrated using 1% Aerosol OT (Ayensu, 1967). They were then dissected in order to yield measurements for corolla, calyx, stamen, and pistil characters. Characters measured are recorded in Table 6. In order to measure characters at similar developmental stages across all specimens, a number of characters to be measured

6 were based on nodal position. Unless the flowers or leaves were too underdeveloped, the third node from the apex was chosen. The involucres and stipules of these species are laciniate; lengths measured for these organs are depicted in Fig. 1. Specimens exhibiting all measured characters and representing the full range of morphological and geographical diversity were selected for use in the statistical analyses. Within T. variegatum s.l., 275 specimens were chosen. The full set of measurements for all specimens, with updated nomenclature, is in Appendix 2. Principle components analysis (PCA; Pearson, 1901; Hotelling, 1933) was carried out using several different computer programs: R (R Core Team, 2013), NTSYSpc (Rolf, 2012), and JMP (SAS Institute Inc., 2013). All R code is provided in Appendix 3. PCA was carried out using the ‘princomp’ function, and produced in R. Characters selected for use in this analysis represent a subset of all characters measured. They were selected for their consistency (it was easy to make measurements of the same object at the same developmental stage for all specimens), variability (for some characters, measurements were more or less invariant, and therefore were not included in such analyses [Sneath and Sokal, 1973]), and their assumed developmental independence (logically correlated characters were left out [all characters left out of CA and PCA, as shown in Table 6, were unused for this reason], those highly correlated but of different organs remained in the analysis [Sneath and Sokal, 1973]). The package ‘scatterplod3d’ (Ligges and Mächler, 2011) was used to project 3 principle components onto 3- dimensional space. Cluster analyses (CA) were carried out in NTSYSpc and JMP (Rolf, 2012; SAS Institute Inc., 2013). Figures given here were produced in JMP. The same characters used in PCA were used in these analyses. Average linkage (also known as UPGMA; Sokal and Michener, 1958) and Ward’s methods (Ward, 1963) were used to produce different phenograms. Average linkage links clusters together that have the least average distance between them; Ward’s method creates clusters that have the least possible sum of the squares deviations, and often results in more compact clusters (Marhold, 2011). These two clustering methods perform well overall according to Dunn and Everitt (1982), in comparison with other methods. In all CA’s, the dataset was standardized before analyses were conducted. Canonical discriminant analyses (DA; Fisher, 1936) were also carried out in JMP using the same characters for CA and PCA. Despite Marhold’s warning that “the characters that define the groups should not be involved in the analyses to avoid circular argumentation” (Marhold, 2011), important characters are not left out in the initial DA’s. Discriminant analyses including groups not recognized by this study are used as evidence against them. Despite the DA reducing axes in order to maximize the distance between predefined groups, its inability to do so can be viewed as a sign that such groups are not well supported. Additionally, a restricted DA, leaving out characters used to identify species, was also performed. These characters are recorded in Table 6. This last DA was used to evaluate group hypotheses and discover previously unused characters for identification.

Character measurements were looked at in a simpler way using the ‘beanplot’ (Kampstra, 2008) package in R. In ‘beanplot’, the ‘beanplot’ function produced beanplots, which are figures depicting individual measurements as lines in a one-dimensional scatterplot upon which an estimated density distribution and average line are superposed. Beanplots are useful because they show actual overlap in measurements between groups. In sections of this paper arguing against the recognition of certain groups, the five hypothetical groups are presented in each beanplot. In sections arguing for the recognition of groups as species, only the recognized taxa are presented, so that, for instance, when comparing T. appendiculatum to T. variegatum, the T. variegatum being compared to is the wider and not the stricter circumscription. All characters were tested for normality using the Shapiro-Wilk test (Shapiro and Wilk, 1965). Since it is valuable to present measurements in their original units, the dataset was not transformed. Thus, the non-parametric Kruskal-Wallis rank sum test (Kruskal and Wallis, 1952) was used to test equivalence between multiple groups. In the package ‘pgirmess’ (Giraudoux, 2014), the function ‘kruskalmc’ was used to perform multiple comparisons between groups. These results are listed in Appendix 1. They are not given in the thesis body because their usefulness is doubtful. P-values, obtained from the Kruskal multiple comparison test, indicate the “probability of obtaining the difference observed, or one that is more extreme, considering the null is true” and not “the probability that the null hypothesis is true” (Biau et al., 2010). Thus, they are simply a form of evidence against the null hypothesis. Since the data itself is also evidence against the null hypothesis, simply looking at it is enough for the evaluation of taxonomic hypotheses. Amount of overlap and degree of difference within the data are more important for taxonomy than statistically significant differences. Distribution maps were produced in R using packages ‘maps’ (Becker et al., 2014a) and ‘mapdata’ (Becker et al., 2014b). For specimens without latitude and longitude data on the label, estimations of coordinates were made in Google Earth for those where approximation could be made to an area smaller than a county. Google Earth was also used to estimate elevation when that information was not on the label. Table 6. Characters and measurements used in this study for multivariate analyses.

Character PCA/CA Restricted DA Description

Stem Diameter X X Measurement of the diameter of the stem halfway between the chosen node and the node above it Stipule Length X Measurement of the length of one stipule at the chosen node Stipule Lobe X X Measurement of the length of the longest stipule Length free segment at the chosen node Petiole Length X X Measurement of the length of the petiole at the chosen node Petiole X X Measurement of the diameter of the petiole at the Diameter chosen node

Table 6. (continued) Leaflet Base X X Measurement of the angle of the base of the Angle terminal leaflet Terminal Leaflet X X Measurement of the length of the terminal leaflet at Length the chosen node Terminal Leaflet X X Measurement of the width of the terminal leaflet at Width the chosen node Petiolule Length X X Measurement of the length of the petiolule at the chosen node Petiolule X X Measurement of the diameter of the petiolule at the Diameter chosen node Head Length X Measurement of the length of the inflorescence at the chosen node Flower Number X Number of flowers in one inflorescence at the chosen node Peduncle X X Measurement of the length of the peduncle at the Length chosen node Peduncle X X Measurement of the diameter of the peduncle at Diameter the chosen node Involucre X Measurement of the radius of the involucre at the Radius chosen node Involucral Lobe X X Measurement of the length of the longest involucral Length lobe at the chosen node Calyx Length Measurement of the length of the calyx Calyx X Measurement of the circumference of the calyx Circumference Calyx Lobe X Measurement of the length of the calyx lobe Length Calyx Lobe X Measurement of the width of the calyx lobe Width Calyx Tube X Measurement of the length of the calyx tube Length Calyx Lobe X Number of calyx lobe teeth per calyx Tooth Number Banner Petal X Measurement of the length of the banner petal Length Banner Petal X Measurement of the maximum width of the banner Width petal Wing Petal Measurement of the length of the wing petal Length Wing Petal X Measurement of the length of the wing petal lamina Lamina Length Wing Petal X Measurement of the maximum width of the wing Lamina Width petal lamina Keel Petal Measurement of the length of the keel petal Length

Table 6. (continued) Keel Petal X Measurement of the length of the keel petal lamina Lamina Length Keel Petal X Measurement of the maximum width of the keel Lamina Width petal lamina Connate X Measurement of the length of the connate filaments Filaments Length Style Length X Measurement of the length of the style

a b Fig. 1. Drawings of laciniate organs. a. The involucre from a specimen of T. variegatum is depicted (Gregory et al. 3236 (SD)). The involucre radius is marked by a line and the involucral lobe length is shown by an arrow. This latter measurement was made on the lobe with the longest distance between apex and sinus. b. A stipule from a specimen of T. appendiculatum is depicted (Straw et al 2620 (MU)). The stipule length is marked by a line and stipule lobe length by an arrow. The latter is the length of the longest free segment on the stipule. The partially shown organs appearing, from left to right, are the stem, the peduncle, and the petiole. The peduncle is free from the stipule, while the petiole is adnate to it. In this particular stipule, the connection between stipule and petiole has partially torn.

Results and Discussion PCA and CA supported the recognition of 3 taxa, Trifolium polyodon, T. appendiculatum, and T. variegatum. Other groups of specimens with diverging morphologies were not supported (“T. geminiflorum”, and “T. subsalinum”). These groups were further examined in order to verify this result and will be discussed later in this section. In Fig. 2, all operational taxonomic units (OTU’s) are projected onto a two-dimensional space defined by principle components 1 and 2. Principle Component (PC) 1 (50.9%) reflects general plant size, since almost every

10 character is loaded onto it in the same direction (Table 7; Dunn and Everett, 1982). PC 2 (11.2%) has a suite of reproductive characters loaded in one direction and vegetative in the other. The projection (Fig. 2 and Fig. 3) yields a tight cluster of OTU’s on the right and a more scattered cluster on the left, with a relatively small gap between the two, cutting from the bottom left part of the graph to the top right. This gap bisects the right angle between PC 1 and PC 2, indicating that the separation is due to more than overall plant size. Trifolium appendiculatum predominates the left cluster, with the three T. polyodon specimens intermixed, and T. variegatum predominates the right cluster, with “T. subsalinum” scattered diffusely throughout and “T. geminiflorum” marginally connected. Figure 3 shows the same projection identified only to taxa recognized in this study. Intermixed OTU’s were examined and determined to belong to their designated group based on post hoc identifying characteristics. Several specimens of T. appendiculatum fell into the T. variegatum cluster. This also occurred in CA, and will be discussed there. OTU’s were also projected onto the first 3 PC’s (Fig. 4). PC 3 (5.9%) is difficult to interpret as there is no pattern in the proportion or direction of loadings. Plotting PC 2 against PC 3 yields one mass of specimens without any gaps (not shown). CA using average linkage produced Fig. 5 and Fig. 6. In Fig. 5, OTU’s are colored according to accepted and potential groups. Trifolium polyodon is shown to be separated by a large distance from the other taxa (the rightmost cluster). The next branch separates T. appendiculatum from T. variegatum + “T. geminiflorum” + “T. subsalinum”. Several specimens of T. variegatum are found in the T. appendiculatum cluster, and several specimens of T. appendiculatum are found in the T. variegatum cluster. The misplaced T. variegatum specimens (Andrews 344 (ISC), Heller 6727 (ISC), Calder & MacKay 30139 (CAS), Morgan 1065 (MU), Vincent 8191 (MU), and Morgan 2033 (MU)) were likely growing in very moist localities. This is known for a fact with Vincent 8191 (MU; personal communication). They all have large leaves and are generally robust plants. However, they retain classic T. variegatum oblanceolate leaflets and have small flowers. The misplaced T. appendiculatum specimens (Evermann s.n. (CAS), Howell 20825 (CAS), Heller 10710 (CAS), and Heller 10030 (CAS)) all share classic T. appendiculatum obovate leaflets and have long flowers. They were likely young undeveloped plants or were growing in dry areas. “T. geminiflorum” and “T. subsalinum” are found dispersed throughout T. variegatum. Figure 6 depicts OTU’s colored according to recognized taxa. It shows the intermixture between T. appendiculatum and T. variegatum better. Figure 7 is a CA produced using Ward’s method colored according to potential groups. It exhibits less intermixing than the previous figures. Trifolium polyodon is found within the T. appendiculatum cluster. As mentioned previously, this is likely due to this method’s tendency to avoid small clusters at upper hierarchies in favor of compactness. Discriminant analyses yielded similar results. Figure 8 depicts a DA performed using all hypothetical taxa as its predefined groups. While T. appendiculatum and T. polyodon are separated from all other groups, T. variegatum s.s., “T. geminiflorum” and “T. subsalinum” overlap considerably. It is surprising that “T. geminiflorum” was not depicted as a separate group, since the characters used to identify it were also measured and used in this analysis.

“Trifolium subsalinum”, however, was selected based on a qualitative character, and thus this analysis is good evidence that no other characters support “T. subsalinum” as a separate species. In Fig. 9, DA was performed using accepted taxa as predefined groups. Since calyx lobe tooth number is a determining character for T. polyodon, this graph is tautological. It does show that, unlike “T. geminiforum”, the characters chosen to distinguish these groups actually do so. In Fig. 10, the most important characters, as determined by DA and through personal experience, were removed. The analysis still returned 3 separate groups, and important discriminating characters are depicted. This analysis supported conclusions, formed elsewhere, that stipule length and peduncle length are useful for identifying T. polyodon and T. appendiculatum from T. variegatum, respectively. In order to make sense of these analyses, all groups were investigated further. Putative diagnostic characters were compared to actual measurements and their geographical distribution. Emphasis was placed on stable characters for delimitation, following Cronquist’s advice of delimiting species through the valleys of morphological distribution (Cronquist, 1988). Hypothesized segregates of T. variegatum s.l. are here discussed in the following order: “T. geminiflorum”, “T. subsalinum” and “T. ultramontanum”, T. polyodon, and T. appendiculatum. Groups not recognized by this study as separate taxa are marked with quotation marks. Representative specimens are listed for them in this section, whereas the representative taxa of accepted taxa are listed in the taxonomic treatment.

Table 7. Character loadings on PC’s.

Character Comp.1 Comp.2 Comp.3 Comp.4 Comp.5 Stem Diameter -0.175 -0.321 Stipule Lobe Length -0.112 -0.266 -0.107 -0.272 0.345 Petiole Length -0.121 -0.163 0.469 0.257 -0.158 Petiole Diameter -0.194 -0.285 0.151 -0.143 Petiolule Length -0.160 -0.252 -0.157 -0.133 Petiolule Diameter -0.187 -0.257 0.102 Leaflet Base Angle -0.117 0.537 0.283 0.114 Leaflet Length -0.194 -0.308 -0.114 Leaflet Width -0.207 -0.253 0.176 Head Length -0.237 0.150 Flower Number -0.191 -0.214 -0.150 Peduncle Length -0.205 -0.259 Peduncle Diameter -0.230 Involucral Lobe Length -0.168 -0.122 -0.165 0.280 Calyx Circumference -0.181 0.160 -0.264 0.334 Calyx Lobe Length -0.199 -0.104 0.225 Calyx Lobe Width -0.122 -0.396 0.523 Calyx Tube Length -0.183 -0.198 0.284 -0.113 Calyx Lobe Tooth Number -0.126 0.326 0.735 Banner Petal Length -0.238 0.213 Banner Petal Width -0.225 0.150 -0.142

Table 7. (continued) Wing Petal Lamina Length -0.239 0.200 Wing Petal Lamina Width -0.213 0.175 0.166 -0.170 Keel Petal Lamina Length -0.228 0.215 Keel Petal Lamina Width -0.204 0.199 -0.129 Connate Filaments Length -0.236 0.213 Style Length -0.234 0.197

Standard Deviation 3.7062760 1.7376794 1.7376794 1.11479071 1.05369599 Proportion of Variance 0.5087586 0.1118344 0.05854164 0.04602809 0.04112131 Cumulative Proportion 0.5087586 0.6205930 0.67913465 0.72516274 0.76628404

Fig. 2. Principle components analysis projection of all T. variegatum s.l. OTU’s, colored according to potential groups. PC 1 explains 50.9% of variation, PC 2 explains 11.2%.

Fig. 3. Principle components analysis projection of all T. variegatum s.l. OTU’s, colored according to accepted groups. PC 1 explains 50.9% of variation, PC 2 explains 11.2%.

Fig. 4. Principle components analysis 3-dimensional projection of all T. variegatum s.l. OTU’s, colored according to accepted groups. PC 1 explains 50.9% of variation, PC 2 explains 11.2%, and PC 3 explains 5.9%.

Fig. 5. Cluster analysis using average linkage of all OTU’s, colored according to potential groups. Orange is T. polyodon, dark blue T. appendiculatum, red “T. geminiflorum”, green “T. subsalinum”, and cyan T. variegatum. Horizontal black lines below the phenogram denote clusters consisting largely of one taxon, as labeled below the lines. Letters correspond to OTU’s not identified as belonging to the same group as the others in the cluster. The letter “v” stands for T. variegatum, the letter “a” stands for T. appendiculatum, the letter “g” stands for “T. geminiflorum”, and the letter “s” for “T. subsalinum”. The number of letters does not correspond exactly to the number of included OTU’s.

Fig. 6. Cluster analysis using average linkage of all OTU’s, colored according to accepted groups. Orange is T. polyodon, dark blue T. appendiculatum, and cyan T. variegatum. Horizontal black lines below the phenogram denote clusters consisting largely of one taxon, as labeled below the lines. Letters correspond to OTU’s not identified as belonging to the same group as the others in the cluster. The letter “v” stands for T. variegatum and the letter “a” stands for T. appendiculatum. The number of letters does not correspond exactly to the number of included OTU’s.

Fig. 7. Cluster analysis using Ward’s method of all OTU’s, colored according to potential groups. Orange is T. polyodon, dark blue T. appendiculatum, red “T. geminiflorum”, green “T. subsalinum”, and cyan T. variegatum. Horizontal black lines below the phenogram denote clusters consisting largely of one taxon, as labeled below the lines. Letters correspond to OTU’s not identified as belonging to the same group as the others in the cluster. The letter “v” stands for T. variegatum, the letter “p” stands for T. polyodon, the letter “a” stands for T. appendiculatum, the letter “g” stands for “T. geminiflorum”, and the letter “s” for “T. subsalinum”. The number of letters does not correspond exactly to the number of included OTU’s.

Fig 8. Discriminant analysis of hypothetical groups. Operational taxonomic units are colored and labeled according to group membership. The outer circle contains an estimated 50% of the population for each group, while the inner circle represents a 95% confidence limit for the mean. There is considerable overlap between T. variegatum s.s., “T. geminiflorum”, and “T. subsalinum”, all found in the bottom left part of the graph. Biplot rays indicate the loading of characters onto the axes. Calyx lobe tooth number is the most prominent discriminating character for T. polyodon, because this character is used for identification. Discriminant

18 analyses without calyx lobe tooth number still retain T. polyodon quite separate from the other groups, although with the group center at 7 instead of 10 on the y-axis.

Fig. 9. Discriminant analysis of accepted groups. Operational taxonomic units are colored and labeled according to group membership. The outer circle contains an estimated 50% of the population for each group, while the inner circle represents a 95% confidence limit for the

19 mean. No circles intersect at all. This graph does not provide explicit support for the recognition of these groups, since the characters used to diagnose OTU’s were also used in the DA.

Fig. 10. Discriminant analysis of recognized groups, with restricted characters. Operational taxonomic units are colored and labeled according to group membership. The outer circle contains an estimated 50% of the population for each group, while the inner circle represents a 95% confidence limit for the mean. No circles overlap, and important discriminating characters are shown. Relative length of biplot rays indicate discriminating power. “Trifolium geminiflorum” “Trifolium geminiflorum” is characterized by having 1, 2, or 3 flowers and a generally delicate appearance (a shorter height, thinner stems and peduncles, smaller leaflets, and, at times, a reduced involucre). It is only found above 2,000 feet in the Sierra Nevada Mountains and in the southern mountain ranges of San Diego and Riverside Counties, California (Fig. 11). Often, it is located within the tree line on these mountains, a different habitat than what is seen for the other groups of T. variegatum s.l. It is also found near normal-sized T. variegatum, and the two are connected morphologically by a grade of specimens. Figure 12 depicts the overlapping range of “T. geminiflorum” banner petal length compared to T. variegatum s.s. In

20 fact, their averages only differ by ~1 mm. This indicates that floral morphologies do not differ drastically between “T. geminiflorum” and T. variegatum s.s. Differences are apparent in vegetative morphologies (e.g. stipule length, Fig. 13, and leaflet length, Fig. 14). Even though these vegetative characters are used to identify specimens as “T. geminiflorum”, there is considerable overlap in the ranges. The chief character associated with “T. geminiflorum” is flower number (Fig. 15). This trait, however, is not closely correlated with any other characters, and can even vary within a single plant. Numerous specimens were encountered during this study that could easily have been called either T. variegatum or “T. geminiflorum”. Occasionally, “T. geminiflorum” co-occurs with T. variegatum s.s. in the same population. This is documented in Fig. 16 and Fig. 17. In these situations, while there are differences in morphology, the specimens seem to have more in common with each other than they do with other taxa, such as T. appendiculatum. This set of evidence is a good argument against the recognition of “T. geminiflorum” as its own species. Within “T. geminiflorum”, there are a few specimens with a unique character: a significantly reduced involucre. As shown in Fig. 18, the involucre is dissected completely to the peduncle and only has 2 or 3 lobes. Normally, the involucre has at least a 2-3 mm radius of tissue between the lobes and the peduncle, with upwards of 5 lobes. Figure 8 depicts the delicate aspect of this morphology, only found in the Yosemite region of California. This character, however, is not consistent enough to be recognized on its own, since, as Fig. 19 shows, it co-occurs on the same plant with normal involucres. The habitat data suggests that the smaller size characteristic of “T. geminiflorum” is a plastic response to the environment. Moreover, close inspection of the specimens did not indicate a gap in morphology essential for distinguishing between two species. Therefore, “T. geminiflorum” will not to be recognized as a separate species. Representative specimens exhibiting “T. geminiflorum” morphology: R. M. Austen s.n. (NDG, TYPE), Bacigalupi et al. 2587 (CAS), Boyd and Ross 7597 (SD), Dudley 1769 (CAS), Eastwood 317 (CAS), Gander s.n. (SD), Gibbs s.n. (BM), Gregory and Marshall 3253 (SD), Griesel s.n. (CAS), Hansen s.n. (NDG, TYPE), Heller and Kennedy 8873 (ISC), Howell 27046 (CAS), Howell and True 48997 (RENO), Howell and True 48568 (CAS), Moran s.n. (SD), Parry 45 (BM), Parry 451 (ISC), Penalosa 2619 (CAS), C. Quick and A. Quick 66-12 (CAS), Raven 3377 (CAS), Rebman and Hollingsworth 7300 (SD), Rebman et al. 9327 (SD), C.F. Sonne 3 (NDG, TYPE), C.F. Sonne 5 (NDG, TYPE), Spencer s.n. (MU), Spencer s.n. (MU), Spencer s.n. (MU), Spencer 2389 (WIS), Thomas 8744 (CAS), Twisselmann 7349 (CAS), Twisselmann 5538 (CAS), Twisselmann 16180 (CAS), Wiggins 20746 (MU), Williams 73-20-16 (RENO), Wolley-Dod 178 (BM).

Fig. 11. Distribution map of “T. geminiflorum” specimens. They have been collected mostly in the Sierra Nevada Mountains and the mountains in the eastern part of San Diego County.

Fig. 12. Beanplot of banner petal length. “T. geminiflorum” has a slightly smaller average banner petal length than that of T. variegatum s.s. The majority of banner petals of “T. geminiflorum”, besides the smallest few, would fall within the central block of the T. variegatum s.s. range.

Fig. 13. Beanplot of stipule length. “T. geminiflorum” has an average stipule length 2 mm shorter than that of T. variegatum s.s. There not much overlap between the ranges of the two groups of measurements.

Fig. 14. Beanplot of leaflet length. “T. geminiflorum” has a much shorter average leaflet length than that of T. variegatum s.s.

Fig. 15. Beanplot of flower number. “T. geminiflorum” has fewer flowers on average than T. variegatum s.s.

Fig. 16. Specimen with two plants, one exhibiting “T. geminiflorum” morphology and the other “T. variegatum” morphology (Botti 154 (CAS)). Despite overall plant size and flower number, the two plants share many common characters, including the length of corolla characters.

Fig. 17. Specimen exhibiting “T. geminiflorum” morphology and “T. variegatum” morphology (Keck 1237 (CAS)).

Fig. 18. “Trifolium geminiflorum” morph. This is one of the specimens that would have been called T. pusillum by Greene (Eastwood 317 (CAS)). Close up view of reduced involucre (arrow) is shown.

a b

Fig. 19. “Trifolium geminiflorum” morph. a. Specimen exhibits (arrow) reduced involucre and “normal” involucre on the same plant (Raven 3377 (CAS)). b. Drawing depicts a peduncle and involucre of a “normal” inflorescence on the left and a peduncle, involucre, and calyx (cut off) of a reduced inflorescence on the right. Both inflorescences are on the same plant.

“Trifolium subsalinum” and “Trifolium ultramontanum” “T. subsalinum” is characterized by a zig-zag pattern in the upper stem when pressed (Figs. 20, 21). Hypothetically, this pattern could be due to a thickening of the upper stem. Most specimens come from the Great Basin region (Fig. 22), where stem thickness might represent a xerophytic adaptation, but specimens were also found from Solano County and San Diego County, CA. At first it seemed that the California specimens differed from the Great basin specimens in some sort of floral relationship (Fig. 23), putatively the ratio of calyx length to corolla length. However, this did not hold up in measurements comparing calyx to corolla length (Figs. 24 and 25), and it was determined that this difference was only superficial. In fact, perceived differences were chiefly due to a lack of pigmentation on the calyx lobes of the Great Basin specimens. Likewise, perceived stem thickness could not be linked to stem diameter (Fig. 26). Stem rigidity also appears to be a grade, with many specimens exhibiting a morphology intermediate between “T. subsalinum” and the rest of T. variegatum s.l. Figures 27 and 28 depict specimens in this intermediate range. The previously mentioned calyx pigmentation was investigated further, and certain specimens also bearing blunt leaflet apices but lacking zig-zagged stems were called “T. ultramontanum”. Figures 29 and 30 depict its habit. The difference between this group and T. variegatum s.s. is most apparent in the state of Washington, where it is easy to determine if a specimen came from east or west of the Cascades. However, pigmentation varies throughout the entire range of T. variegatum s.l. (Fig. 31), as does leaflet bluntness. Specimens at higher elevations in the Great Basin were more similar to T. variegatum s.s. than “T. ultramontanum”, suggesting that the morphology of the latter may be a plastic response to drier lowland conditions. “Trifolium ultramontanum” was not assessed using multivariate analyses. Neither “T. subsalinum” nor “T. ultramontanum” are recognized here as species because of lack of support in measurements and instability in their defining characters. Ultimately, no gap in morphology was found in order to definitively establish these groups as separate species. Representative specimens exhibiting “T. subsalinum” morphology: Allen 414 (RENO), Archer 7229 (RENO), Clark 255 (CAS), de Nevers 8173 (CAS), E.L. Greene s.n. (NDG, TYPE), E.L. Greene s.n. (NDG, TYPE), Heller 9129 (ISC), Howard s.n. (BRY), Jones 1046 (BM), Kennedy 1944 (WIS, WIS), Leonard s.n. (MU), Naczi and Thieret 3399 (MU), Scofield 226 (WIS), Tiehm 3660 (RENO), Tiehm and Williams 1109 (RENO), Williams and Tiehm 83-97-1 (RENO), Williams et al. 83-84-6 (BRY, RENO), Williams et al. 76-90-14 (RENO), Wosley s.n. (WIS). Representative specimens exhibiting “T. ultramontanum” morphology: Archer 6526 (RENO), DeDecker 6101 (MU), Fiker 855 (CAS), E.L. Greene s.n. (NDG, TYPE), E.L. Greene s.n. (NDG, TYPE), Hitchcock 21590 (CAS), Hoffmann s.n. (CAS), Kirkwood 1756 (CAS (2 sheets), ISC, MU), Mancuso 1708 (SRP), Sandberg 401 (BM), Thompson 10 (CAS), Tiehm 14991 (MU), Weldert and Boner 151 (CAS).

Fig. 20. “Trifolium subsalinum” morph (Williams 83-84-6 (RENO)).

Fig. 21. “Trifolium subsalinum” morph. Plant is from Sonoma County, CA (de Nevers 8173 (CAS)), and is exhibiting zig-zag stem pattern.

Fig. 22. Distribution map of “T. subsalinum” specimens. An additional specimen, labeled “Antimony Springs” (Wosley s.n., WIS), is likely from eastern Oregon.

a b Fig. 23. “Trifolium subsalinum” morphs compared. a. Specimen is from Sonoma County, CA (de Nevers 8173 (CAS)). Picture depicts a close-up of the purple-pigmented calyx lobes. b. Picture depicts a close-up of the calyx lobes lacking purple pigment (Williams 83-84-6 (RENO)).

Fig. 24. Beanplot of the difference between banner petal length and calyx length. Despite the banner petal superficially exceeding the calyx more in “T. subsalinum” than it does in T. variegatum s.s., the difference between these two measurements does not change much.

Fig. 25. Beanplot of banner petal length to calyx length ratio. Like the difference as shown in Fig. 30, the ratio between banner petal length and calyx length does not appear to change much when comparing “T. subsalinum” to T. variegatum s.s.

Fig. 26. Beanplot of stem diameter. Stem diameter, at least where measured (usually below the third or fourth most distal node) does not increase much in “T. subsalinum”.

Fig. 27. Specimen from Arizona exhibiting intermediate morphology between “T. subsalinum” and T. variegatum s.s. (Buegge 873 (ASU)). Stems are not uniformly rigid.

Fig. 28. Nevada specimen exhibiting intermediate morphology between “T. subsalinum” and T. variegatum s.s. (Kennedy 3056 (CAS)).

Fig. 29. “Trifolium ultramontanum” morph. Specimen exhibiting blunt leaflet apices and calyx lobes without purple pigment (Tiehm 14991 (RENO)).

Fig. 30. “Trifolium ultramontanum” morph. Close-up with calyx lobes lacking purple pigmentation (Tiehm 14991 (RENO)).

Fig. 31. Map showing the distribution of specimens with pigmented or unpigmented calyx lobes within the Trifolium variegatum complex. Groups included are T. variegatum s.s., “T. subsalinum” and “T. ultramontanum”. Trifolium polyodon Found solely in Monterey County, CA (Fig. 32), T. polyodon is often recognized a distinct species by other authors (e.g., Zohary and Heller, 1984; Vincent, 2012). It is demarcated from T. variegatum by having consistently-toothed calyx lobes (Fig 33). Each calyx lobe usually bears two or more teeth. Trifolium variegatum rarely has calyx lobe teeth (75% of T. variegatum specimens measured lacked teeth, 13% had only one per calyx, and the remaining 12% range from 2 to 8 teeth per calyx), and T. appendiculatum even more rarely. Trifolium polyodon is generally robust and has wide leaflets (Fig. 34), a character not common in T. variegatum.

Trifolium polyodon also has longer stipules than either T. variegatum or T. appendiculatum (Fig. 13). Other specimens of T. variegatum s.l. with up to 8 calyx lobe teeth per calyx have been collected outside of Monterey County, California (Fig. 35). The erratic distribution of this character warranted further investigation. If it could be shown that the calyces in non- T. polyodon plants were homologous to those in T. polyodon, T. polyodon ought not be recognized as a separate species. Trifolium variegatum specimens with many calyx lobe teeth can be distinguished from T. polyodon by having fewer and shorter teeth, as well as by being smaller both in vegetative and reproductive characters. Figures 36 and 37 depict how the five T. variegatum specimens with the most calyx lobe teeth differ from T. polyodon in terms of both tooth number per calyx and tooth length. Despite the pseudoreplication, these measurements offer a compelling case that the calyx lobe teeth present on T. polyodon are of a different origin than those on these specimens of T. variegatum. Tooth length was not used in establishing the identity of these species, nor in any statistical analyses. Figures 38 and 39 provide pictures of Neese 9097 (BRY), the most toothed specimen of T. variegatum. Figure 40, of Yadon s.n. (MU), a specimen of T. polyodon, depicts how this species is comparatively larger overall. Since PC 5 has a significant loading from calyx lobe tooth number, T. polyodon is clearly separated from T. appendiculatum in a plot with PC 1 and PC 5 as axes (Fig. 41). This group is well-supported as a species given its unique and consistent morphology and distribution suggesting a single origin.

Fig. 32. Distribution map of T. polyodon specimens.

Fig. 33. Drawing of dissected calyx of a specimen of T. polyodon (Heller 6759 (ISC)).

Fig. 34. Beanplot of leaflet width. Trifolium polyodon has, on average, wider leaflets than both T. variegatum and T. appendiculatum.

Fig. 35. Distribution map of specimens with more than 5 calyx lobe teeth per calyx, all belonging to T. variegatum.

Fig. 36. Beanplot of tooth number per calyx. For T. polyodon, 10 calyces from 5 plants from 4 specimens were used to count the number of teeth. For T. variegatum, 15 calyces from 12 plants from 5 specimens, representing those specimens with the highest number of teeth (most specimens of T. variegatum have none), were used to count the number of teeth. One calyx of each species was shown to have 8 teeth per calyx. On other calyces on each of these specimens, the number of calyces returned closer to its species average.

Fig. 37. Beanplot of tooth length. All teeth (n=109) present on 10 calyces from 5 plants from 4 specimens were measured for their length in T. polyodon specimens. All teeth (n=66) present on 15 calyces from 12 plants from 5 specimens were measured for their length in T. variegatum specimens.

Fig. 38. Specimen of T. variegatum with many calyx lobe teeth (Neese 9097 (BRY)). Specimen is not T. polyodon.

a b

Fig. 39. Specimen of T. variegatum with many calyx lobe teeth (Neese 9097 (BRY)). Specimen is not T. polyodon. a. A photograph shows a close-up of the flowers. b. A drawing exhibits this calyx with 6 calyx lobe teeth.

Fig. 40. Cultivated specimen of T. polyodon (Yadon s.n. (MU)).

Fig. 41. PCA projection of all T. variegatum s.l. OTU’s, colored according to accepted groups. Principle component 1 explains 50.8% of variation, principle component 5 explains 4.1%. On PC 5, the character with the highest loading is calyx lobe tooth number. Trifolium appendiculatum Trifolium appendiculatum has not been recognized as a species separate from T. variegatum in many recent floristic works (e.g., the Jepson Manual, Hickman, 1993, and Baldwin et al., 2012). These publications follow Isely (1993), who was the first person not to recognize T. appendiculatum as a separate species. Before his work, however, T. appendiculatum was included in the original Jepson Manual (Jepson, 1925) and Munz’s California flora (Munz and Keck, 1959). It is found in cismontane California and also in southern Oregon (Fig. 42). It is chiefly distinguished from T. variegatum by having larger reproductive parts, mainly those of the corolla. Figures 43, 44, and 45 all depict T. appendiculatum’s increased lengths of corolla characters, as does Fig. 46 for connate stamen length. All of these characters are highly correlated with each other throughout the T. variegatum complex. Trifolium appendiculatum often has larger vegetative organs, but this is not always the case, especially early in the growing season. Trifolium appendiculatum also has, on average, longer peduncles (Fig. 47). Seed length was determined to be a useful diagnostic character for T. appendiculatum (Fig. 48). This character was identified after specimens were assigned to species. Seeds taken from mature fruits (those that have dehisced and often bear purple speckles) are 1.9 to 2.8 mm long in T. appendiculatum (seeds n=18; plants n=10), while those of T. variegatum are 1.1 to 1.6 mm long (seeds n=12; plants n=6). Seeds were sampled from across the morphological and geographic extent of both taxa. Seeds and floral organs of T. appendiculatum and T. variegatum

45 are depicted in Fig. 49. If T. variegatum seeds can be described as mittens, T. appendiculatum seeds are more like boxing gloves. Some plants of T. appendiculatum have beaked keel petals (Fig. 50). There is a minute elongation of tissue at the apex of the connate petal laminas. This morphology is consistently found within the same plant, but there are no correlated morphological characters. Figure 51 is a picture of a beaked T. appendiculatum and Fig. 52 is a picture of one without a beak. Beaks are not visible, but these figures depict that there are no macromorphological differences associated with having or not having a beak. Both specimens are from the same county. Trifolium appendiculatum flowers earlier than T. variegatum, although this is probably due to habitat differences. In Table 8, it is clear that the majority of T. appendiculatum specimens (n=57) were collected earlier than the majority of T. variegatum specimens (n=226). In the later months, however, specimens of T. variegatum are predominantly from states in the Great Basin region, which has a higher elevation and therefore warms later than areas further west. Based on the results of multivariate analyses, consistent morphological characters different from those of T. variegatum, and a well-defined distributional pattern, T. appendiculatum is here recognized as a species. It is sympatric with T. variegatum, and collection data does not indicate any differences in habitat preference. Divergent floral morphologies could indicate pollination barriers.

Table 8. Proportion of specimens collected by month.

Month Proportion of T. Proportion of T. Proportion of T. variegatum in Nevada, appendiculatum collected variegatum collected Arizona, Idaho, Montana, Utah March 0.087719 0.004425 0 April 0.350877 0.088496 0.05 May 0.473684 0.230088 0.057692308 June 0.087719 0.300885 0.411764706 July 0 0.287611 0.529411765 August 0 0.061947 0.785714286 September 0 0.022124 0.4 October 0 0.004425 0

Fig. 42. Distribution map of T. appendiculatum specimens.

Fig. 43. Beanplot of banner petal length. The banner petal length of T. appendiculatum is longer than that of T. variegatum by 4 mm.

Fig. 44. Beanplot of wing petal length. The average wing petal length of T. appendiculatum exceeds that of T. variegatum by 3 mm.

Fig. 45. Beanplot of keel petal length. The average keel petal length of T. appendiculatum exceeds that of T. variegatum by almost 3 mm.

Fig. 46. Beanplot of connate filaments length. The average connate stamens length of T. appendiculatum exceeds that of T. variegatum by 2 mm.

Fig. 47. Beanplot of peduncle length. It is, on average, 2 cm longer in T. appendiculatum than in T. variegatum.

Fig. 48. Beanplot of seed length. For T. appendiculatum, 10 plants were sampled for a total of 18 seeds. Sampled plants included those from 3 counties and ranged from small to large. For T. variegatum, 6 plants for a total of 12 seeds were sampled. Sampled plants were from British Columbia, Idaho, and California with morphologies ranging from large to minute.

a b

Fig. 49. Comparisons of floral organs and seeds between T. appendiculatum and T. variegatum. a. Floral organs and a seed are drawn from a specimen of T. appendiculatum (Thorne & Evertt 36989 (ISC)). Organs, from top left to bottom right are: calyx, banner petal, wing petal, keel petal, connate stamens, pistil, and seed. b. Floral organs and a seed are drawn from a specimen

50 of T. variegatum (Rebman 15030 (SD)). Organs, from top left, to top right, to bottom, are: calyx, banner petal with free stamen attached, wing petals, keel petals, pistil, and seed.

Fig. 50. Beak of T. appendiculatum keel petals. Flower taken from horticulturally-grown plant. Plant was grown from seed provided by the USDA National Plant Germplasm System (PI 593317). Keel petals were left intact, and the top wing petal was removed; the other wing petal was not removed (on the far side of the keel petals).

Fig. 51. Trifolium appendiculatum specimen with beaked keels, from Tulare County, CA (Straw 2620 (MU)).

Fig. 52. Trifolium appendiculatum specimen without beaked keels, from Tulare County, CA (Thorne 36989 (ISC)).

Further Discussion Based on the evidence discussed in the preceding section, three taxa are here recognized. These taxa are T. polyodon, T. appendiculatum, and T. variegatum, the last of which includes as synonyms “T. geminiflorum”, “T. subsalinum”, and “T. ultramontanum”. These species are presented in more detail in the following section. The taxonomic decisions made in this study are consistent with what was previously mentioned regarding patterns of diversity in western North America. Almost two of the three species are California endemics. Trifolium polyodon is a true endemic. Trifolium appendiculatum is found throughout cismontane California, and its one collection outside that state (Josephine County, OR) is still within the California Floristic Province. Annual dicots are an important constituent of all California endemics (Raven and Axelrod, 1978). According to the most recently published molecular phylogeny of Trifolium (Ellison et al., 2006) and assuming that T. appendiculatum would fall out near T. variegatum, T. variegatum is a part of a clade of four annual species (the fourth is T. buckwestiorum), three of which are endemic to California. Given this viewpoint, it is T. variegatum’s widespread distribution, and not the narrow distributions of T. polyodon and T. appendiculatum, that calls for explanation. Assuming T. variegatum originated in California, a likely proposition based on the distribution of its hypothesized relatives, it had to be dispersed a great distance and cross the Sierra Nevada Mountains in order to reach its present distribution. It is unclear if there are any traits that indicate an increase in dispersal ability, but, during the course of this study, it was noticed that T. variegatum had a larger proportion of inflorescences in fruit than the other species. Either these plants are producing fruits after flowering more quickly, or they produce more inflorescences overall. In a similar manner, T. variegatum’s phenotypic plasticity, such a confounding factor for taxonomic studies, might also help explain its distribution. The Sierra Nevada Mountains and Great Basin Region have very different climates, and the ability to change habit could be advantageous. Thus, this species is certainly worth further investigation outside of a taxonomic context.

Taxonomic Treatment Key to species 1. Calyx lobes usually many-toothed, averaging (1.6) 1.8 - 2.8 teeth per lobe, teeth 0.8 mm long on average, distal stipules 8 - 10 mm long; Monterey County, CA ...... 1. T. polyodon 1. Calyx lobes usually entire, if toothed, with 0 - 1 (1.6) teeth per lobe, teeth 0.3 mm long on average, distal stipules 2 - 8 (10) mm long; western United States, southwestern British Columbia, Canada, and northern Baja California, Mexico ...... 2 2. Banner petals (6) 8 - 16 mm long, exerted beyond tips of calyx lobes (1.5) 3-7 (8.4) mm, mature seeds 1.9 - 2.8 mm long, keel petals beaked or not; California and southern Oregon ...... 2. T. appendiculatum 2. Banner petals (3) 4 - 10 mm long, included in or exerted beyond tips of calyx lobes (0.1) 1 - 4 (5.7) mm, mature seeds 1.1 - 1.6 mm long, keel petals not beaked; British Columbia, Canada, south through Washington, Oregon, Idaho, Montana, California, Nevada, Utah, and Arizona, USA, to Baja California, Mexico ...... 3. T. variegatum

1. Trifolium polyodon Greene, Figs. 40 and 53. Trifolium polyodon Greene, Pittonia 3: 215, 1897. Trifolium tridentatum var. polyodon (Greene) Jeps., Fl. Calif. 2: 292. 1936. TYPE: USA, California: Monterey County, Pacific Grove, 27 May 1895, E.L. Greene s.n. (holotype: NDG 67254!; isotype: NDG 67260 – electronic image!)

Plants annual, entirely glabrous; roots a taproot, woody, nodulated; stems ascending to decumbent, 10-52 cm in length, 1-2.3 mm in diameter; leaves trifoliate, dimorphic with gradual transition between proximal and distal leaves; stipules adnate to petiole, free portion laciniate, usually with one tooth longer than the rest; proximal stipules browning with age, each lobe rectangular proximally and triangular distally, 6-14×2-3 mm, margins +/- entire proximally, laciniate distally, apices acute with 1-2 lobes, the longest lobe 2-3.1 mm; distal stipules quarter- circled to ovate, recurved, 4-7x5-6 mm, margins laciniate, with 0-3 apical lobes, the longest lobe 2.5-3.2 mm; petioles 9-55×0.2-0.9 mm; petiolules 0.4-0.7x≤0.4 mm; proximal leaflets obovate- obcordate, bases cuneate, margins serrate, apices emarginate-mucronate, terminal leaflets 4- 10×4-6 mm; distal leaflets elliptic-obovate, bases cuneate, margins serrate, apices rounded, terminal leaflets 13-18.7x6.4-9.2 mm; peduncles 13-54×0.3-0.8 mm; inflorescences axillary, racemose-umbellate, involucrate, 8.2-16×10-17 mm; involucre flat, laciniate, 4.3-6.7 mm radius, lobes indistinguishable from teeth, longest tooth 1.7-2.7 mm; flowers 12-22, in 1-4 whorls; pedicels 0.5-1×<0.5 mm; calyces 5-7 mm long, tubes 2.3-2.9 mm in circumference, 1.9- 2.4 mm long, lobes awl-like to shouldered, tips purple, 3.1-4.6 mm long, apices acute, each bearing 0-4 teeth 0.1-3.1 mm long; petals pink-purple with paler to white tips; banner petals obovate, apices emarginate, 8.3-8.8×2.1-3.2 mm, exerted beyond the tips of calyces 1.5-3.6 mm; wing petals 7.5-8.4 mm long, asymmetrically clawed, claws 4-4.5×≤0.1 mm, laminae elongate-elliptical, 3.2-4.4x1-1.4 mm, bases shortly auriculate, auricles rounded, petal margins entire, apices rounded; keel petals 6.1-7.1 mm long, asymmetrically clawed, claws 4.4-4.8×≤0.1 mm, laminae ovate, 1.7-2.4x1-1.3 mm, base rounded, margins entire, apices rounded; stamens diadelphous, connate filaments 6-6.6×≤0.5 mm; pistils sessile, ovaries 2.3-2.5×0.6-0.7 mm, styles 3.6-4.6×≤0.1 mm, ovules 2; legumes laterally compressed, 4×2 mm; seeds 2, globular-- mitten-shaped, blackish-brown, dark-speckled, 1.9×1.1 mm. 2n=?

Representative Specimens USA. California: Monterey County. Pacific Grove, 21 May 1903, Heller 6759 (ISC); Pacific Grove, 1 May 1909, Heller 28022 (WIS); Pacific Grove, 9 May 1909, Heller 6707 (WIS); horticulturally grown specimens, originally from Indian Village, 20 June 1994, Yadon s.n. (MU); Indian Village near Bird Rock, off 17-Mile Drive, 9 April 1998, Vincent et al. 8195 (MU).

Distribution and Habitat Specimens have been collected from 2 localities, Pacific Grove (the type locality) and Indian Village, which is approximately 3 miles away (Fig. 51). It has been found in wet grassy

56 locations. All of these collections were made at ~122 m in elevation. It has been collected, flowering, in April, May and June.

Comments Duane Isely (1998) suggested that T. polyodon is the result of hybridization between T. variegatum and T. willdenovii because the latter often has many-toothed calyx lobes. Accordingly, he lumped T. polyodon within T. variegatum, albeit as its own “phase”. In his Flora of California (Jepson, 1936) Jepson treated T. polyodon as a variety of T. tridentatum, a current synonym for T. willdenovii (Isely, 1993). This is odd because in 1925 Jepson’s Manual of the Flowering Plants of California recognized T. polyodon as a separate species. In the Flora of California (Jepson, 1936), Jepson twice mentions how T. tridentatum var. polyodon (Greene) Jeps. is more like T. variegatum than T. tridentatum. Isely in 1998 and 1993 and Jepson in 1936 are the only instances where T. polyodon has not been treated a distinct species. Based on the existence of T. variegatum with numerous calyx lobe teeth, it does not seem that the hybridization hypothesis is necessary. These specimens indicate that T. variegatum has the genetic mechanism or is only a few genetic switches away from the ability to form calyx lobe teeth. Given this interpretation, and the different nature of T. polyodon calyx lobe teeth from those of T. variegatum, it is reasonable to assume that T. polyodon’s distinctive teeth are from an independent evolutionary origin than those of T. variegatum. Moreover, the calyx lobe teeth found on T. willdenovii are smaller than those of T. polyodon, and T. polyodon has no other characters suggestive of T. willdenovii. Trifolium willdenovii is recognizable by its narrow, at times linear, leaflets, whereas T. polyodon, as well as T. variegatum and T. appendiculatum, has wider leaflets. In Ellison et al.’s (2006) molecular phylogenetic analysis, T. variegatum comes out as sister to T. polyodon in a clade consisting of five species. Trifolium willdenovii is in a different clade consisting of eight species, though all three fall within a clade of involucrate species. One specimen of T. variegatum with many calyx lobe teeth, Neese 9097 (BRY), was collected in Nye County, Nevada. In this study, no specimens of T. willdenovii (or its synonym, T. tridentatum) have been seen from Nevada, and neither Zohary and Heller (1984) nor Tidestrom (1925) list Nevada as part of its range. Neese 9097 (BRY) cannot be of hybrid origin, since one of the parent species is not present. Independent mutational events seem to be more parsimonious explanation than hybridization events.

Fig. 53. Distribution map of T. polyodon specimens.

2. Trifolium appendiculatum Lojac., Figs. 51 and 54. Trifolium appendiculatum Lojac. Nuovo Giorn. Bot. Ital. 15: 181, 1883. TYPE: USA, California: Coast field, Lemmon s.n. (not located at PAL or elsewhere) NEOTYPE (to be designated): USA, California: Contra Costa County, St. Mary’s College, 14 May 1933, J.T. Howell 11227 (neotype: CAS!; isoneotype: WIS!). Trifolium variegatum var. major Lojac. Nuovo Giorn. Bot. Ital. 15: 183, 1883. Trifolium variegatum var. melananthum fo. major (Lojac.) McDermott, Ill. Key Amer. Trifolium 76. 1910. TYPE: USA, California: Lemmon s.n. Inot located at PAL or elsewhere) NEOTYPE (to be designated): USA, California: Tulare County, hills east of Yokohl Valley, 9 May 1969, E.C. Twisselman 15246 (neotype: CAS!; isoneotypes: SBB, RSA). Trifolium trilobatum Jeps., Bull. Torrey Bot. Club 18: 322. 1891. Trifolium variegatum var. trilobatum (Jeps.) Jeps. in McDermott, Ill. Key Amer. Trifolium 73. 1910. TYPE: USA, California: Sutter County, base of South Peak, 20 April 1891, W.L. Jepson 14854 (holotype: JEPS - electronic image!) Trifolium rostratum Greene, Proc. Acad. Nat. Sci. Philad. 47: 547. 1895. Trifolium appendiculatum fo. rostratum (Greene) McDermott, Ill. Key Amer. Trifolium 92. 1910. Trifolium appendiculatum var. rostratum (Greene) Jeps., Man. Fl. Pl. Calif. 539. 1925. Trifolium variegatum var. rostratum (Greene) C.L. Hitch. Vasc. Pl. Pacif. N.W. 3: 370. 1961. TYPE: USA, California: Alameda County, Oakland, Lake Merritt, 1889, V.K. Chestnut s.n. (holotype: NDG!). Trifolium morleyanum Greene, Erythea 3: 47. 1895. Trifolium variegatum var. melananthum fo. morleyanum (Greene) McDermott, Ill. Key Amer. Trifolium 76. 1910. TYPE: USA, California: Modoc County, Morley’s Station, 1894, Baker & Nutting s.n. (holotype: NDG!). Trifolium calophyllum Greene, Pittonia 3: 213. 1897. TYPE: USA, California: Alameda County, Berkeley, Botanic Garden, 1895, E.L. Greene s.n. (lectotype: NDG!, NDG67170 (selected by Gillett, 1966); syntypes (same locality and date): NDG 67171!, NDG 67172!, NDG 67214!, NDG 67176!). Trifolium phaeocephalum Greene, Pittonia 3: 216. 1897. Trifolium variegatum var. pauciflorum fo. phaeocephalum (Greene) McDermott, Ill. Key Amer. Trifolium 78. 1910. TYPE: USA, California: Butte County, east of Chico, May 1883, R.M. Austin s.n. (lectotype: NDG! (selected by Gillett, 1966)). Trifolium splendens A.Heller, Muhlenbergia 1: 115, 1905. TYPE: USA, California: Monterey County, Pacific Grove, 7 May 1903, A. Heller 6691 (holotype: BKL; isotypes: AC - electronic image!, BM!, CAS - electronic image! (3 sheets), F - electronic image!, G - electronic image!, ISU!, K, MICH - electronic image!, MSC - electronic image!, NY - electronic image! (2 sheets), P - electronic image!, US - electronic image!, WIS!)

Plants annual, entirely glabrous; roots a taproot, woody, nodulated; stems ascending, 7-82 cm in length, 0.5-3 mm in diameter; leaves trifoliate, dimorphic with gradual transition between proximal and distal leaves; stipules adnate to petiole, free portion laciniate, usually with one tooth longer than the rest; proximal stipules browning with age, each lobe rectangular proximally and triangular distally, 7-13×2-5 mm, margins +/- entire proximally, laciniate distally,

59 apices acute with 1-2 lobes, the longest lobe 1.5-3 mm; distal stipules quarter-circled to ovate, recurved, 3.4-10.3x5-6 mm, margins laciniate, with 0-many apical lobes, the longest lobe 0.8- 4mm; petioles 2-73×0.2-1 mm; petiolules 0.3-0.9x≤0.5 mm; proximal leaflets obovate- obcordate, bases cuneate, margins serrate, apices emarginate-mucronate, terminal leaflets 2.9- 7×1.5-5 mm; distal leaflets elliptic-obovate, bases cuneate, margins serrate, apices rounded and mucronate, terminal leaflets 5.8-25.9x2.8-10.9 mm; peduncles 17-73×0.3-1.3 mm; inflorescences axillary, racemose-umbellate, involucrate, 7.3-21×8-31 mm; involucre flat, laciniate, 3.2-6.5 mm radius, 4-7 lobes, lobes at time indistinguishable from teeth, longest tooth 1.1-3 mm; flowers 4-25, in 1-4 whorls; pedicels 0.3-0.5×<0.5 mm; calyces 3.5-7.4 mm long, tubes 2.2-4.6 mm in circumference, 1.5-3.3 mm long, lobes awl-like to shouldered, tips and shoulders purple, 2.2-5.2 mm long, apices acute, usually toothless, teeth < 1 mm; petals deep purple, white-tipped; banner petals obovate, apices emarginate, 6.3-15.8×1.5-4 mm, exerted beyond the tips of calyces 1.5-8.4 mm; wing petals 5.3-14.2 mm long, asymmetrically clawed, claws 2.9-7×≤0.1 mm, laminae elongate-elliptical, 3-7.3x0.8-1.9 mm, bases shortly auriculate, auricles rounded, petal margins entire, apices rounded; keel petals 4.6-10.6 mm long, asymmetrically clawed, claws 2.8-6.7×≤0.1 mm, laminae ovate, 1.8-3.9x0.8-1.8 mm, base rounded, margins entire, apices rounded to beaked, beaks ≤ 1 mm ; stamens diadelphous, connate filaments 3.9-9.8×≤0.5 mm; pistils sessile, ovaries 1.4-4.1×0.3-1.1 mm, styles 2.4- 6.2×≤0.3 mm, ovules 2; legumes laterally compressed, 3.6-3.8×2-2.4 mm; seeds 2, globular-- mitten-shaped, blackish-brown, black-speckled, 1.9-2.8×1.2-2 mm. 2n=?

Representative Specimens USA California: Butte County, hills near Big Chico Creek east of Chico, 6 April 1913, Heller 10710 (CAS); Kern County, high slope 1 mile east of Poso Creek Narrows, 17 May 1963, Twisselmann 8251 (CAS); Monterey County, west of S. Fran., June 1876, Palmer 71 (GH); San Diego County, Cuyamaca Rancho State Park. Approx. 04 mile NE of intersection of State hwy., 26 June 2005, Hendrickson 1084 (SD); San Joaquin County, eastern edge of Wallace, 2 May 1959, Smith 1063 (CAS); Sutter County, found growing on damp soil along the edge of West Butte Road, about 1/8 mile north of the Sacramento Outing Club, Sutter Buttes, 19 April 1984, Ahart 4566 (CAS). Oregon: Josephine County, Grant’s Pass, 16 May 1910, Heller 10030 (CAS), Heller 10031 (ISC, CAS (2 sheets))

Distribution and Habitat Trifolium appendiculatum is found throughout California, though never east of the Sierra Nevada Mountains, and into southern Oregon (Fig. 54). It has not often been found much in the central valley of California, although this is likely due to habitat loss. It is often collected in open fields, meadows, ditches, marshes, swamps, oak woodlands, and digger pine woodlands, in rocky, sandy, or loamy soil. Its elevation ranges from 0 to 1400 m. It flowers in March, April, May, and June (Table 8).

In Hitchcock et al.’s Vascular plants of the Pacific Northwest (1961), it is reported that T. variegatum var. rostratum (Greene) C.L. Hitch. was found on Vancouver Island, British Columbia. Since no voucher specimens were cited, it is unclear what species was actually present. No specimens of T. appendiculatum have been seen from Vancouver Island during this study.

Comments The types of T. melananthum and T. variegatum var. β, though the latter is not a validly published name, are among the larger specimens of T. variegatum, but are not included within T. appendiculatum because their flowers are too short.

Fig. 54. Distribution map of T. appendiculatum specimens.

3. Trifolium variegatum Nutt., Figs. 55 and 56. Trifolium variegatum Nuttall in Torrey & A. Gray, Fl. N. Amer. 1: 317. 1838. TYPE: USA, Oregon: springy places near the mouth of the Wahlamet, T. Nuttall s.n. (holotype: NY - electronic image!; isotypes: BM!, GH, K). Trifolium spinulosum var. triste Torrey and A. Gray, Fl. N. Amer. 1: 318, 1838. TYPE: USA, California: Santa Barbara County, Santa Barbara, T. Nuttall s.n. (holotype: BM!). Trifolium melananthum Hook. & Arn., Bot. Beechey Voy. 331. 1838. Trifolium variegatum var. melananthum (Hook. & Arn.) Greene, Fl. Francisc. 1: 29. 1891. Trifolium tridentatum var. melananthum (Hook. & Arn.) S. Watson, Proc. Amer. Acad. Arts 11: 130. 1876. TYPE: USA, California: D. Douglas s.n. (lectotype (to be designated): K 001051289 – electronic image!; isolectotype: BM 010892 - photocopy!). Trifolium dianthum Greene, Pittonia 3: 217. 1897. TYPE: Canada, British Columbia: Vancouver Island, vicinity of Victoria, 13 May 1893, J. Macoun 94 (holotype: NDG 67320!; isotype: MO!). Trifolium geminiflorum Greene, Pittonia 3: 216. 1897. Trifolium variegatum var. geminiflorum (Greene) Vincent, Madroño 56: 208. 2009. TYPE: USA, California: Shasta County, Lassen’s Peak, August 1879, R. M. Austen s.n. (lectotype (to be designated): NDG 67301A!; syntype: USA, California: Nevada County, Donner Lake, July 1890, C.F. Sonne 3 (NDG 67301B!); syntype: USA, California: Nevada County, Donner Lake, August 1890, C.F. Sonne 5 (NDG 67301C!); syntype: USA, California, Amador County, 1891, Hansen s.n. (NDG 67208!). Trifolium pusillum Greene, Pittonia 3: 217. 1897. TYPE: USA, California: Yosemite Valley, 1881, C.C. Parry 45 (holotype: NDG!; isotype: MO!). Trifolium subsalinum Greene, Pittonia 3: 219. 1897. TYPE: USA, Nevada: Eureka County, Palisade, 24 July 1893, E.L. Greene s.n. (lectotype: NDG! (selected by Gillett, 1966)); syntype: USA, Nevada: Eureka County, Palisade, 24 August 1896, E.L. Greene s.n. (NDG!). Trifolium ultramontanum Greene, Pittonia 3: 218. 1897. TYPE: USA, Nevada: Elko County, Holborn, 16 July 1896, E.L. Greene s.n. (lectotype: NDG 67201! (selected by Gillett, 1966); isolectotype: NDG 67202!, NDG 67204!); syntype: USA, Nevada: Star Valley, 20 July 1896, E.L. Greene s.n. (NDG!).

Plants annual, entirely glabrous; roots a taproot, woody, nodulated; stems prostrate to erect, 2.5-48+ cm in length, 0.1-2.3 mm in diameter; leaves trifoliate, dimorphic, with gradual or abrupt transition between proximal and distal leaves; stipules adnate to petiole, free portion laciniate, usually with one tooth longer than the rest; proximal stipules browning with age, each lobe rectangular proximally and triangular distally, 2.4-10×0.5-2.1 mm, margins +/- entire proximally, laciniate distally, apices acute with 1-2 lobes, the longest lobe 1-2.5 mm; distal stipules quarter-circled to ovate, recurved or not, 1.7-9.3x0.5-2.5 mm, margins laciniate, with 0- 3 apical lobes, the longest lobe 0.6-4.5 mm; petioles 1-95×0.1-1.1 mm; petiolules 0.1-0.8x≤0.5 mm; proximal leaflets obovate-obcordate-cuneate, bases cuneate, margins serrate, apices

62 emarginate-mucronate, terminal leaflets 1.9-5×0.9-2.1 mm; distal leaflets elliptic-obovate, bases cuneate, margins serrate, apices rounded and mucronate, terminal leaflets 2-29x1-11.4 mm; peduncles 4-53×0.1-0.7 mm; inflorescences axillary, racemose-umbellate, involucrate, 4.6- 12.6×0.8-10 mm; involucre flat, laciniate, 1.5-6 mm radius, 2-7 lobes, at times indistinguishable from teeth, longest tooth 0.6-3 mm; flowers 1-25, in 1-4 whorls; pedicels 0.1-0.5×<0.1 mm; calyces 2.1-6.8 mm long, tubes 1.4-4.1 mm in circumference, 1.9-2.4 mm long, lobes awl-like to shouldered, tips purple or stramineus, 1.5-4.8 mm long, apices acute, occasionally bearing up to 3 teeth per lobe, teeth < 0.6 mm long; petals purple, usually white-tipped; banner petals obovate, apices emarginate, 3.2-9.7×0.6-2.6 mm, included in or exerted beyond the tips of calyces 0.1-5.7 mm; wing petals 3.1-9 mm long, asymmetrically clawed, claws 1.6-5.1×≤0.1 mm, laminae elongate-elliptical, 1.3-4x0.2-1.4 mm, bases shortly auriculate, auricles rounded, petal margins entire, apices rounded; keel petals 3-7.3 mm long, asymmetrically clawed, claws 1.4- 4.8×≤0.1 mm, laminae ovate, 1-2.5x0.4-1.8 mm, base rounded, margins entire, apices rounded- short tipped; stamens diadelphous, connate filaments 2.2-6.6×≤0.5 mm; pistils sessile, ovaries 2-2.9×0.2-1.4 mm, styles 1.1-4.2×≤0.1 mm, ovules 2; legumes laterally compressed, 2.2-4×1.3-2 mm; seeds 2, globular--mitten-shaped, blackish-brown, dark-speckled or not, 1.1-1.6×1-1.2 mm. 2n=16.

Representative Specimens CANADA. British Columbia: Vancouver Island, very Common especially in wet areas on rocky- grassy bluffs above river, Sproat River Falls north of Alberni, 7 June 1961, Calder and MacKay 30139 (CAS); Victoria Three miles northeast of Duncan on road to Maple Bay, 19 June 1961, Calder and MacKay 30732 (CAS); Okanagan, Penticton, Lake Okanagan, 11 July 1889, Macoun s.n. (BM). MEXICO. Baja California: Ensenada: Sierra San Pedro Martir, meadows along road heading south of Vallecitos towards La Encantada, base of Cerro Botella Azul, 19 July 1988, Boyd et al. 2683 (MU); Laguna Hanson, Constitucion National park, Sierra de Juarez; north end of lake on sandy marsh delta and wet margins of lake, 28 May 1983, Thorne et al. 55739 (WIS). USA. Arizona: Graham County: Coronado National Forest, Holdout Creek, just above jcn with Black Rock wash along trail splitting off by corrals, 10 June 1998, Buegge et al. 338 (ASU); 4 mi. below Pine Crest, Pinaleno Mts., 19 May 1935, B. Maguire and R. Maguire 11476 (CAS). California: San Bernadino County, Vicinity of Bonanza King Mine, East Slope of Providence Mointains, Mojave Desert, 21 May 1920, Munz et al. 4020 (MU, CAS); Santa Barbara County, Santa Cruz Island, Creek below Main Ranch (toward Prisoner's Harbor), 12 October 1958, Balls 11820 (BM); Inyo County, Wild Rose Canyon, Panamont Mts., 30 June 1931, Hoffmann s.n. (CAS); Tuolumne County, Tioga Road from Harden Lake to Pareuphene Flat, Yosemite National Park, 13 August 1907, Eastwood 317 (CAS); San Diego County, Cleveland National Forest: Northeast of El Cajon Mountain, west of the San Diego River, and south of Barona Mesa, just off of El Cajon Mountain Truck Trail, 7 April 2010, Rebman et al. 18976 (SD); Kern County, Saddle Springs, north end of the crest of Piute Mountain, 22 June 1962, Twisselmann 7349 (CAS); Monterey County, "The Indians" near Jolon, 6 April 1988, Morgan 1065 (MU). Idaho: Blain County, Tikura, 22 July 1911, Nelson and Macbride 1303 (CAS); Kootenai County, June 1889,

Sandberg s.n. (MU); Owyhee County: S Quicksilver Mountain above Boone Peak, 19 July 1996, Atwood 21000 (MU, BRY); YP desert, lower Little Owyhee River, ca 41 air miles west of Riddle, 23 July 1998, Mancuso 1708 (SRP). Montana: Missoula County, Missoula, Hay Spur, Geo. F. Fox Ranch, 19 June 1924, Kirkwood 1756 (MU, ISC, CAS (2 sheets)). Nevada: Nye County, Along Eden Creek, 2 mi. east on road past Eden Ranch, about 15 mi. due south of Warm Springs, 28 August 1980, Neese and White 9744 (BRY); Elko County, Deeth, 23 July 1908, Heller 9129 (ISC); Washoe County: Above the reservoir on Heinz Ranch, northwest slope of Peavine Mountain about 10 miles north of Reno, 21 September 1974, Howell et al. 50973 (CAS); creek 0.5 mile south of Racetrack Reservoir, 5 June 1978, Rogers and Tiehm 991 (RENO, CAS); Growing on the drying mud flats at the north end of Washoe Lake, 4 August 1976, Williams et al. 76-90-14 (RENO). Oregon: Wasco County, Dalles City, June 8 1897, Suksdorf 2583 (GH); Lane County, West of Science building Lane Community College, 9 May 1992, Love 930 (MU); Clackamas County, Gladstone, May 1894, Howell 135 (WIS); Coos County, 2 mi. west of Charleston opposite Rhodo-Azalea Gds. Nursery, 29 June 1961, Kimber 122 (WIS). Utah: Box Elder County, southwest corner of the County, 16 mi. south of Lucine, east of the Pilot Range, Patterson Spring at edge of Great Salt Lake Desert in greasewood belt, 5 July 1976, Arnow 5013 (BRY, MU); Salt Lake County, Salt Lake City, July 1880, Howard s.n. (BRY). Washington: Ferry County, Open, springy, gravelly spot 1/4 mi. west of Columbia River, Kettle Falls, 17 June 1939, Weldert and Boner 151 (CAS (2 sheets)); San Juan County, Lopez Island, 17 July 1937, Evans 11524 (MU); Cowlitz County, Moist bluffs along the Columbia River near Kalama, 7 April 1934, Thompson 10129 (CAS).

Distribution and Habitat Trifolium variegatum is the most widespread of the three recognized here (Fig. 56), encompassing the ranges of the other two. It does not appear to have any unique habitat requirements other than some moisture, occurring in dry grasslands and moist vernal pools. It appears in grasslands, sandy stream beds, and other open areas, but can also be found in forests. It occurs in elevations ranging from sea level to 3,000 m. It has been collected in every month from March to October, although this varies depending on the geographic area (Table 8). Isely (1998) reports a specimen of T. variegatum from southwestern Wyoming in his Map 387. Unfortunately, no voucher specimen is cited and no specimens from Wyoming have been seen during the course of this study. It is entirely possible that T. variegatum could be found in Wyoming. There are several specimens from Salt Lake City, Utah, which is less than fifty miles from the border. One specimen, Whittier s.n. (ISC), collected in 1903 in St. Michael, Alaska, is approximately 1,700 miles from the nearest known population. No specimens in the intermediate region have yet to be found in any other herbarium. Interestingly, in 1897 St. Michael was an important port of entry for the Alaskan gold rush (State of Alaska, Department of Commerce, Community, and Economic Development, website). Many of the prospectors

64 came from California, possibly carrying Californian soil with them. Thus, this specimen is likely an introduction and the population did not likely persist long due to the colder climate.

Comments Trifolium variegatum is the correct and legitimate name for this species, despite T. melananthum, its synonym, being published in a different publication during the same year. The latter was published in December of 1838 as part 7 of the entire publication (Stafleu and Cowan, 1979), whereas the former was published as part two of its volume, in October of 1838. Trifolium variegatum var. β, so-called, is not a validly published name, since names must be constructed of Latin letters (art. 32). It is best regarded as an unpublished variety. It is mentioned in the protologue for T. variegatum (Fl. N. Amer. (Torr. & A. Gray) 1: 317. 1838), and a specimen is cited: D. Douglas s.n. (NY - electronic image!), a paratype of T. variegatum. Trifolium geminiflorum was not lectotypified by Gillett, because he thought the name was based on Lojacono’s T. pauciflorum, which is not a published name, and therefore did not think Greene could have been in possession of the type. However, Lojacono merely commented what he thought was a strange specimen of Nuttall’s T. pauciflorum and did not intend to publish a new name. It was incorrectly lectotypified by Vincent (2009) because the specimen selected (G. Hansen 1, Amador County, April 1892, cited as at NDG) does not exist at NDG. There is a Hansen specimen from 1891 (NDG67208) at NDG, and there are Hansen specimens of Hansen 1 at BM and K. Since Hansen 1 does not exist at NDG, it is unlikely that Greene saw that collection, and instead was referring to Hansen s.n. (1891, Amador County). Thus, Vincent’s incorrect lectotypification is set aside. Several interesting specimens were discovered during the course of this study. Sanders et al. 20707 (SD; California, Riverside County: San Jacinto Mountains, north side of the range, Twin Pines Creek below the confluence with Brown Creek) is a putative hybrid between T. variegatum and T. oliganthum. The inflorescence looks like normal T. variegatum, whereas the rest of the plant has leaflets similar to those of T. oliganthum. A similar situation exists with Smith & Wheeler 6352 (CAS; California, Mendocino County: large flood plain on Rancheria Creek, 7.3 km from junction with Rte. 128 at Yorkville, first bridge, on big curve of creek), which has T. oliganthum-like stipules in addition to leaflets. A specimen from Table Mountain, Butte County, CA, (Vincent et al. 8110 (MU)) appears to be a hybrid between T. variegatum and T. jokerstii, having the former’s flower shape and light purplish color with the latter’s rounded stipules and bowl-shaped involucre. A number of specimens collected by Randall Morgan (Morgan 1622 (MU), Morgan 1597 (MU), and Morgan 2033 (MU)) in Santa Cruz County, CA, bear the name “T. pseudovariegatum” on their labels. These specimens have inflorescences that seem more racemose than umbellate, long stipules, long petioles, and a higher number of distal secondary stems. Other specimens from Idaho (Atwood 20319 (BRY), and Ertter 4101 (CAS, ISC) and from San Mateo County, CA (Abrams 5077 (MU)) also share these features. Due to this strange distribution and the lack of other distinguishing characters, this group has been included in T. variegatum.

Fig. 55. Specimen of T. variegatum (Gillett and Crompton 12951 (MU)).

Fig. 56. Distribution map of T. variegatum specimens.

Excluded Taxon Trifolium variegatum var. parunuweapensis S.L. Welsh, Utah Flora (ed. 3) 425. 2003. TYPE: USA, Utah: Kane County, ca. 1.5 mi S of The Barracks, subsequent drainage of Parunuweap Canyon, 1450m, 16 July 1992, S.L. Welsh & K.H. Thorne 25120 (holotype: BRY!). After comparing the type specimen of T. variegatum var. parunuweapensis to the type specimen of T. mucronatum subsp. lacerum (Greene) Gillett (holotype: US - electronic image!, isotype: NY - electronic image!), it is evident that the former is a synonym of the latter. This is because T. variegatum var. parunuweapensis, like T. mucronatum subsp. lacerum, produces rhizomes (visible on paratypes, difficult to distinguish on holotype), has stipules up to 30 mm long, and has 3 or 4 ovules. None of these features are ever found on any of the species recognized as a part of the Trifolium variegatum species complex.

References Axelrod, D.I. 1973. History of the Mediterranean ecosystem in California. Mediterranean Type Ecosystems. (ed. by F. diCastri and H.A. Mooney). Springer, Berlin, Germany. pp. 225–305.

Ayensu, E.S. 1967. Aerosol OT solution—an effective softener of herbarium specimens for anatomical study. Stain Technology 42: 155–156.

Baldwin, B. G., D. H. Goldman, D. J. Keil, R. Patterson, T. J. Rosatti, and D. H. Wilken, editors. 2012. The Jepson manual: vascular plants of California, second edition. University of California Press, Berkeley, California.

Becker, R.A., A.R. Wilks, R. Brownrigg, and T.P. Minka. 2014a. maps: Draw Geographical Maps. R package version 2.3-9. http://CRAN.R-project.org/package=maps.

Becker, R.A., A.R. Wilks, and R. Brownrigg. 2014b. mapdata: Extra Map Databases. R package version 2.2-3. http://CRAN.R-project.org/package=mapdata.

Biau, D.J., B.M. Jolles, and R. Porcher. 2010. P Value and the Theory of Hypothesis Testing: An Explanation for New Researchers. Clinical Orthopaedics and Related Research 468: 885-892.

Cronquist, A. 1988. The Evolution and Classification of Flowering Plants. Second Edition. The New York Botanical Garden, Bronx, New York, USA.

Dunn, G., and B.S. Everitt. 1982. An introduction to mathematical taxonomy. Cambridge University Press, Cambridge, United Kingdom.

Ellison, N.W., A. Liston, J.J. Steiner, W.M. Williams, and N.L. Taylor. 2006. Molecular phylogenetics of the clover genus (Trifolium—Leguminosae). Molecular Phylogenetics and Evolution 39: 688-705.

Fisher, R.A. 1936. The use of multiple measurements in taxonomic problems. Annals of Eugenics 7: 179-188.

Gillett, J.M. 1966. Type Collections of Trifolium in the Greene Collection at Notre Dame. The American Midland Naturalist 76: 468-474.

Giraudoux, P. 2014. Pgirmess: Data analysis in ecology. R package version 1.5-9. http://perso.orange.fr/giraudoux.

Hickman, J.C. 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley, California, USA.

Hitchcock, C. L., A. Cronquist, M. Owenby, and J. W. Thompson. 1955–1964. Vascular plants of the Pacific Northwest. University of Washington Press, Seattle, Washington, USA.

Hotelling, H. 1933. Analysis of a complex of statistical variables into principal components. The Journal of Educational Psychology 24: 417-441, 498-520.

Isely, D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States (exclusive of Alaska and Hawaii). Monte L. Bean Life Science Museum Press, Provo, Utah, USA.

Jepson, W.L. 1925. Manual of the Flowering Plants of California. University of California Press, Berkeley and Los Angeles, California, USA.

Jepson, W.L. 1936. A Flora of California. Vol. 2: Capparidaceae to Cornaceae. Associated Students Store, Berkeley, California, USA.

Kampstra, P. 2008. Beanplot: A Boxplot Alternative for Visual Comparison of Distributions. R package version 1.2. Journal of Statistical Software, Code Snippets 28: 1-9. http://www.jstatsoft.org/v28/c01/.

Kruskal, W.H., and W.A. Wallis. 1952. Use of Ranks in One-Criterion Variance Analysis. Journal of the American Statistical Association 47: 583-621.

Ligges, U. and M. Machler. 2011. scatterplot3d: 3D Scatter Plot. R package version 0.3-35. http://cran.r-project.org/web/packages/scatterplot3d.

Marhold, K. 2011. Multivariate morphometrics and its application to monography at infraspecific levels. In T.F. Stuessy and H.W. Lack [eds.], Monographic Plant Systematics: Fundamental Assessment of Biodiversity. Regnum Vegetabile 153: 73-99.

McDermott, L.F. 1910. An illustrated key to the North American species of Trifolium. Cunningham, Curtis, and Welch, San Francisco, California, USA.

Munz, P.A., and D.D. Keck. 1959. A California Flora. University of California Press, Berkeley and Los Angeles, California, USA.

Pearson, K. 1901. On lines and planes of closest fit to systems of points in space. Philosophical Magazine 2: 559-572.

R Core Team. 2013. R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna, Austria. http://www.R-project.org/.

Raven, P.H. 1963. Amphitropical Relationships in the Floras of North and South America. The Quarterly Review of Biology 38: 151-177.

Raven, P.H., and D.I. Axelrod. 1978. Origin and relationships of the California flora. University of California Publications in Botany, vol. 72. University of California Press, Berkeley and Los Angeles, California, USA.

Rohlf, F. J. 2012. NTSYSpc: numerical taxonomy system. ver. 2.21q. Exeter Software. Setauket, New York, USA.

SAS Institute. 2013. JMP Pro 11.2.0. SAS Institute Inc. Cary, North Carolina, USA.

Shapiro, S.S., and M.B. Wilk. 1965. An analysis of variance test for normality (complete samples). Biometrika 52: 591-611.

Sneath, P.H.A., and R.R. Sokal. 1973. Numerical Taxonomy. W.H. Freeman, San Francisco, California, USA.

Sokal, R.R., and C.D. Michener. 1958. A Statistical Method for Evaluating Systematic Relationships. The University of Kansas Scientific Bulletin 38: 1409-1438.

Stafleu, F.A., and R.S. Cowan. 1979. Taxonomic literature 2. Ed. 2. Bohn, Scheltema & Holkema, Utrecht, Netherlands.

State of Alaska, Department of Commerce, Community, and Economic Development, Community and Regional Affairs. Community: St. Michael [online]. Website http://commerce.state.ak.us/cra/DCRAExternal/community/Details/e565d757-3b9d-4dbf- 90b4-fcdd2b696ab8 [accessed 23 April 2015].

Takhtajan, A.L. 1986. Floristic Regions of the World. University of California Press, Berkeley and Los Angeles, California, USA.

Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden's Virtual Herbarium. Website http://sweetgum.nybg.org/ih/ [accessed 8 May 2015].

Tidestrom, I. 1925. Flora of Utah and Nevada. Government Printing Office, Washington D.C., USA.

Vincent, M.A. 2009. A New Combination in Trifolium variegatum (Fabaceae). Madroño 56: 208.

Zohary, M., and D. Heller. 1984. The genus Trifolium. Israel Academy of Sciences and Humanities, Jerusalem, Israel.

Appendices Appendix 1. Kruskal-Wallace tests and multiple comparisons for accepted taxa and potential taxa. False differences indicate that the comparison is not considered significant, true differences are those comparisons considered significant.

Character Accepted T. appendiculatum vs. T. appendiculatum vs. T. variegatum vs Taxa: KW p T. polyodon T. variegatum T. polyodon

Stem Diameter (cm) 1.51E-07 FALSE TRUE FALSE Internode Length (cm) 0.3362 N/A N/A N/A Stipule Length (cm) 3.29E-08 FALSE TRUE TRUE Stipule Lobe Length (cm) 0.0006716 FALSE TRUE TRUE Petiole Length (cm) 8.11E-05 FALSE TRUE FALSE Petiole Diameter (cm) 6.27E-10 FALSE TRUE TRUE Petiolule Length (cm) 5.82E-07 FALSE TRUE FALSE Petiolule Diameter (cm) 5.28E-11 FALSE TRUE TRUE Leaflet Base Angle (degrees) 4.66E-07 FALSE TRUE FALSE Leaflet Length (cm) 2.08E-09 FALSE TRUE TRUE Leaflet Width (cm) 2.95E-13 FALSE TRUE TRUE Head Length (cm) < 2.2e-16 FALSE TRUE FALSE Flower Number 2.16E-13 FALSE TRUE TRUE Peduncle Length (cm) 2.20E-16 TRUE TRUE FALSE Peduncle Diameter (cm) 2.20E-16 FALSE TRUE FALSE Involucre Radius (cm) 2.20E-16 FALSE TRUE TRUE Involucral Lobe Length (cm) 1.83E-08 FALSE TRUE FALSE Calyx Length (cm) < 2.2e-16 FALSE TRUE TRUE Calyx Circumference (cm) 2.32E-14 FALSE TRUE FALSE Calyx Lobe Length (cm) 1.72E-15 FALSE TRUE TRUE Calyx Lobe Width (cm) 3.04E-05 FALSE TRUE TRUE Calyx Tube Length (cm) 2.93E-11 FALSE TRUE FALSE Calyx Lobe Tooth Number 4.34E-06 TRUE FALSE TRUE Banner Petal Length (cm) < 2.2e-16 FALSE TRUE FALSE Banner Petal Width (cm) < 2.2e-16 FALSE TRUE TRUE Wing Petal Length (cm) < 2.2e-16 FALSE TRUE TRUE Wing Petal Lamina Length (cm) < 2.2e-16 FALSE TRUE FALSE Wing Petal Lamina Width (cm) < 2.2e-16 FALSE TRUE TRUE Keel Petal Length (cm) < 2.2e-16 FALSE TRUE TRUE Keel Petal Lamina Length (cm) < 2.2e-16 FALSE TRUE FALSE Keel Petal Lamina Width (cm) < 2.2e-16 FALSE TRUE FALSE Connate Filaments Length (cm) < 2.2e-16 FALSE TRUE TRUE Style Length (cm) < 2.2e-16 FALSE TRUE TRUE Style Diameter (cm) 3.11E-11 FALSE TRUE FALSE

Appendix 1. (continued)

Character Potential "T. "T. "T. "T. "T. Taxa KW p geminiflorum geminiflorum" geminifloru geminifloru subsalinum" " vs. "T. vs. T. m" vs. T. m" vs. T. vs. T. subsalinum" appendiculatum polyodon variegatum appendiculat um Stem Diameter (cm) 8.22E-14 TRUE TRUE TRUE TRUE FALSE Internode Length (cm) 0.00414 FALSE TRUE FALSE TRUE FALSE Stipule Length (cm) 9.48E-15 TRUE TRUE TRUE TRUE FALSE Stipule Lobe Length 5.61E-08 FALSE TRUE TRUE TRUE FALSE (cm) Petiole Length (cm) 4.79E-05 FALSE TRUE FALSE FALSE TRUE Petiole Diameter (cm) 1.91E-14 TRUE TRUE TRUE TRUE FALSE Petiolule Length (cm) 2.92E-10 FALSE TRUE TRUE TRUE FALSE Petiolule Diameter 3.41E-15 TRUE TRUE TRUE TRUE FALSE (cm) Leaflet Base Angle 9.62E-07 FALSE TRUE FALSE FALSE TRUE (degrees) Leaflet Length (cm) 1.54E-15 TRUE TRUE TRUE TRUE TRUE Leaflet Width (cm) < 2.2e-16 FALSE TRUE TRUE TRUE TRUE Head Length (cm) < 2.2e-16 TRUE TRUE FALSE FALSE TRUE Flower Number < 2.2e-16 TRUE TRUE TRUE TRUE FALSE Peduncle Length (cm) < 2.2e-16 FALSE TRUE FALSE TRUE TRUE Peduncle Diameter < 2.2e-16 TRUE TRUE TRUE TRUE FALSE (cm) Involucre Radius (cm) < 2.2e-16 TRUE TRUE TRUE TRUE FALSE Involucral Lobe Length 1.41E-10 TRUE TRUE TRUE TRUE FALSE (cm) Calyx Length (cm) < 2.2e-16 TRUE TRUE TRUE TRUE FALSE Calyx Circumference 1.71E-15 TRUE TRUE FALSE TRUE FALSE (cm) Calyx Lobe Length (cm) < 2.2e-16 FALSE TRUE TRUE TRUE TRUE Calyx Lobe Width (cm) 2.68E-06 TRUE TRUE TRUE FALSE FALSE Calyx Tube Length (cm) 9.05E-15 TRUE TRUE FALSE TRUE FALSE Calyx Lobe Tooth 2.50E-05 FALSE FALSE TRUE FALSE FALSE Number Banner Petal Length < 2.2e-16 FALSE TRUE TRUE FALSE TRUE (cm) Banner Petal Width < 2.2e-16 FALSE TRUE TRUE TRUE TRUE (cm) Wing Petal Length (cm) < 2.2e-16 FALSE TRUE TRUE FALSE TRUE Wing Petal Lamina < 2.2e-16 FALSE TRUE TRUE TRUE TRUE Length (cm) Wing Petal Lamina < 2.2e-16 FALSE TRUE TRUE FALSE TRUE Width (cm) Keel Petal Length (cm) < 2.2e-16 TRUE TRUE TRUE FALSE TRUE Keel Petal Lamina < 2.2e-16 FALSE TRUE FALSE FALSE TRUE Length (cm) Keel Petal Lamina < 2.2e-16 FALSE TRUE TRUE FALSE TRUE Width (cm) Connate Filaments < 2.2e-16 TRUE TRUE TRUE FALSE TRUE Length (cm)

Appendix 1. (continued)

Style Length (cm) < 2.2e-16 TRUE TRUE TRUE FALSE TRUE Style Diameter (cm) 5.33E-10 FALSE TRUE FALSE FALSE TRUE

Appendix 1. (continued)

Character "T. "T. T. T. T. polyodon vs. subsalinum" subsalinum" vs. appendiculatum appendiculatum T. variegatum vs. T. polyodon T. variegatum vs. T. polyodon vs. T. variegatum Stem Diameter (cm) FALSE FALSE FALSE TRUE FALSE Internode Length (cm) FALSE FALSE FALSE FALSE FALSE Stipule Length (cm) TRUE FALSE FALSE TRUE TRUE Stipule Lobe Length (cm) FALSE FALSE FALSE FALSE FALSE Petiole Length (cm) TRUE FALSE FALSE TRUE FALSE Petiole Diameter (cm) FALSE FALSE FALSE TRUE FALSE Petiolule Length (cm) FALSE FALSE FALSE TRUE FALSE Petiolule Diameter (cm) FALSE FALSE FALSE TRUE FALSE Leaflet Base Angle TRUE FALSE FALSE TRUE FALSE (degrees) Leaflet Length (cm) FALSE FALSE FALSE TRUE FALSE Leaflet Width (cm) TRUE FALSE FALSE TRUE FALSE Head Length (cm) FALSE FALSE FALSE TRUE FALSE Flower Number FALSE FALSE FALSE TRUE FALSE Peduncle Length (cm) FALSE FALSE FALSE TRUE FALSE Peduncle Diameter (cm) FALSE FALSE FALSE TRUE FALSE Involucre Radius (cm) FALSE FALSE FALSE TRUE TRUE Involucral Lobe Length FALSE FALSE FALSE TRUE FALSE (cm) Calyx Length (cm) FALSE FALSE FALSE TRUE FALSE Calyx Circumference (cm) FALSE FALSE FALSE TRUE FALSE Calyx Lobe Length (cm) FALSE FALSE FALSE TRUE FALSE Calyx Lobe Width (cm) FALSE FALSE FALSE TRUE FALSE Calyx Tube Length (cm) FALSE TRUE FALSE TRUE FALSE Calyx Lobe Tooth Number FALSE FALSE TRUE FALSE TRUE Banner Petal Length (cm) FALSE FALSE FALSE TRUE FALSE Banner Petal Width (cm) FALSE FALSE FALSE TRUE FALSE Wing Petal Length (cm) FALSE FALSE FALSE TRUE FALSE Wing Petal Lamina Length FALSE FALSE FALSE TRUE FALSE (cm) Wing Petal Lamina Width FALSE FALSE FALSE TRUE FALSE (cm) Keel Petal Length (cm) FALSE FALSE FALSE TRUE FALSE Keel Petal Lamina Length FALSE FALSE FALSE TRUE FALSE (cm) Keel Petal Lamina Width FALSE FALSE FALSE TRUE FALSE (cm) Connate Filaments Length FALSE FALSE FALSE TRUE FALSE (cm) Style Length (cm) FALSE FALSE FALSE TRUE TRUE Style Diameter (cm) FALSE FALSE FALSE TRUE FALSE

Appendix 2. Label and measurement data for specimens used in multivariate analyses.

Numb Her Collector Name Collector Month State/Region County Species er bari Number Collected um 167 ISC Parry s.n. May NA San Luis Obispo T. variegatum 70 ISC Whittier s.n. NA Alaska Nome T. variegatum 306 CAS B. Maguire and R. 11476 May Arizona Graham T. variegatum Maguire 424 ASU Buegge et al. 338 June Arizona Graham T. variegatum 425 ASU Buegge 873 May Arizona Graham T. variegatum 319 CAS Pringle s.n. April Arizona NA T. variegatum 69 MU Boyd et al. 2683 July Baja Ensenada T. variegatum California 229 WIS Thorne et al. 55739 May Baja Ensenada T. variegatum California 26 MU Macoun 78941 June British Vancouver Island T. variegatum Columbia 41 MU Macoun 78942 July British Vancouver Island T. variegatum Columbia 292 CAS Calder and 30139 June British Vancouver Island T. variegatum MacKay Columbia 322 CAS Calder and 29518 May British Vancouver Island T. variegatum MacKay Columbia 303 CAS Calder and 30732 June British Victoria T. variegatum MacKay Columbia 304 CAS Eastwood 9821 June British Victoria T. variegatum Columbia 305 CAS Eastwood 9707 June British Victoria T. variegatum Columbia 380 CAS S. Parish and W. 749a May California NA T. variegatum Parish 225 WIS Walker 550 April California Alameda T. variegatum 359 CAS Beet s.n. June California Alpine T. variegatum 226 WIS Heller 11818 April California Butte T. variegatum 327 CAS Ahart 6212 May California Butte T. appendiculatum 328 CAS Heller 10710 April California Butte T. appendiculatum 369 CAS Eastwood 11587 April California Calaveras T. appendiculatum 220 WIS Howell 11227 May California Contra Costa T. appendiculatum 352 CAS Mexia 2379 April California Contra Costa T. appendiculatum 353 CAS Howell 11227 May California Contra Costa T. appendiculatum 368 CAS Robbins 1856 April California El Dorado T. appendiculatum 382 CAS Raven and 9094 April California El Dorado T. appendiculatum Robbins 325 CAS Shannon s.n. April California Fresno T. appendiculatum 337 CAS Beane 36 July California Fresno T. variegatum 356 CAS Heller 11347 May California Glenn T. appendiculatum 75 ISC Tracy 16238 April California Humboldt T. appendiculatum 182 ISC Gunn 2701 June California Humboldt T. variegatum 189 ISC Gunn 2701 June California Humboldt T. variegatum

Appendix 2. (continued)

12 MU DeDecker 6101 May California Inyo T. variegatum 311 CAS Hoffmann s.n. June California Inyo T. variegatum 171 ISC Heller 7821 May California Kern T. appendiculatum 339 CAS Twisselmann 7349 June California Kern T. variegatum 370 CAS Twisselmann 9465 May California Kern T. appendiculatum 371 CAS Twisselmann 8251 May California Kern T. appendiculatum 372 CAS Twisselmann 12209 May California Kern T. appendiculatum 373 CAS Twisselmann 5146 May California Kern T. variegatum 374 CAS Evermann s.n. April California Kern T. appendiculatum 375 CAS Twisselmann 7505 July California Kern T. variegatum 376 CAS Heller 7821 May California Kern T. appendiculatum 377 CAS Breedlove 3954 July California Kern T. variegatum 87 MU Crampton 2767 May California Lake T. appendiculatum 38 MU Wheeler s.n. September California Los Angeles T. variegatum 180 ISC Thorne 38233 July California Los Angeles T. variegatum 361 CAS Harwood 3297 March California Los Angeles T. appendiculatum 362 CAS Sheet and 2496 April California Los Angeles T. appendiculatum Williams 384 CAS Griesel s.n. June California Los Angeles T. variegatum 21 MU Evans 11433 July California Madera T. variegatum 22 MU Evans 11432 July California Madera T. variegatum 366 CAS Kawhara 675 April California Madera T. appendiculatum 367 CAS Howell 41087 April California Madera T. variegatum 386 CAS Raven 3377 July California Madera T. variegatum 30 MU Vincent 8600 June California Marin T. appendiculatum 170 ISC Rose 46037 April California Marin T. appendiculatum 336 CAS Howell 20825 May California Marin T. appendiculatum 378 CAS Howell 23064 April California Marin T. appendiculatum 329 CAS Breedlove 17375 April California Mariposa T. appendiculatum 400 CAS Kennedy 3030 July California Mariposa T. variegatum 46 MU Gillett and 12951 June California Mendocino T. variegatum Crompton 74 ISC Tracy 15752 June California Mendocino T. appendiculatum 176 ISC Kennedy 10309 May California Mendocino T. appendiculatum 185 ISC Gunn 2666 June California Mendocino T. variegatum 187 ISC Gunn 2666 June California Mendocino T. variegatum 350 CAS Smith and 6340 May California Mendocino T. variegatum Wheeler 50 MU Morgan 1065 April California Monterey T. variegatum 76 ISC Heller 6722 May California Monterey T. variegatum 78 ISC Heller 8504 May California Monterey T. variegatum 143 MU Vincent et al. 8195 April California Monterey T. polyodon 181 ISC Heller 6727 May California Monterey T. variegatum 186 ISC Andrews 344 California Monterey T. variegatum

Appendix 2. (continued)

222 WIS Heller 28022 May California Monterey T. polyodon 223 WIS Heller 6707 May California Monterey T. polyodon 332 CAS Kennedy s.n. May California Monterey T. variegatum 175 ISC Jepson 9177 May California Napa T. variegatum 179 ISC Ottley 1326 May California Napa T. variegatum 166 ISC Isely 9624 May California Nevada T. variegatum 326 CAS Heller 9830 July California Nevada T. variegatum 55 MU Crampton 7647 April California Placer T. appendiculatum 155 ISC Heller and 8873 July California Plumas T. variegatum Kennedy 351 CAS Janeway 4135 August California Plumas T. variegatum 18 MU Boyd et al. 3636 June California Riverside T. variegatum 257 SD Boyd et al. 7332 May California Riverside T. variegatum 258 SD White and Carey 10331 May California Riverside T. variegatum 333 CAS Lathrop 4674 March California Riverside T. variegatum 47 MU Crampton 1078 April California Sacramento T. appendiculatum 354 CAS Ramaley 11078 April California Sacramento T. appendiculatum 355 CAS Copeland 1601 April California Sacramento T. variegatum 15 MU Vincent et al. 8191 April California San Benito T. variegatum 65 MU Munz et al. 4020 May California San Bernadino T. variegatum 309 CAS Munz et al. 4020 May California San Bernadino T. variegatum 62 MU Spencer s.n. June California San Diego T. variegatum 67 MU Banks and 1922 April California San Diego T. variegatum Steinmann 218 SD Gander s.n. May California San Diego T. variegatum 219 SD Hendrickson 1084 June California San Diego T. appendiculatum 259 SD Boyd and Ross 7597 June California San Diego T. variegatum 260 SD Banks and Boyd 587 June California San Diego T. variegatum 261 SD Rebman et al. 9320 June California San Diego T. variegatum 262 SD Lightner 246 June California San Diego T. variegatum 263 SD Gregory et al. 3236 June California San Diego T. variegatum 264 SD Rebman et al. 23601 May California San Diego T. variegatum 265 SD Cain et al. 753 April California San Diego T. appendiculatum 267 SD Rebman et al. 18976 April California San Diego T. variegatum 268 SD Rebman et al. 19657 June California San Diego T. variegatum 269 SD Calcarone 169 March California San Diego T. appendiculatum 270 SD Rebman et al. 21862 May California San Diego T. variegatum 271 SD Gregory and 3253 June California San Diego T. variegatum Marshall 272 SD Rebman et al. 21966 May California San Diego T. variegatum 273 SD Hendrickson 43b May California San Diego T. variegatum 274 SD Rebman and 7300 April California San Diego T. variegatum Hollingsworth 275 SD Pigniolo 1348 June California San Diego T. variegatum 276 SD Rebman et al. 9327 June California San Diego T. variegatum

Appendix 2. (continued) 277 SD Moran s.n. May California San Diego T. variegatum 278 SD Lauri and 193 May California San Diego T. variegatum Rebman 279 SD Rebman et al. 10478 June California San Diego T. variegatum 280 SD J. Gregory and J. 1807 July California San Diego T. variegatum Gregory 281 SD Hirshberg 1001 April California San Diego T. variegatum 282 SD Hendrickson and 707 May California San Diego T. appendiculatum Jee 283 SD Hendrickson and 666 May California San Diego T. appendiculatum Jee 284 SD Rebman et al. 12071 June California San Diego T. variegatum 286 SD Rebman 15030 May California San Diego T. variegatum 288 SD Rebman et al. 17906 May California San Diego T. variegatum 290 SD Beauchamp and 2783 June California San Diego T. variegatum Howe 291 SD Nenow and Glacy 1206 May California San Diego T. variegatum 310 CAS Wiggins 2734 September California San Diego T. variegatum 360 CAS Abrams 3844 June California San Diego T. appendiculatum 358 CAS Smith 1063 May California San Joaquin T. appendiculatum 49 MU Keck 1434 April California San Mateo T. variegatum 61 MU Abrams 5077 April California San Mateo T. variegatum 64 MU Rose 37251 May California San Mateo T. variegatum 48 MU Dudley s.n. April California Santa Clara T. variegatum 52 MU Dudley s.n. May California Santa Clara T. variegatum 178 ISC Heller 8600 May California Santa Clara T. variegatum 1 MU Morgan 2033 May California Santa Cruz T. variegatum 4 MU Morgan 1597 May California Santa Cruz T. variegatum 6 MU Morgan 1999 March California Santa Cruz T. appendiculatum 7 MU Morgan 1433 April California Santa Cruz T. appendiculatum 8 MU Morgan 1387 March California Santa Cruz T. appendiculatum 36 MU Morgan 1622 May California Santa Cruz T. variegatum 42 MU Morgan 2031 April California Santa Cruz T. variegatum 51 MU Morgan 1458 April California Santa Cruz T. variegatum 330 CAS Thomas 3149 May California Santa Cruz T. variegatum 13 MU Ahart and Dittes 10155 May California Shasta T. appendiculatum 165 ISC Heller 7892 May California Shasta T. variegatum 80 ISC Heller 7964 June California Siskiyou T. variegatum 56 MU Crampton 7671 April California Solano T. variegatum 357 CAS Kennedy s.n. May California Solano T. variegatum 73 ISC Rubtzoff 6023 May California Sonoma T. appendiculatum 334 CAS de Nevers 8173 May California Sonoma T. variegatum 389 CAS Rubtzoff 4990 July California Sonoma T. variegatum 390 CAS Rubtzoff 5750 June California Sonoma T. variegatum 391 CAS Rubtzoff 6022 May California Sonoma T. variegatum

Appendix 2. (continued)

335 CAS Ahart 5310 May California Sutter T. appendiculatum 379 CAS Ahart 4566 April California Sutter T. appendiculatum 392 CAS Ahart 3305 April California Sutter T. appendiculatum 29 MU Straw et al. 2620 May California Tulare T. appendiculatum 153 ISC Thorne and 36989 June California Tulare T. appendiculatum Everett 154 ISC DeBuhr et al. 969 April California Tulare T. variegatum 341 CAS Howell and True 48568 July California Tulare T. variegatum 342 CAS Twisselmann 5538 July California Tulare T. variegatum 343 CAS Twisselmann 16180 October California Tulare T. variegatum 344 CAS Twisselmann 16151 September California Tulare T. variegatum 345 CAS Bacigalupi et al. 2587 July California Tulare T. variegatum 346 CAS Dudley 1769 July California Tulare T. variegatum 347 CAS Howell 27046 July California Tulare T. variegatum 348 CAS Howell 44429 May California Tulare T. variegatum 349 CAS Howell and True 47122 July California Tulare T. variegatum 393 CAS Twisselmann 15246 May California Tulare T. appendiculatum 394 CAS Twisselmann 5205 May California Tulare T. appendiculatum 395 CAS Norris 424 May California Tulare T. appendiculatum 53 MU Wiggins 20746 June California Tuolumne T. variegatum 338 CAS Penalosa 2619 June California Tuolumne T. variegatum 381 CAS Keck 1237 June California Tuolumne T. variegatum 396 CAS Thomas 8744 May California Tuolumne T. variegatum 397 CAS Keck 5598 May California Tuolumne T. variegatum 398 CAS C. Quick and A. 66-12 May California Tuolumne T. variegatum Quick 399 CAS Eastwood 317 August California Tuolumne T. variegatum 401 CAS Botti 154 June California Tuolumne T. variegatum 363 CAS Pollard s.n. March California Ventura T. appendiculatum 364 CAS Pollard s.n. May California Ventura T. appendiculatum 365 CAS Pollard s.n. April California Ventura T. appendiculatum 101 SRP Ertter 21078 June Idaho Ada T. variegatum 383 CAS Clark 255 August Idaho Ada T. variegatum 318 CAS Nelson and 1303 July Idaho Blain T. variegatum Macbride 151 SRP DeBolt 1252 July Idaho Camas T. variegatum 33 MU Smith 8089 June Idaho Elmore T. variegatum 77 ISC Ertter 4101 June Idaho Elmore T. variegatum 88 BRY Smith 8089 June Idaho Elmore T. variegatum 146 SRP Smith 8089 June Idaho Elmore T. variegatum 164 ISC Hitchcock and 10194 July Idaho Elmore T. variegatum Muhlick 388 CAS Ertter 4101 June Idaho Elmore T. variegatum 149 SRP Brown CGB 75-100 June Idaho Gooding T. variegatum 3 MU Sandberg s.n. June Idaho Kootenai T. variegatum

Appendix 2. (continued)

84 MU Atwood 21000 July Idaho Owyhee T. variegatum 91 BRY Atwood 20319 July Idaho Owyhee T. variegatum 92 BRY Atwood 21000 July Idaho Owyhee T. variegatum 152 SRP Mancuso 1708 July Idaho Owyhee T. variegatum 232 WIS Woods and 2599 July Idaho NA T. variegatum Tidestrom 320 CAS Kirkwood 1756 June Montana Missoula T. variegatum 321 CAS Kirkwood 1756 June Montana Missoula T. variegatum 200 REN Tiehm and Lott 3680A July Nevada Churchill T. variegatum O 195 REN Bair and Tiehm 218 June Nevada Douglas T. variegatum O 201 REN Tiehm 3660 July Nevada Douglas T. variegatum O 324 CAS Rose s.n. July Nevada Douglas T. variegatum 32 MU Naczi and Thieret 3399 August Nevada Elko T. variegatum 94 BRY Tiehm and 12871 June Nevada Elko T. variegatum Nachlinger 157 ISC Heller 9129 July Nevada Elko T. variegatum 193 REN Tiehm and 12871 June Nevada Elko T. variegatum O Nachlinger 199 REN Tiehm and 7153 June Nevada Elko T. variegatum O Williams 387 CAS Tiehm and 7153 June Nevada Elko T. variegatum Williams 95 BRY Morefield 4855 August Nevada Esmeralda T. variegatum 215 REN Archer 7229 September Nevada Esmeralda T. variegatum O 97 BRY Gentry and 1620 June Nevada Humboldt T. variegatum Davidse 190 REN Tiehm and Rogers 4461 June Nevada Humboldt T. variegatum O 203 REN Williams and 83-97-1 July Nevada Humboldt T. variegatum O Tiehm 205 REN Tiehm and 1109 June Nevada Humboldt T. variegatum O Williams 207 REN Lewis 3582 July Nevada Humboldt T. variegatum O 323 CAS Gentry and 1620 June Nevada Humboldt T. variegatum Davidse 217 REN Tiehm and Tucker 6725 July Nevada Lyon T. variegatum O 98 BRY Neese and White 9744 August Nevada Nye T. variegatum 99 BRY Neese and Welsh 9097 July Nevada Nye T. variegatum 191 REN Niles et al. 6278 June Nevada Nye T. variegatum O 54 MU Tiehm 14991 June Nevada Washoe T. variegatum 93 BRY Tiehm 14991 June Nevada Washoe T. variegatum 96 BRY Williams et al. 83-84-6 July Nevada Washoe T. variegatum 100 SRP Tiehm 14991 June Nevada Washoe T. variegatum 103 BM Jones s.n. June Nevada Washoe T. variegatum

Appendix 2. (continued) 194 REN Williams and 75-113-3 August Nevada Washoe T. variegatum O McPherson 196 REN Tiehm 14991 June Nevada Washoe T. variegatum O 197 REN Williams 73-20-16 July Nevada Washoe T. variegatum O 202 REN Williams et al. 83-84-6 July Nevada Washoe T. variegatum O 204 REN Howell and True 48997 July Nevada Washoe T. variegatum O 208 REN Williams et al. 76-90-14 August Nevada Washoe T. variegatum O 210 REN Breene 544 July Nevada Washoe T. variegatum O 212 REN Archer 6526 August Nevada Washoe T. variegatum O 213 REN Archer 6527 August Nevada Washoe T. variegatum O 216 REN Archer 6309 July Nevada Washoe T. variegatum O 307 CAS Howell et al. 50973 September Nevada Washoe T. variegatum 308 CAS Kennedy 3056 July Nevada Washoe T. variegatum 312 CAS Williams and 74-57-3 August Nevada Washoe T. variegatum Tiehm 313 CAS Williams and 75-82-5 July Nevada Washoe T. variegatum Tiehm 314 CAS Kennedy 1290 August Nevada Washoe T. variegatum 315 CAS Rogers and Tiehm 991 June Nevada Washoe T. variegatum 316 CAS Kennedy 1298 May Nevada Washoe T. variegatum 317 CAS Kennedy 1290 August Nevada Washoe T. variegatum 385 CAS Howell et al. 49597 June Nevada Washoe T. variegatum 228 WIS Wosley s.n. August Oregon Antimony Springs T. variegatum 224 WIS Howell 135 May Oregon Clackamas T. variegatum 231 WIS Kimber 122 June Oregon Coos T. variegatum 188 ISC Heller 10031 May Oregon Josephine T. appendiculatum 293 CAS Heller 10030 May Oregon Josephine T. appendiculatum 294 CAS Heller 10031 May Oregon Josephine T. appendiculatum 295 CAS Heller 10031 May Oregon Josephine T. appendiculatum 5 MU Love 930 May Oregon Lane T. variegatum 233 WIS Kimber 68 June Oregon Lane T. variegatum 85 MU Arnow 5013 July Utah Box Elder T. variegatum 89 BRY Arnow 5013 July Utah Box Elder T. variegatum 31 MU Leonard s.n. July Utah Salt Lake T. variegatum 331 CAS Thompson 10129 April Washington Cowlitz T. variegatum 296 CAS Weldert and 151 June Washington Ferry T. variegatum Boner 298 CAS Weldert and 151 June Washington Ferry T. variegatum Boner 300 CAS Thompson 10 June Washington Grant T. variegatum 9 MU Anonymous s.n. May Washington King T. variegatum

Appendix 2. (continued)

230 WIS Shumway 801 NA Washington King T. variegatum 299 CAS Heller s.n. May Washington Klickitat T. variegatum 302 CAS Thompson 11562 May Washington Klickitat T. variegatum 297 CAS Fiker 855 June Washington Okanogan T. variegatum 19 MU Evans 11526 July Washington San Juan T. variegatum 20 MU Evans 11527 July Washington San Juan T. variegatum 23 MU Evans 11525 July Washington San Juan T. variegatum 24 MU Evans 11524 July Washington San Juan T. variegatum 301 CAS Hitchcock and 21590 May Washington Yakima T. variegatum Muhlick

Appendix 2. (continued)

Number Stem Internode Stipule Stipule Petiole Petiole Petiolule Petiolule Diameter Length Length Lobe Length Diameter Length Diameter (cm) (cm) (cm) Length (cm) (cm) (cm) (cm) (cm) 167 0.04 0 0.424 0.156 0.9 0.032 0.03 0.016 70 0.048 0.37 0.336 0.144 0.6 0.036 0.028 0.012 306 0.024 0.27 0.328 0.144 0.3 0.016 0.03 0.01 424 0.02 0.39 0.336 0.096 0.2 0.016 0.018 0.012 425 0.028 0.63 0.216 0.084 0.1 0.02 0.016 0.01 319 0.072 3.62 0.624 0.236 1.5 0.044 0.046 0.02 69 0.072 3.08 0.392 0.18 0.4 0.024 0.026 0.016 229 0.052 1.51 0.512 0.148 0.2 0.032 0.04 0.022 26 0.056 4.15 0.608 0.228 1.3 0.028 0.032 0.018 41 0.076 2.65 0.408 0.204 1.7 0.04 0.042 0.02 292 0.12 0.95 0.848 0.256 3.2 0.112 0.072 0.038 322 0.052 0.34 0.544 0.176 1.3 0.028 0.04 0.036 303 0.036 1.11 0.584 0.272 0.3 0.036 0.044 0.022 304 0.048 1.21 0.432 0.164 0.3 0.024 0.028 0.02 305 0.032 0.46 0.352 0.14 0.7 0.024 0.022 0.018 380 0.036 0.8 0.336 0.128 0.4 0.032 0.034 0.016 225 0.052 1.11 0.592 0.292 0.9 0.044 0.064 0.03 359 0.02 0.2 0.36 0.176 0.5 0.012 0.024 0.012 226 0.044 2.45 0.36 0.088 1.4 0.024 0.026 0.014 327 0.052 1.36 0.488 0.24 0.2 0.04 0.044 0.02 328 0.06 0.65 0.496 0.196 0.9 0.028 0.036 0.024 369 0.12 1.14 0.52 0.18 2.1 0.06 0.05 0.028 220 0.06 1.82 0.512 0.172 0.8 0.032 0.034 0.02 352 0.108 0.83 0.76 0.26 1.2 0.044 0.076 0.038 353 0.052 0.41 0.624 0.304 0.3 0.036 0.036 0.022 368 0.072 1.04 0.392 0.132 0.3 0.032 0.05 0.02 382 0.072 1.09 0.408 0.16 0.9 0.044 0.054 0.03 325 0.084 0.88 0.648 0.248 0.5 0.048 0.072 0.046 337 0.024 0.2 0.352 0.168 0.1 0.012 0.02 0.016 356 0.136 1 0.728 0.244 0.3 0.056 0.084 0.034 75 0.06 0.43 0.449 0.166 0.5 0.028 0.024 0.016 182 0.04 0.7 0.472 0.204 0.2 0.032 0.036 0.02 189 0.04 0.48 0.392 0.16 0.3 0.032 0.032 0.028 12 0.096 2.67 0.48 0.22 1.5 0.04 0.06 0.03 311 0.092 0.75 0.536 0.172 1.1 0.052 0.062 0.04 171 0.124 4.15 0.776 0.136 4.8 0.088 0.032 0.03 339 0.028 0.34 0.312 0.132 0.6 0.016 0.022 0.012 370 0.056 0.63 0.568 0.204 0.6 0.032 0.054 0.03 371 0.06 1.39 0.552 0.188 2 0.032 0.044 0.022 372 0.048 0.31 0.424 0.12 0.2 0.044 0.038 0.028

Appendix 2. (continued)

373 0.048 0.82 0.344 0.104 0.9 0.028 0.046 0.018 374 0.044 0.41 0.448 0.14 0.2 0.02 0.036 0.016 375 0.04 0.73 0.52 0.172 0.4 0.032 0.042 0.02 376 0.092 0.92 0.84 0.288 1.5 0.072 0.042 0.028 377 0.048 0.66 0.448 0.164 0.5 0.032 0.044 0.018 87 0.068 0.85 0.6 0.244 1 0.044 0.042 0.032 38 0.14 2.55 0.496 0.196 0.3 0.04 0.06 0.02 180 0.036 0.48 0.344 0.196 0.3 0.04 0.05 0.018 361 0.1 0.9 0.792 0.308 4.1 0.052 0.05 0.024 362 0.168 2.33 0.664 0.296 1.6 0.06 0.078 0.032 384 0.056 0.9 0.424 0.224 0.7 0.032 0.038 0.022 21 0.068 2.64 0.44 0.164 0.2 0.036 0.012 0.02 22 0.06 2.21 0.432 0.144 0.7 0.024 0.03 0.018 366 0.096 0.88 0.624 0.22 0.5 0.052 0.07 0.03 367 0.092 0 0.648 0.244 1.3 0.044 0.084 0.022 386 0.02 1.38 0.304 0.144 0.6 0.02 0.022 0.012 30 0.056 1.05 0.456 0.128 3.5 0.032 0.052 0.03 170 0.052 0.87 0.608 0.22 0.9 0.064 0.078 0.022 336 0.048 0.58 0.528 0.188 1.3 0.036 0.038 0.018 378 0.088 1.12 0.52 0.24 1.5 0.068 0.088 0.03 329 0.052 0.87 0.544 0.172 1.4 0.036 0.04 0.036 400 0.064 2.18 0.576 0.208 0.1 0.036 0.05 0.022 46 0.06 0.9 0.384 0.148 0.4 0.024 0.032 0.014 74 0.056 1.11 0.584 0.268 0.5 0.056 0.044 0.03 176 0.084 1.17 0.6 0.176 0.5 0.048 0.03 0.02 185 0.028 0.6 0.464 0.224 0.4 0.028 0.032 0.02 187 0.052 1.14 0.592 0.272 0.8 0.04 0.03 0.018 350 0.048 1.21 0.496 0.276 0.9 0.036 0.026 0.016 50 0.092 1.6 0.6 0.26 1.8 0.04 0.036 0.02 76 0.068 1.67 0.456 0.2 1.3 0.04 0.034 0.022 78 0.024 0.37 0.28 0.1 1.1 0.024 0.018 0.018 143 0.128 0.9 0.912 0.32 2.3 0.052 0.052 0.03 181 0.228 3.5 0.688 0.136 2 0.084 0.07 0.03 186 0.12 1.16 0.8 0.188 0.3 0.084 0.072 0.036 222 0.076 0.95 0.848 0.288 2.1 0.064 0.07 0.04 223 0.092 1.56 0.832 0.248 0.9 0.072 0.044 0.04 332 0.06 1.09 0.568 0.2 1.7 0.044 0.032 0.022 175 0.072 0.66 0.56 0.156 1 0.032 0.044 0.026 179 0.084 0.54 0.6 0.22 1.1 0.06 0.07 0.022 166 0.044 0.82 0.472 0.24 0.5 0.036 0.034 0.018 326 0.048 1.02 0.368 0.16 0.9 0.028 0.038 0.024 55 0.048 0.71 0.52 0.34 3.5 0.032 0.048 0.02 155 0.024 0.58 0.232 0.06 0.2 0.012 0.012 0.014

Appendix 2. (continued)

351 0.04 0.99 0.392 0.168 0.4 0.032 0.036 0.018 18 0.084 3.74 0.64 0.288 0.1 0.04 0.034 0.024 257 0.04 2.07 0.528 0.216 0.4 0.028 0.058 0.016 258 0.08 2.04 0.528 0.156 0.1 0.028 0.03 0.02 333 0.052 1.02 0.448 0.116 0.8 0.024 0.026 0.018 47 0.052 0.32 0.544 0.292 1 0.04 0.038 0.028 354 0.036 0.37 0.496 0.18 1.4 0.028 0.036 0.018 355 0.068 1.19 0.552 0.18 0.2 0.044 0.064 0.034 15 0.2 6.04 0.872 0.34 3 0.08 0.06 0.04 65 0.164 3.08 0.488 0.24 2.6 0.064 0.05 0.048 309 0.068 0.46 0.52 0.236 0.5 0.032 0.058 0.022 62 0.064 1.55 0.536 0.24 0.3 0.04 0.048 0.02 67 0.032 0.27 0.432 0.224 0.2 0.02 0.02 0.012 218 0.02 0.56 0.272 0.064 0.3 0.016 0.034 0.014 219 0.144 1.73 0.72 0.248 0.7 0.052 0.056 0.03 259 0.04 0.82 0.352 0.14 0.5 0.02 0.026 0.016 260 0.032 1.31 0.488 0.224 0.6 0.028 0.038 0.018 261 0.028 1.05 0.456 0.192 0.1 0.02 0.012 0.012 262 0.108 1.21 0.472 0.24 0.4 0.036 0.056 0.022 263 0.052 0.56 0.544 0.22 0.2 0.028 0.03 0.016 264 0.052 1.14 0.696 0.256 0.6 0.032 0.054 0.024 265 0.128 1.38 0.872 0.204 0.4 0.064 0.08 0.04 267 0.1 1.34 0.76 0.28 0.8 0.048 0.05 0.03 268 0.032 0.53 0.352 0.12 0.2 0.02 0.038 0.016 269 0.06 0.85 1 0.396 3.1 0.048 0.08 0.03 270 0.036 0.75 0.408 0.144 0.2 0.028 0.042 0.012 271 0.028 1.29 0.392 0.16 0.3 0.02 0.034 0.012 272 0.032 0.46 0.552 0.22 0.4 0.028 0.044 0.012 273 0.076 3.4 0.648 0.24 0.9 0.04 0.056 0.022 274 0.032 0.54 0.384 0.172 0.6 0.024 0.02 0.012 275 0.088 1.87 0.68 0.228 1.4 0.052 0.078 0.032 276 0.02 0.24 0.248 0.108 0.2 0.016 0.012 0.008 277 0.028 0.6 0.28 0.12 0.4 0.016 0.018 0.01 278 0.08 1.16 0.512 0.176 0.3 0.024 0.036 0.016 279 0.04 1.79 0.344 0.152 0.4 0.02 0.066 0.014 280 0.028 1.11 0.416 0.176 0 0.012 0.026 0.014 281 0.096 1.14 0.736 0.26 0.8 0.036 0.042 0.024 282 0.124 1.04 1.032 0.32 2.7 0.08 0.07 0.038 283 0.12 1.56 0.648 0.136 2 0.044 0.04 0.026 284 0.072 1.63 0.504 0.136 0.1 0.052 0.048 0.018 286 0.056 0.88 0.552 0.16 0.5 0.028 0.056 0.02 288 0.056 0.53 0.688 0.268 0.4 0.036 0.064 0.022 290 0.044 0.46 0.408 0.136 0.1 0.032 0.034 0.012

Appendix 2. (continued)

291 0.032 0.88 0.544 0.224 0.1 0.028 0.04 0.02 310 0.152 1.33 0.672 0.3 0.4 0.052 0.052 0.032 360 0.072 1.51 0.584 0.244 0.3 0.048 0.06 0.026 358 0.024 0.48 0.424 0.168 0.2 0.04 0.056 0.024 49 0.076 1.68 0.816 0.26 0.9 0.036 0.046 0.024 61 0.1 1.24 0.792 0.16 2.3 0.064 0.042 0.026 64 0.048 1.73 0.472 0.16 2 0.024 0.062 0.024 48 0.1 2.65 0.664 0.196 2.5 0.048 0.042 0.038 52 0.056 0.44 0.36 0.204 1 0.048 0.032 0.022 178 0.076 1.84 0.64 0.264 0.9 0.052 0.028 0.024 1 0.064 2.47 0.808 0.26 1.1 0.048 0.046 0.024 4 0.072 0.54 0.934 0.164 2.7 0.068 0.048 0.034 6 0.084 1.02 0.4 0.136 3.2 0.04 0.04 0.024 7 0.116 2.72 0.44 0.088 4.5 0.1 0.05 0.044 8 0.042 1.11 0.496 0.2 2.8 0.076 0.06 0.042 36 0.06 1.72 0.544 0.136 1.2 0.036 0.034 0.03 42 0.036 1.34 0.432 0.136 0.5 0.02 0.022 0.016 51 0.036 2.11 0.312 0.124 1.3 0.032 0.026 0.02 330 0.076 2.84 0.552 0.212 1.3 0.04 0.034 0.022 13 0.08 4 0.448 0.156 1.4 0.044 0.058 0.03 165 0.044 0.34 0.4 0.156 0.4 0.024 0.028 0.014 80 0.044 0.48 0.408 0.172 0.2 0.02 0.032 0.016 56 0.056 1.43 0.528 0.236 2 0.048 0.054 0.03 357 0.072 0.36 0.576 0.232 0.4 0.028 0.034 0.024 73 0.14 1.36 0.584 0.168 0.9 0.064 0.062 0.024 334 0.068 0 0.56 0.192 0.2 0.032 0.022 0.022 389 0.096 2.33 0.568 0.1 1.6 0.048 0.054 0.03 390 0.048 0.97 0.456 0.236 0.3 0.024 0.028 0.024 391 0.092 0.82 0.624 0.224 0.8 0.048 0.05 0.036 335 0.072 1.38 0.72 0.348 1.8 0.06 0.08 0.03 379 0.052 0.78 0.4 0.08 0.7 0.04 0.036 0.018 392 0.116 1.38 0.608 0.224 0.8 0.04 0.064 0.03 29 0.136 2.21 0.344 0.166 2 0.052 0.06 0.052 153 0.112 1.84 0.48 0.196 0.4 0.036 0.04 0.03 154 0.1 1.38 0.576 0.228 0.7 0.048 0.05 0.034 341 0.024 1.51 0.2 0.076 0.6 0.016 0.04 0.016 342 0.012 0.48 0.312 0.18 0.5 0.012 0.022 0.008 343 0.02 0.92 0.28 0.1 1.3 0.02 0.022 0.014 344 0.04 0.32 0.312 0.152 0.6 0.02 0.02 0.01 345 0.04 0.54 0.4 0.164 0.4 0.048 0.034 0.016 346 0.024 0.2 0.296 0.128 0.3 0.016 0.024 0.01 347 0.032 1.43 0.328 0.128 1.6 0.02 0.03 0.018 348 0.036 1.09 0.248 0.112 0.4 0.02 0.02 0.016

Appendix 2. (continued) 349 0.06 1.96 0.336 0.136 0.9 0.032 0.042 0.022 393 0.108 1.48 0.48 0.164 1.6 0.048 0.06 0.022 394 0.076 1.58 0.624 0.276 0.5 0.032 0.04 0.024 395 0.104 1.75 0.616 0.24 0.4 0.052 0.03 0.04 53 0.044 0.54 0.408 0.172 0.4 0.032 0.028 0.016 338 0.032 0.65 0.216 0.108 0.2 0.016 0.03 0.014 381 0.048 1.36 0.496 0.208 0.3 0.036 0.042 0.018 396 0.028 0.39 0.36 0.12 0.6 0.016 0.028 0.01 397 0.04 0.97 0.344 0.16 1 0.02 0.048 0.014 398 0.028 0.24 0.216 0.064 0.2 0.016 0.014 0.008 399 0.012 0.51 0.168 0.06 0.5 0.016 0.02 0.01 401 0.036 0.87 0.288 0.08 0.2 0.024 0.036 0.014 363 0.064 0.65 0.68 0.264 1.2 0.044 0.028 0.03 364 0.052 1.11 0.456 0.156 0.5 0.036 0.046 0.026 365 0.152 1.67 0.8 0.248 1.1 0.068 0.066 0.046 101 0.04 1.39 0.48 0.136 1.6 0.024 0.028 0.016 383 0.056 0.34 0.392 0.088 0.2 0.028 0.042 0.016 318 0.092 0.94 0.56 0.168 0.2 0.044 0.036 0.02 151 0.044 1.46 0.4 0.16 0.6 0.036 0.046 0.022 33 0.044 0.95 0.296 0.116 1 0.016 0.036 0.018 77 0.088 2.77 0.6 0.26 1.1 0.052 0.052 0.03 88 0.032 0.51 0.304 0.076 0.4 0.024 0.044 0.018 146 0.032 0.71 0.32 0.12 0.9 0.028 0.042 0.024 164 0.028 0.51 0.36 0.16 0.3 0.02 0.026 0.014 388 0.12 1.28 0.6 0.228 1.3 0.052 0.084 0.028 149 0.032 0.6 0.36 0.132 0.4 0.024 0.02 0.014 3 0.048 0.31 0.368 0.22 0.1 0.02 0.04 0.012 84 0.02 0.29 0.296 0.14 0.2 0.02 0.026 0.016 91 0.116 1.48 0.664 0.172 0.3 0.06 0.046 0.03 92 0.02 0.97 0.296 0.1 0.4 0.02 0.014 0.014 152 0.04 0.43 0.48 0.124 0.2 0.032 0.026 0.024 232 0.06 0 0.448 0.2 0.4 0.028 0.056 0.016 320 0.068 0.92 0.568 0.22 1.2 0.044 0.032 0.026 321 0.076 0.56 0.68 0.324 1.8 0.048 0.044 0.02 200 0.06 0.05 0.6 0.236 0.3 0.04 0.046 0.026 195 0.044 0.41 0.384 0.16 0.2 0.02 0.03 0.016 201 0.04 0.66 0.384 0.172 0.2 0.04 0.034 0.022 324 0.064 1.24 0.496 0.208 2.3 0.032 0.046 0.024 32 0.108 2.75 0.52 0.22 0.5 0.06 0.052 0.026 94 0.048 0.63 0.464 0.128 0.2 0.024 0.01 0.016 157 0.04 0.36 0.408 0.168 0.2 0.028 0.03 0.018 193 0.064 0 0.4 0.112 0.5 0.036 0.042 0.022 199 0.064 0.78 0.608 0.24 3 0.036 0.022 0.04

Appendix 2. (continued) 387 0.076 0.43 0.76 0.364 2 0.052 0.03 0.03 95 0.06 2.47 0.568 0.212 0.4 0.048 0.044 0.03 215 0.056 0.29 0.392 0.16 0.3 0.032 0.026 0.02 97 0.076 0.71 0.408 0.2 1.9 0.04 0.042 0.022 190 0.048 0.41 0.344 0.104 0.7 0.032 0.022 0.022 203 0.08 1.48 0.472 0.24 0.4 0.048 0.05 0.032 205 0.044 0.53 0.448 0.14 0.2 0.028 0.022 0.016 207 0.028 0.75 0.448 0.212 0.2 0.016 0.03 0.018 323 0.068 1.16 0.472 0.24 2.2 0.044 0.044 0.024 217 0.06 1.9 0.44 0.176 0.1 0.048 0.044 0.036 98 0.096 1.33 0.408 0.144 0.2 0.032 0.05 0.024 99 0.104 2.31 0.552 0.448 1.1 0.048 0.052 0.03 191 0.06 2.14 0.52 0.244 0.7 0.036 0.032 0.022 54 0.08 2.96 0.504 0.184 2.6 0.036 0.036 0.014 93 0.044 1.12 0.464 0.224 1 0.028 0.054 0.018 96 0.056 0.8 0.48 0.212 1 0.04 0.03 0.02 100 0.068 0.99 0.456 0.256 0.8 0.04 0.058 0.04 103 0.06 1.19 0.424 0.12 3.7 0.04 0.04 0.034 194 0.052 0.24 0.472 0.144 0.6 0.04 0.044 0.022 196 0.032 0.43 0.392 0.184 0.4 0.02 0.048 0.016 197 0.028 0.51 0.248 0.088 0.7 0.024 0.03 0.016 202 0.072 0 0.416 0.156 0.4 0.028 0.032 0.024 204 0.04 0.41 0.328 0.148 0.2 0.036 0.036 0.026 208 0.108 1.5 0.504 0.208 0.3 0.06 0.066 0.02 210 0.072 0.83 0.44 0.204 0.1 0.032 0.064 0.018 212 0.076 1.19 0.608 0.284 0.3 0.044 0.034 0.03 213 0.048 1.39 0.416 0.196 0.6 0.024 0.042 0.016 216 0.072 0.78 0.52 0.2 1 0.044 0.064 0.034 307 0.048 0 0.448 0.228 0.4 0.036 0.02 0.018 308 0.04 0 0.36 0.1 0.4 0.036 0.038 0.022 312 0.036 1.53 0.392 0.2 0.4 0.024 0.05 0.016 313 0.032 0.41 0.456 0.232 0.5 0.02 0.034 0.018 314 0.044 0.54 0.416 0.18 0.2 0.028 0.044 0.016 315 0.076 2.09 0.56 0.16 3.1 0.06 0.03 0.022 316 0.06 0.63 0.48 0.18 0.5 0.036 0.052 0.03 317 0.044 0.7 0.464 0.26 0.3 0.036 0.036 0.026 385 0.036 0.41 0.424 0.18 0.3 0.036 0.032 0.02 228 0.088 0.7 0.488 0.208 0.1 0.06 0.042 0.028 224 0.072 2.47 0.624 0.244 0.4 0.024 0.058 0.02 231 0.02 0.6 0.424 0.18 0.4 0.024 0.044 0.016 188 0.12 2.18 0.552 0.312 0.3 0.044 0.062 0.028 293 0.056 0.58 0.528 0.208 0.7 0.036 0.026 0.022 294 0.112 1.05 0.648 0.28 0.5 0.048 0.032 0.028

Appendix 2. (continued) 295 0.108 1.29 0.592 0.224 0.4 0.052 0.05 0.032 5 0.104 1.77 0.688 0.34 0.9 0.036 0.048 0.026 233 0.032 1.89 0.376 0.2 1 0.024 0.024 0.014 85 0.048 0.88 0.392 0.168 0.3 0.048 0.028 0.026 89 0.048 1.09 0.36 0.112 0.3 0.036 0.024 0.028 31 0.068 1.04 0.464 0.172 0.9 0.044 0.034 0.026 331 0.096 2.48 0.736 0.324 2.4 0.044 0.05 0.03 296 0.088 2.53 0.624 0.284 1.1 0.044 0.05 0.018 298 0.084 1.5 0.768 0.364 1.6 0.06 0.044 0.032 300 0.1 0 0.584 0.2 0.3 0.068 0.028 0.028 9 0.064 3.16 0.4 0.2 0.9 0.044 0.042 0.024 230 0.072 1.41 0.44 0.16 0.2 0.028 0.046 0.018 299 0.104 2.26 0.608 0.212 1.7 0.048 0.044 0.028 302 0.096 1.72 0.624 0.344 0.7 0.06 0.066 0.042 297 0.072 0.77 0.496 0.204 1.1 0.04 0.036 0.018 19 0.044 1 0.376 0.196 1.5 0.04 0.04 0.022 20 0.04 2.21 0.36 0.136 1.2 0.028 0.04 0.016 23 0.056 1.62 0.408 0.28 0.5 0.02 0.036 0.014 24 0.08 2.07 0.424 0.22 0.2 0.028 0.014 0.012 301 0.08 2.96 0.688 0.224 0.1 0.044 0.054 0.02

Appendix 2. (continued)

Number Leaflet Leaflet Leaflet Head Flower Peduncle Peduncle Involucre Base Length Width Length Number Length Diameter Radius Angle (cm) (cm) (cm) (cm) (cm) (cm) (degrees) 167 60 0.552 0.384 0.83 5 1.5 0.034 0.344 70 40 0.544 0.288 0.83 7 1.5 0.02 0.336 306 30 0.376 0.144 0.6 3 0.6 0.02 0.224 424 30 0.552 0.232 0.68 10 1.9 0.02 0.296 425 30 0.504 0.16 0.65 12 1.4 0.024 0.216 319 30 1.016 0.416 0.77 10 4.3 0.038 0.368 69 30 0.616 0.24 0.71 6 2.2 0.032 0.368 229 40 0.808 0.4 0.82 9 3.8 0.036 0.36 26 50 0.976 0.48 0.92 18 4 0.034 0.384 41 30 0.8 0.336 0.66 6 3.4 0.03 0.296 292 70 1.824 1.136 0.83 9 2.8 0.068 0.408 322 60 0.64 0.44 0.66 5 1.5 0.03 0.344 303 40 0.752 0.44 0.75 7 2.4 0.036 0.408 304 30 0.728 0.28 0.83 10 1.3 0.024 0.368 305 40 0.744 0.464 0.71 8 2.1 0.026 0.304 380 30 0.624 0.328 0.65 4 1.8 0.026 0.28 225 50 1.064 0.68 1.14 12 2.9 0.044 0.36 359 30 0.368 0.208 0.53 2 0.9 0.012 0.288 226 40 0.576 0.328 0.88 5 4.8 0.024 0.368 327 30 1.128 0.584 1.14 9 3.7 0.056 0.416 328 50 0.728 0.384 0.94 11 2.5 0.03 0.376 369 50 1 0.536 1.19 15 4.3 0.086 0.424 220 30 0.848 0.384 1.16 9 3.5 0.026 0.384 352 40 1.752 0.664 1.04 25 3.1 0.056 0.544 353 30 0.96 0.28 1.33 19 4 0.04 0.472 368 50 0.792 0.408 1.16 12 3.3 0.038 0.392 382 40 0.912 0.6 1.39 17 4.1 0.048 0.328 325 20 1.52 0.528 1.39 23 3.6 0.054 0.44 337 20 0.616 0.176 0.75 3 1.3 0.018 0.288 356 30 2.088 0.96 1.24 13 2.9 0.06 0.512 75 30 1.04 0.296 0.61 9 2.3 0.036 0.288 182 30 0.68 0.256 0.82 11 1.2 0.028 0.36 189 30 0.608 0.232 0.7 10 0.9 0.03 0.352 12 30 1.136 0.368 0.58 5 1.9 0.036 0.24 311 40 1.168 0.528 0.88 11 3 0.044 0.384 171 70 1.344 0.84 1.05 16 7.3 0.08 0.648 339 30 0.488 0.216 0.78 2 0.9 0.02 0.264 370 40 0.96 0.536 1.11 8 2.9 0.044 0.432 371 60 0.936 0.592 1.24 13 3.9 0.042 0.36

Appendix 2. (continued)

372 70 0.584 0.368 0.73 8 1.7 0.034 0.336 373 20 1.008 0.176 0.95 9 2.8 0.034 0.256 374 30 0.696 0.28 1.11 10 2.5 0.034 0.352 375 40 1.12 0.48 0.87 6 1.8 0.036 0.344 376 70 1.936 1.072 1.22 14 3 0.08 0.616 377 30 1.24 0.408 0.83 9 2.7 0.04 0.408 87 40 1.032 0.488 1.46 20 3.9 0.064 0.536 38 30 1.648 0.512 0.73 16 3.1 0.06 0.344 180 20 0.728 0.128 0.49 3 1.6 0.016 0.256 361 60 1.44 0.736 1.29 10 5.3 0.054 0.44 362 40 2.024 0.84 1.36 17 3.9 0.08 0.64 384 20 0.872 0.192 0.82 2 1.7 0.03 0.312 21 30 0.808 0.328 1.05 15 2.9 0.056 0.36 22 30 0.672 0.32 0.88 13 2.5 0.044 0.36 366 30 2.592 0.512 1.24 15 4.5 0.05 0.528 367 30 1.752 0.488 0.85 14 3.2 0.038 0.408 386 30 0.448 0.256 0.73 1 1.2 0.018 0.264 30 30 1.016 0.464 1.62 16 6.1 0.052 0.496 170 30 1.184 0.528 1.34 13 3.5 0.06 0.464 336 30 1.088 0.536 1.21 12 2.8 0.05 0.44 378 40 1.504 0.84 1.67 21 5 0.114 0.576 329 60 1 0.52 1.39 16 3.3 0.048 0.44 400 30 1.056 0.384 0.88 9 2.1 0.036 0.36 46 20 0.681 0.248 0.78 3 2.7 0.026 0.288 74 40 1.312 0.552 1.7 17 4.4 0.076 0.496 176 40 1.192 0.464 1.36 12 3.5 0.062 0.56 185 30 0.624 0.256 0.73 11 1.8 0.022 0.36 187 40 0.912 0.344 0.87 12 1.7 0.034 0.432 350 20 0.656 0.24 0.75 10 3.3 0.024 0.28 50 40 1.52 0.584 1.21 13 4.8 0.048 0.6 76 40 0.864 0.48 0.61 9 2.8 0.03 0.224 78 50 0.68 0.36 0.63 7 1.9 0.018 0.208 143 50 1.4 0.64 1.02 22 1.8 0.072 0.432 181 50 2.896 0.896 0.88 18 5.3 0.05 0.424 186 50 2.304 1.04 0.85 24 2.9 0.064 0.448 222 80 1.872 0.92 0.82 12 1.3 0.034 0.6 223 40 1.304 0.768 1.17 21 2.8 0.082 0.672 332 40 0.912 0.496 0.73 7 2.4 0.028 0.288 175 40 1.216 0.432 0.68 9 1.7 0.038 0.376 179 50 1.36 0.592 0.94 9 2.2 0.05 0.424 166 40 1.008 0.416 0.78 11 2.2 0.036 0.36 326 30 0.648 0.304 1.11 6 2.7 0.026 0.336 55 60 0.68 0.44 1.02 11 6.1 0.058 0.408

Appendix 2. (continued) 155 30 0.2 0.096 0.56 2 0.9 0.012 0.248 351 40 0.472 0.264 0.65 3 0.9 0.03 0.264 18 40 1.36 0.467 0.85 12 3 0.052 0.392 257 30 0.928 0.432 0.71 15 2.9 0.04 0.296 258 30 0.608 0.296 0.92 13 3.2 0.038 0.392 333 40 0.56 0.312 0.63 6 1.5 0.022 0.312 47 40 0.952 0.52 1.28 17 3 0.046 0.536 354 60 0.6 0.424 1.28 10 2 0.036 0.432 355 40 1.336 0.584 0.97 12 3 0.038 0.504 15 20 1.32 0.384 0.77 8 4 0.044 0.376 65 50 1.616 0.672 0.83 11 4.6 0.052 0.312 309 30 1.32 0.552 0.66 8 2.2 0.022 0.352 62 20 1.216 0.288 0.7 10 3.3 0.038 0.352 67 30 0.448 0.16 0.6 2 0.9 0.028 0.32 218 30 0.328 0.16 0.6 2 1.1 0.016 0.216 219 50 1.784 0.64 1.19 23 4.7 0.078 0.56 259 30 0.512 0.112 0.54 3 1.3 0.02 0.24 260 40 0.584 0.288 0.61 7 2.7 0.028 0.288 261 40 0.472 0.192 0.61 8 1.9 0.024 0.248 262 30 0.96 0.4 0.77 18 2.3 0.04 0.336 263 30 0.64 0.256 0.7 13 1.6 0.028 0.4 264 30 1.12 0.432 0.58 9 3.5 0.028 0.328 265 50 1.92 0.976 1.28 9 4.1 0.068 0.544 267 30 1.592 0.624 0.75 10 2.9 0.046 0.304 268 10 0.648 0.104 0.77 7 0.9 0.03 0.256 269 50 1.68 0.608 1.14 15 5 0.058 0.632 270 30 0.64 0.272 0.7 9 3 0.02 0.304 271 30 0.472 0.176 0.66 2 1.6 0.022 0.28 272 30 0.848 0.28 0.77 8 2.3 0.02 0.352 273 30 1.344 0.456 0.78 19 3.3 0.048 0.44 274 50 0.272 0.192 0.56 3 0.4 0.02 0.28 275 30 1.36 0.44 0.83 13 4.1 0.05 0.376 276 30 0.28 0.128 0.61 2 1.1 0.02 0.232 277 20 0.24 0.136 0.51 3 1.2 0.02 0.232 278 30 0.568 0.232 0.7 14 1.1 0.03 0.336 279 20 0.816 0.304 0.68 11 3.1 0.028 0.232 280 20 0.416 0.112 0.56 2 1.7 0.02 0.312 281 30 1.04 0.496 0.83 15 2.7 0.042 0.464 282 40 1.824 0.768 1.09 16 4.4 0.062 0.512 283 50 1.304 0.704 1.26 11 5 0.062 0.464 284 20 1.072 0.32 0.83 14 4 0.04 0.32 286 20 0.936 0.184 0.78 20 2.4 0.034 0.328 288 30 1.176 0.36 0.85 13 1.2 0.052 0.488

Appendix 2. (continued) 290 20 0.608 0.232 0.71 13 1.8 0.032 0.328 291 20 0.792 0.208 0.87 11 2.7 0.034 0.312 310 30 1.848 0.752 0.88 10 3.7 0.052 0.432 360 30 1.392 0.44 0.8 15 3 0.044 0.384 358 40 0.96 0.44 1.39 15 4 0.068 0.36 49 30 1.39 0.536 0.9 18 3.2 0.042 0.328 61 30 1.488 0.672 1.11 13 3.3 0.054 0.408 64 60 0.848 0.544 0.85 8 4.6 0.032 0.312 48 40 1.128 0.608 0.94 13 5 0.052 0.496 52 50 0.912 0.528 0.83 11 3.8 0.042 0.568 178 40 1.296 0.464 0.94 18 3.3 0.05 0.44 1 40 1.168 0.592 1.26 25 2.8 0.052 0.48 4 60 1.288 0.76 1.02 15 2.5 0.046 0.456 6 60 0.664 0.472 1.28 10 3.6 0.04 0.36 7 70 1.744 0.992 1.29 16 5.5 0.07 0.336 8 60 1.112 0.752 1.12 17 6.2 0.076 0.624 36 40 0.984 0.544 1.19 17 2.7 0.052 0.472 42 40 0.496 0.32 0.83 9 2 0.03 0.296 51 30 0.848 0.416 0.58 8 2.8 0.026 0.288 330 50 0.832 0.376 0.68 8 2 0.022 0.368 13 50 0.72 0.416 1.02 4 3.5 0.032 0.4 165 30 0.496 0.184 0.68 11 1.2 0.034 0.264 80 20 0.752 0.328 1.11 5 1 0.022 0.36 56 50 0.84 0.432 0.94 15 4.2 0.04 0.392 357 40 1.008 0.344 0.85 11 1.6 0.048 0.48 73 60 1.72 0.832 1.48 25 4.8 0.128 0.52 334 30 0.792 0.24 0.85 15 1.3 0.038 0.424 389 40 1.04 0.504 0.94 15 3.3 0.044 0.496 390 20 0.776 0.24 0.77 14 2.1 0.038 0.4 391 30 1.416 0.672 0.82 14 2.9 0.056 0.432 335 30 1.968 0.472 1.02 14 4.7 0.05 0.496 379 30 0.912 0.4 1.17 15 3.3 0.054 0.376 392 30 1.2 0.344 1.34 10 3.6 0.06 0.504 29 50 1.928 0.8 1.36 19 4 0.064 0.448 153 40 1.168 0.432 1.43 17 6.1 0.08 0.464 154 20 1.56 0.256 0.73 14 4 0.068 0.368 341 20 0.344 0.096 0.66 2 0.9 0.018 0.24 342 30 0.456 0.176 0.48 2 1.5 0.016 0.256 343 30 0.376 0.176 0.71 2 1.8 0.02 0.248 344 20 0.28 0.112 0.56 2 0.8 0.014 0.296 345 60 0.68 0.376 0.78 5 1.6 0.02 0.344 346 20 0.496 0.168 0.77 3 1.2 0.018 0.232 347 70 0.456 0.416 0.78 2 1.8 0.018 0.296

Appendix 2. (continued) 348 30 0.544 0.152 0.65 7 2.3 0.026 0.168 349 20 0.608 0.192 0.82 4 3.3 0.024 0.32 393 30 1.52 0.464 1.34 13 4.8 0.064 0.424 394 40 0.752 0.288 1.28 16 2.5 0.046 0.424 395 30 1.248 0.488 1.33 16 4.4 0.078 0.408 53 40 0.616 0.352 0.78 4 2 0.036 0.296 338 30 0.296 0.176 0.66 3 1.2 0.02 0.208 381 30 0.856 0.328 0.8 3 1.6 0.018 0.368 396 40 0.408 0.272 0.71 2 1.1 0.014 0.272 397 30 0.736 0.288 0.97 7 2.4 0.028 0.248 398 30 0.336 0.184 0.73 4 0.7 0.026 0.256 399 30 0.256 0.12 0.7 1 1.3 0.01 0.152 401 30 0.88 0.256 0.85 8 2 0.018 0.304 363 40 1.128 0.56 1.12 8 3.5 0.042 0.552 364 50 1.376 0.632 1.24 13 3.2 0.032 0.424 365 70 1.952 1.096 1.53 20 5 0.098 0.56 101 30 0.512 0.216 0.65 8 1.2 0.022 0.288 383 30 0.84 0.296 0.92 9 1.9 0.034 0.384 318 20 1.152 0.296 0.95 12 3.1 0.05 0.392 151 40 1.088 0.416 0.77 11 2.5 0.022 0.32 33 40 0.52 0.256 0.66 2 1.8 0.014 0.264 77 40 1.416 0.544 0.71 14 2.5 0.034 0.328 88 30 0.696 0.288 0.63 8 1.2 0.02 0.272 146 40 0.68 0.336 0.66 3 1.7 0.02 0.28 164 30 0.576 0.312 0.65 6 1.4 0.022 0.328 388 40 1.816 0.704 0.7 10 3.4 0.044 0.32 149 40 0.592 0.256 0.7 5 2 0.02 0.36 3 20 0.64 0.2 0.83 9 1.3 0.034 0.344 84 30 0.4 0.16 0.61 6 1.1 0.016 0.256 91 30 1.92 0.488 1.11 17 3.3 0.05 0.44 92 20 0.528 0.24 0.51 4 1.9 0.014 0.24 152 20 1.04 0.264 0.78 7 2.4 0.026 0.392 232 20 0.832 0.28 0.71 7 1.3 0.024 0.304 320 30 1.12 0.424 0.8 7 2.3 0.03 0.376 321 20 1.56 0.48 0.65 10 1.8 0.04 0.408 200 20 0.928 0.272 0.73 8 2.8 0.036 0.344 195 40 0.56 0.208 0.65 10 1.2 0.03 0.28 201 20 0.752 0.16 0.88 15 2.5 0.038 0.272 324 50 1.152 0.592 0.83 9 3.4 0.018 0.352 32 30 1.152 0.376 1 15 2.8 0.054 0.472 94 50 0.352 0.2 0.63 6 0.7 0.034 0.336 157 30 0.552 0.256 0.73 4 1.9 0.04 0.4 193 50 0.952 0.528 0.65 9 0.8 0.022 0.328

Appendix 2. (continued) 199 50 0.704 0.464 0.53 2 2.1 0.02 0.368 387 50 1.16 0.648 0.7 3 2.3 0.046 0.408 95 30 0.96 0.336 0.87 7 2 0.038 0.328 215 30 0.64 0.232 0.85 9 0.8 0.032 0.312 97 30 0.928 0.32 0.63 4 2.6 0.028 0.312 190 40 0.584 0.312 0.61 2 0.7 0.018 0.24 203 40 0.872 0.264 0.99 7 1.4 0.054 0.464 205 20 0.656 0.208 0.46 8 2.1 0.028 0.344 207 40 0.704 0.352 0.7 8 1.7 0.022 0.312 323 30 0.976 0.4 0.7 6 2.6 0.036 0.376 217 30 1.088 0.312 0.75 10 2.2 0.036 0.36 98 30 0.728 0.352 0.87 10 1.6 0.036 0.336 99 50 0.76 0.392 0.78 13 1.8 0.032 0.384 191 40 0.968 0.448 0.85 15 2.6 0.034 0.288 54 40 1.136 0.464 0.78 7 3.7 0.03 0.296 93 40 0.944 0.496 0.87 8 3.1 0.024 0.328 96 30 0.8 0.288 0.94 8 2.2 0.044 0.4 100 40 1.24 0.56 0.92 10 3.5 0.024 0.376 103 60 0.984 0.56 0.75 3 4 0.038 0.32 194 30 0.8 0.28 0.83 6 1 0.028 0.28 196 50 0.96 0.416 0.7 8 2.4 0.03 0.288 197 30 0.4 0.208 0.58 4 0.9 0.016 0.264 202 30 0.624 0.208 0.9 11 1.7 0.038 0.32 204 40 0.8 0.288 0.78 6 2 0.022 0.408 208 30 0.928 0.232 0.85 8 2.4 0.044 0.424 210 20 0.848 0.288 0.73 5 2 0.026 0.368 212 30 1.2 0.384 0.87 18 2.1 0.044 0.408 213 30 0.928 0.32 0.85 7 2.3 0.028 0.256 216 20 1.56 0.408 0.8 12 2.6 0.018 0.328 307 30 0.384 0.168 0.58 2 0.7 0.026 0.392 308 30 0.712 0.288 0.77 10 1.3 0.028 0.296 312 20 0.744 0.232 0.8 9 2.3 0.026 0.352 313 30 0.512 0.264 0.7 2 1.3 0.02 0.32 314 20 0.824 0.232 0.82 10 1.8 0.024 0.336 315 60 0.424 0.408 0.73 2 2.1 0.026 0.352 316 40 1.256 0.712 0.82 9 2.3 0.042 0.4 317 30 0.656 0.28 0.8 9 1.4 0.026 0.344 385 20 0.568 0.208 0.7 4 1.3 0.032 0.28 228 20 0.816 0.232 0.97 14 1.4 0.05 0.368 224 30 0.824 0.328 0.83 10 3.3 0.034 0.344 231 30 0.68 0.288 0.58 5 2 0.014 0.264 188 30 0.96 0.408 1.31 13 3.6 0.068 0.44 293 40 0.792 0.304 0.99 9 2.3 0.036 0.432

Appendix 2. (continued) 294 30 1.288 0.408 1.19 16 4.1 0.052 0.504 295 30 1.536 0.592 1.22 13 3.6 0.048 0.432 5 40 1.44 0.68 0.95 22 3.8 0.032 0.48 233 30 0.592 0.272 0.71 3 2.1 0.018 0.288 85 60 0.568 0.368 0.85 5 1.2 0.024 0.376 89 40 0.672 0.392 0.92 12 2 0.04 0.392 31 30 0.912 0.328 0.7 10 2 0.04 0.328 331 50 1.144 0.8 0.83 9 3.6 0.048 0.384 296 40 1.112 0.568 0.8 7 3.3 0.026 0.352 298 40 1.424 0.632 0.77 6 3.6 0.032 0.496 300 30 1.16 0.44 0.95 11 1.8 0.046 0.392 9 30 0.976 0.416 0.85 10 4 0.03 0.352 230 30 0.896 0.368 1.02 11 3.6 0.03 0.328 299 40 1.024 0.456 0.85 6 2.9 0.038 0.36 302 30 1.288 0.472 0.68 12 3 0.05 0.472 297 60 1.232 0.728 0.82 7 2 0.032 0.48 19 40 0.792 0.328 0.65 3 1.1 0.026 0.24 20 40 0.608 0.288 0.77 3 1.8 0.024 0.28 23 40 0.56 0.216 0.7 8 1.3 0.03 0.24 24 30 0.84 0.352 0.77 16 2.3 0.036 0.288 301 30 1.456 0.448 0.88 12 3.3 0.03 0.44

Appendix 2. (continued) Number Involucral Calyx Calyx Calyx Calyx Calyx Calyx Banner Lobe Length Circumference Lobe Lobe Tube Lobe Petal Length (cm) (cm) Length Width Length Tooth Length (cm) (cm) (cm) (cm) Number (cm) 167 0.136 0.432 0.24 0.28 0.04 0.136 0 0.7 70 0.192 0.52 0.272 0.336 0.048 0.176 0 0.75 306 0.096 0.336 0.216 0.184 0.04 0.144 1 0.49 424 0.104 0.4 0.296 0.248 0.056 0.136 6 0.53 425 0.096 0.368 0.24 0.232 0.04 0.136 2 0.54 319 0.16 0.408 0.216 0.24 0.048 0.176 0 0.65 69 0.2 0.48 0.304 0.336 0.064 0.152 0 0.61 229 0.104 0.416 0.232 0.256 0.048 0.168 0 0.66 26 0.216 0.48 0.192 0.256 0.048 0.192 0 0.75 41 0.112 0.4 0.256 0.24 0.056 0.136 0 0.54 292 0.096 0.544 0.256 0.336 0.056 0.2 0 0.73 322 0.152 0.44 0.216 0.28 0.048 0.168 1 0.56 303 0.168 0.424 0.232 0.272 0.048 0.144 0 0.63 304 0.176 0.488 0.24 0.32 0.048 0.168 0 0.75 305 0.16 0.488 0.28 0.312 0.056 0.16 2 0.77 380 0.128 0.4 0.208 0.256 0.04 0.144 3 0.61 225 0.144 0.528 0.296 0.304 0.072 0.232 0 0.73 359 0.168 0.4 0.28 0.216 0.048 0.184 1 0.66 226 0.16 0.448 0.288 0.448 0.056 0.168 0 0.77 327 0.208 0.448 0.264 0.288 0.056 0.16 0 0.78 328 0.128 0.432 0.224 0.272 0.032 0.152 0 0.63 369 0.192 0.544 0.408 0.336 0.056 0.208 0 1.14 220 0.128 0.592 0.312 0.36 0.064 0.24 0 1.11 352 0.232 0.6 0.336 0.408 0.064 0.2 0 0.87 353 0.232 0.672 0.36 0.432 0.08 0.24 0 1.14 368 0.16 0.568 0.352 0.4 0.064 0.2 0 1.09 382 0.112 0.616 0.376 0.4 0.08 0.216 0 1.38 325 0.2 0.352 0.304 0.384 0.056 0.208 0 1.12 337 0.112 0.384 0.304 0.248 0.048 0.144 1 0.68 356 0.216 0.448 0.307 0.448 0.056 0.232 0 1 75 0.128 0.424 0.152 0.264 0.032 0.152 0 0.56 182 0.112 0.456 0.208 0.28 0.032 0.192 0 0.58 189 0.128 0.392 0.184 0.232 0.032 0.16 0 0.56 12 0.16 0.44 0.376 0.232 0.08 0.16 0 0.6 311 0.152 0.472 0.248 0.264 0.064 0.208 0 0.65 171 0.2 0.544 0.28 0.328 0.048 0.2 0 1.17 339 0.112 0.44 0.264 0.256 0.064 0.168 2 0.66 370 0.168 0.64 0.304 0.416 0.056 0.232 0 1.04 371 0.128 0.52 0.256 0.328 0.048 0.192 0 1

Appendix 2. (continued) 372 0.128 0.472 0.344 0.288 0.064 0.192 0 0.92 373 0.112 0.432 0.248 0.264 0.048 0.176 0 0.85 374 0.144 0.464 0.248 0.264 0.04 0.184 0 0.94 375 0.152 0.472 0.272 0.264 0.048 0.192 0 0.71 376 0.208 0.664 0.344 0.408 0.056 0.248 0 1.33 377 0.168 0.504 0.344 0.312 0.064 0.2 7 0.73 87 0.24 0.712 0.336 0.392 0.056 0.312 0 1.46 38 0.264 0.496 0.272 0.288 0.048 0.168 0 0.6 180 0.112 0.456 0.2 0.264 0.04 0.176 0 0.56 361 0.152 0.64 0.256 0.432 0.048 0.2 0 1.16 362 0.272 0.728 0.352 0.48 0.072 0.232 0 1.22 384 0.184 0.368 0.216 0.2 0.04 0.168 1 0.56 21 0.192 0.488 0.288 0.304 0.08 0.192 0 0.83 22 0.216 0.464 0.232 0.28 0.056 0.184 1 0.85 366 0.264 0.6 0.288 0.384 0.04 0.216 0 0.99 367 0.176 0.536 0.312 0.344 0.072 0.16 0 0.77 386 0.176 0.392 0.232 0.224 0.04 0.152 0 0.7 30 0.168 0.656 0.416 0.36 0.096 0.28 1 1.26 170 0.152 0.568 0.32 0.304 0.048 0.256 4 1.09 336 0.16 0.44 0.36 0.216 0.088 0.232 1 0.9 378 0.216 0.736 0.464 0.44 0.088 0.28 0 1.58 329 0.192 0.576 0.312 0.336 0.064 0.24 0 1.11 400 0.16 0.528 0.264 0.32 0.048 0.184 0 0.78 46 0.16 0.472 0.288 0.272 0.072 0.184 1 0.61 74 0.288 0.696 0.392 0.368 0.072 0.328 0 1.41 176 0.216 0.688 0.384 0.44 0.088 0.256 0 1.34 185 0.128 0.472 0.328 0.296 0.056 0.184 0 0.78 187 0.2 0.36 0.232 0.216 0.04 0.152 0 0.71 350 0.112 0.432 0.216 0.28 0.048 0.144 0 0.56 50 0.296 0.681 0.296 0.48 0.032 0.192 0 0.95 76 0.144 0.488 0.288 0.312 0.056 0.168 0 0.63 78 0.168 0.408 0.184 0.272 0.032 0.152 0 0.54 143 0.168 0.52 0.232 0.312 0.064 0.24 12 0.88 181 0.184 0.488 0.24 0.28 0.04 0.2 4 0.71 186 0.216 0.512 0.232 0.328 0.056 0.192 0 0.83 222 0.208 0.68 0.288 0.464 0.072 0.2 10 0.83 223 0.272 0.608 0.28 0.408 0.08 0.192 13 0.83 332 0.056 0.456 0.272 0.28 0.056 0.184 0 0.68 175 0.176 0.448 0.28 0.264 0.056 0.192 1 0.68 179 0.144 0.512 0.24 0.328 0.056 0.176 0 0.56 166 0.176 0.456 0.248 0.312 0.056 0.152 0 0.65 326 0.128 0.552 0.32 0.312 0.064 0.232 0 0.87 55 0.224 0.608 0.328 0.408 0.024 0.184 0 1.14

Appendix 2. (continued)

155 0.104 0.416 0.184 0.248 0.032 0.184 0 0.43 351 0.216 0.4 0.248 0.2 0.048 0.184 0 0.63 18 0.168 0.352 0.192 0.192 0.048 0.176 2 0.51 257 0.128 0.448 0.232 0.232 0.048 0.136 1 0.58 258 0.152 0.496 0.184 0.328 0.032 0.16 0 0.44 333 0.104 0.408 0.208 0.256 0.04 0.152 0 0.56 47 0.24 0.624 0.376 0.376 0.056 0.208 0 1.04 354 0.2 0.536 0.224 0.352 0.04 0.176 0 0.97 355 0.2 0.56 0.272 0.368 0.056 0.184 0 0.82 15 0.16 0.568 0.16 0.36 0.048 0.192 0 0.48 65 0.184 0.48 0.288 0.28 0.04 0.16 3 0.56 309 0.208 0.472 0.288 0.312 0.056 0.16 3 0.66 62 0.128 0.4 0.2 0.24 0.048 0.176 0 0.6 67 0.2 0.456 0.288 0.312 0.056 0.144 1 0.65 218 0.072 0.352 0.208 0.208 0.056 0.136 0 0.54 219 0.136 0.568 0.392 0.328 0.072 0.248 0 1.07 259 0.096 0.328 0.168 0.2 0.032 0.12 0 0.46 260 0.096 0.448 0.192 0.272 0.04 0.168 1 0.49 261 0.072 0.44 0.264 0.272 0.048 0.16 0 0.53 262 0.176 0.408 0.208 0.256 0.048 0.128 0 0.53 263 0.152 0.488 0.272 0.288 0.056 0.184 0 0.58 264 0.136 0.504 0.272 0.32 0.064 0.192 1 0.65 265 0.168 0.672 0.312 0.408 0.056 0.232 0 0.97 267 0.152 0.528 0.24 0.352 0.08 0.176 1 0.56 268 0.096 0.432 0.24 0.232 0.048 0.2 0 0.58 269 0.288 0.712 0.304 0.472 0.048 0.232 0 1.07 270 0.088 0.44 0.256 0.224 0.064 0.184 1 0.58 271 0.096 0.352 0.208 0.216 0.048 0.128 0 0.44 272 0.216 0.592 0.28 0.4 0.064 0.2 0 0.7 273 0.16 0.496 0.288 0.32 0.056 0.192 0 0.6 274 0.136 0.432 0.184 0.264 0.04 0.168 7 0.49 275 0.152 0.48 0.208 0.288 0.048 0.192 0 0.6 276 0.08 0.44 0.192 0.24 0.04 0.168 0 0.49 277 0.104 0.368 0.256 0.232 0.048 0.144 0 0.54 278 0.12 0.512 0.232 0.328 0.048 0.184 6 0.49 279 0.064 0.4 0.248 0.216 0.048 0.144 0 0.58 280 0.184 0.44 0.264 0.28 0.048 0.152 0 0.58 281 0.152 0.536 0.256 0.344 0.056 0.272 0 0.68 282 0.208 0.552 0.28 0.312 0.064 0.224 0 1 283 0.208 0.528 0.296 0.272 0.056 0.24 0 1.11 284 0.136 0.608 0.4 0.416 0.088 0.2 1 0.6 286 0.152 0.424 0.256 0.296 0.04 0.208 0 0.6 288 0.24 0.624 0.2 0.408 0.048 0.168 0 0.63

100

Appendix 2. (continued) 290 0.144 0.416 0.24 0.248 0.056 0.16 0 0.56 291 0.12 0.52 0.24 0.328 0.048 0.184 0 0.63 310 0.176 0.512 0.288 0.376 0.072 0.184 1 0.65 360 0.144 0.488 0.264 0.304 0.048 0.184 0 0.87 358 0.128 0.544 0.368 0.32 0.056 0.208 0 1.05 49 0.176 0.384 0.24 0.232 0.04 0.144 0 0.7 61 0.16 0.552 0.336 0.296 0.064 0.256 0 0.94 64 0.104 0.4 0.216 0.208 0.048 0.192 0 0.75 48 0.2 0.488 0.256 0.32 0.04 0.176 0 0.68 52 0.112 0.432 0.216 0.232 0.056 0.16 0 0.73 178 0.128 0.512 0.256 0.328 0.048 0.184 0 0.73 1 0.208 0.568 0.296 0.36 0.056 0.208 0 0.97 4 0.16 0.592 0.336 0.384 0.056 0.216 1 0.82 6 0.12 0.48 0.36 0.24 0.08 0.244 4 1.05 7 0.24 0.56 0.392 0.288 0.072 0.272 0 1.09 8 0.168 0.64 0.408 0.4 0.096 0.248 0 1.22 36 0.216 0.512 0.288 0.296 0.056 0.184 1 0.87 42 0.136 0.568 0.248 0.248 0.048 0.128 1 0.6 51 0.128 0.416 0.216 0.248 0.056 0.144 0 0.66 330 0.112 0.352 0.312 0.336 0.056 0.16 0 0.92 13 0.304 0.44 0.32 0.28 0.056 0.16 0 0.77 165 0.128 0.432 0.232 0.296 0.048 0.152 0 0.6 80 0.168 0.44 0.288 0.272 0.048 0.176 0 0.94 56 0.128 0.512 0.28 0.288 0.048 0.2 0 0.7 357 0.184 0.544 0.336 0.328 0.072 0.216 0 0.66 73 0.232 0.696 0.28 0.424 0.056 0.28 0 1.39 334 0.176 0.44 0.264 0.28 0.048 0.16 0 0.66 389 0.216 0.352 0.408 0.344 0.096 0.224 0 0.77 390 0.168 0.448 0.216 0.28 0.056 0.176 0 0.66 391 0.176 0.448 0.248 0.28 0.064 0.16 0 0.71 335 0.2 0.632 0.296 0.416 0.064 0.224 0 1.02 379 0.112 0.6 0.336 0.408 0.056 0.2 0 1.04 392 0.168 0.672 0.368 0.432 0.056 0.208 0 1.22 29 0.256 0.56 0.264 0.312 0.056 0.208 0 0.71 153 0.168 0.576 0.288 0.368 0.056 0.216 0 0.92 154 0.168 0.504 0.296 0.32 0.064 0.176 0 0.8 341 0.096 0.36 0.16 0.192 0.024 0.152 0 0.49 342 0.096 0.392 0.16 0.232 0.04 0.16 0 0.37 343 0.08 0.44 0.328 0.264 0.064 0.176 0 0.66 344 0.144 0.36 0.248 0.192 0.04 0.152 0 0.56 345 0.128 0.44 0.224 0.248 0.048 0.176 0 0.48 346 0.104 0.392 0.232 0.232 0.056 0.152 0 0.66 347 0.152 0.416 0.248 0.256 0.048 0.168 0 0.68

101

Appendix 2. (continued) 348 0.056 0.32 0.168 0.152 0.032 0.176 0 0.53 349 0.136 0.52 0.288 0.288 0.064 0.264 0 0.77 393 0.184 0.592 0.304 0.392 0.064 0.2 0 1.11 394 0.192 0.6 0.312 0.4 0.064 0.2 0 1.12 395 0.168 0.592 0.424 0.4 0.064 0.192 0 1.07 53 0.136 0.424 0.272 0.264 0.064 0.176 0 0.7 338 0.056 0.376 0.16 0.224 0.032 0.136 0 0.6 381 0.16 0.504 0.256 0.32 0.048 0.16 0 0.85 396 0.128 0.4 0.24 0.256 0.048 0.144 0 0.61 397 0.152 0.408 0.264 0.24 0.048 0.152 0 0.7 398 0.096 0.424 0.288 0.256 0.056 0.176 0 0.73 399 0.056 0.4 0.216 0.208 0.048 0.168 0 0.61 401 0.08 0.424 0.296 0.24 0.048 0.2 0 0.75 363 0.216 0.576 0.344 0.392 0.064 0.192 0 1.19 364 0.192 0.496 0.28 0.32 0.04 0.168 0 1.02 365 0.288 0.728 0.344 0.488 0.048 0.224 0 1.33 101 0.128 0.408 0.192 0.24 0.056 0.176 0 0.43 383 0.168 0.448 0.312 0.216 0.064 0.224 0 0.66 318 0.128 0.424 0.248 0.2 0.048 0.224 0 0.58 151 0.144 0.488 0.208 0.288 0.048 0.2 0 0.61 33 0.128 0.392 0.216 0.208 0.056 0.152 0 0.56 77 0.16 0.504 0.256 0.312 0.056 0.2 0 0.58 88 0.096 0.432 0.28 0.24 0.048 0.184 0 0.58 146 0.096 0.416 0.272 0.232 0.064 0.216 0 0.6 164 0.16 0.384 0.192 0.208 0.04 0.168 0 0.53 388 0.16 0.512 0.264 0.336 0.04 0.184 0 0.6 149 0.136 0.448 0.288 0.448 0.072 0.184 0 0.7 3 0.16 0.44 0.216 0.272 0.048 0.168 2 0.58 84 0.08 0.336 0.152 0.184 0.032 0.152 0 0.32 91 0.176 0.528 0.352 0.304 0.08 0.248 0 0.87 92 0.072 0.352 0.2 0.168 0.048 0.176 0 0.54 152 0.144 0.52 0.28 0.304 0.072 0.216 0 0.71 232 0.12 0.456 0.232 0.28 0.064 0.176 0 0.6 320 0.144 0.448 0.24 0.28 0.048 0.128 0 0.54 321 0.184 0.44 0.224 0.288 0.048 0.144 0 0.53 200 0.152 0.48 0.328 0.264 0.056 0.232 0 0.68 195 0.136 0.432 0.24 0.264 0.064 0.176 0 0.58 201 0.144 0.504 0.32 0.288 0.056 0.2 0 0.65 324 0.168 0.528 0.216 0.296 0.056 0.208 0 0.71 32 0.208 0.664 0.328 0.424 0.072 0.24 1 0.63 94 0.144 0.4 0.208 0.208 0.04 0.16 0 0.53 157 0.144 0.472 0.28 0.264 0.064 0.184 0 0.61 193 0.128 0.44 0.208 0.248 0.056 0.192 0 0.53

102

Appendix 2. (continued) 199 0.16 0.504 0.296 0.296 0.056 0.184 2 0.66 387 0.24 0.512 0.248 0.336 0.032 0.16 1 0.66 95 0.152 0.568 0.376 0.344 0.056 0.248 0 0.7 215 0.2 0.496 0.216 0.296 0.056 0.192 0 0.66 97 0.16 0.464 0.216 0.272 0.04 0.168 0 0.54 190 0.072 0.424 0.184 0.192 0.04 0.168 0 0.48 203 0.208 0.616 0.296 0.376 0.064 0.24 1 0.71 205 0.104 0.456 0.136 0.272 0.024 0.192 0 0.44 207 0.168 0.416 0.224 0.248 0.064 0.168 4 0.61 323 0.216 0.44 0.224 0.256 0.048 0.184 0 0.63 217 0.176 0.44 0.296 0.288 0.056 0.168 0 0.63 98 0.184 0.52 0.336 0.328 0.064 0.184 4 0.68 99 0.152 0.456 0.264 0.256 0.048 0.216 8 0.68 191 0.16 0.496 0.232 0.296 0.04 0.2 2 0.63 54 0.12 0.528 0.28 0.368 0.048 0.168 3 0.61 93 0.168 0.528 0.256 0.328 0.056 0.216 4 0.71 96 0.176 0.48 0.296 0.208 0.056 0.232 0 0.66 100 0.152 0.536 0.24 0.352 0.048 0.192 3 0.68 103 0.12 0.496 0.256 0.288 0.048 0.208 0 0.7 194 0.144 0.448 0.344 0.216 0.056 0.224 0 0.7 196 0.088 0.504 0.216 0.296 0.048 0.2 2 0.61 197 0.104 0.208 0.184 0.208 0.04 0.184 0 0.61 202 0.128 0.44 0.256 0.224 0.064 0.232 0 0.68 204 0.2 0.496 0.264 0.296 0.056 0.184 0 0.68 208 0.152 0.472 0.336 0.248 0.064 0.24 0 0.58 210 0.184 0.472 0.264 0.288 0.056 0.184 0 0.56 212 0.2 0.456 0.304 0.272 0.072 0.208 0 0.63 213 0.088 0.488 0.296 0.28 0.064 0.2 0 0.71 216 0.128 0.48 0.28 0.28 0.048 0.208 0 0.68 307 0.176 0.48 0.248 0.288 0.064 0.184 1 0.56 308 0.072 0.432 0.28 0.232 0.048 0.192 0 0.65 312 0.168 0.44 0.264 0.264 0.056 0.176 1 0.66 313 0.152 0.44 0.24 0.288 0.056 0.152 0 0.7 314 0.152 0.44 0.248 0.28 0.056 0.16 0 0.6 315 0.16 0.376 0.264 0.216 0.056 0.176 0 0.73 316 0.216 0.488 0.256 0.336 0.056 0.176 0 0.61 317 0.152 0.376 0.264 0.232 0.048 0.144 0 0.58 385 0.096 0.44 0.224 0.28 0.048 0.176 0 0.6 228 0.136 0.512 0.28 0.28 0.08 0.232 0 0.63 224 0.168 0.456 0.216 0.272 0.04 0.2 0 0.77 231 0.128 0.472 0.176 0.304 0.04 0.16 0 0.66 188 0.16 0.6 0.304 0.4 0.056 0.176 0 0.95 293 0.144 0.512 0.264 0.344 0.056 0.176 0 0.82

103

Appendix 2. (continued) 294 0.248 0.68 0.328 0.52 0.056 0.192 0 1.02 295 0.2 0.488 0.256 0.304 0.056 0.16 0 0.94 5 0.16 0.68 0.28 0.44 0.056 0.216 2 0.61 233 0.136 0.432 0.192 0.288 0.04 0.136 0 0.46 85 0.144 0.536 0.312 0.312 0.064 0.216 0 0.61 89 0.168 0.576 0.296 0.32 0.072 0.248 0 0.61 31 0.152 0.48 0.28 0.272 0.056 0.184 1 0.61 331 0.2 0.48 0.224 0.312 0.048 0.16 0 0.63 296 0.232 0.584 0.2 0.384 0.04 0.192 1 0.66 298 0.296 0.472 0.208 0.304 0.048 0.176 2 0.61 300 0.176 0.536 0.408 0.296 0.064 0.24 0 0.9 9 0.192 0.488 0.32 0.28 0.056 0.208 0 0.75 230 0.136 0.528 0.304 0.312 0.072 0.216 0 0.77 299 0.184 0.432 0.232 0.256 0.032 0.176 0 0.73 302 0.208 0.568 0.32 0.376 0.056 0.184 0 0.78 297 0.264 0.568 0.328 0.392 0.056 0.2 3 0.75 19 0.152 0.44 0.28 0.232 0.048 0.2 1 0.78 20 0.104 0.432 0.224 0.248 0.048 0.192 0 0.7 23 0.12 0.4 0.264 0.232 0.064 0.152 0 0.53 24 0.168 0.44 0.264 0.296 0.048 0.16 1 0.78 301 0.168 0.544 0.264 0.296 0.056 0.24 0 0.66

104

Appendix 2. (continued) Number Banner Wing Wing Wing Keel Keel Keel Connate Petal Petal Petal Petal Petal Petal Petal Filaments Width Length Lamina Lamina Length Lamina Lamina Length (cm) (cm) Length Width (cm) Length Width (cm) (cm) (cm) (cm) (cm) 167 0.232 0.592 0.312 0.144 0.512 0.192 0.108 0.344 70 0.248 0.704 0.328 0.08 0.576 0.2 0.104 0.528 306 0.12 0.416 0.216 0.052 0.368 0.144 0.06 0.344 424 0.128 0.416 0.224 0.072 0.376 0.136 0.064 0.4 425 0.152 0.504 0.248 0.08 0.416 0.12 0.08 0.4 319 0.136 0.56 0.248 0.068 0.488 0.144 0.076 0.464 69 0.176 0.552 0.232 0.08 0.432 0.152 0.068 0.456 229 0.144 0.568 0.272 0.068 0.48 0.184 0.052 0.448 26 0.208 0.648 0.296 0.076 0.512 0.2 0.1 0.48 41 0.104 0.472 0.256 0.064 0.456 0.136 0.076 0.4 292 0.192 0.624 0.304 0.088 0.528 0.176 0.088 0.496 322 0.176 0.536 0.288 0.076 0.424 0.16 0.068 0.4 303 0.16 0.528 0.24 0.072 0.416 0.16 0.08 0.408 304 0.168 0.632 0.32 0.06 0.504 0.176 0.072 0.464 305 0.192 0.632 0.32 0.08 0.52 0.176 0.08 0.488 380 0.152 0.536 0.248 0.068 0.44 0.144 0.084 0.456 225 0.256 0.656 0.352 0.08 0.576 0.2 0.12 0.472 359 0.144 0.632 0.28 0.06 0.512 0.144 0.084 0.488 226 0.216 0.664 0.344 0.12 0.496 0.176 0.1 0.456 327 0.272 0.776 0.4 0.152 0.624 0.224 0.132 0.592 328 0.152 0.544 0.296 0.1 0.456 0.184 0.092 0.448 369 0.36 0.92 0.472 0.192 0.728 0.248 0.14 0.68 220 0.296 0.936 0.472 0.14 0.76 0.272 0.124 0.68 352 0.288 0.744 0.36 0.136 0.576 0.2 0.12 0.544 353 0.4 1.024 0.528 0.184 0.8 0.264 0.144 0.744 368 0.256 0.904 0.432 0.144 0.76 0.24 0.1 0.664 382 0.336 1.152 0.56 0.16 0.888 0.272 0.156 0.856 325 0.296 0.936 0.44 0.12 0.752 0.264 0.136 0.72 337 0.144 0.56 0.296 0.072 0.4 0.168 0.072 0.408 356 0.2 0.896 0.432 0.096 0.704 0.216 0.096 0.68 75 0.144 0.496 0.232 0.06 0.408 0.16 0.08 0.36 182 0.184 0.504 0.24 0.08 0.432 0.144 0.08 0.392 189 0.16 0.4 0.176 0.064 0.408 0.152 0.092 0.376 12 0.213 0.488 0.192 0.064 0.432 0.16 0.084 0.4 311 0.152 0.536 0.272 0.06 0.512 0.168 0.08 0.464 171 0.312 1.032 0.464 0.132 0.8 0.224 0.12 0.792 339 0.136 0.544 0.208 0.052 0.488 0.136 0.092 0.472 370 0.304 0.888 0.44 0.18 0.752 0.264 0.144 0.72 371 0.224 0.872 0.416 0.116 0.672 0.232 0.132 0.624

105

Appendix 2. (continued) 372 0.272 0.76 0.384 0.168 0.632 0.256 0.132 0.584 373 0.2 0.688 0.36 0.124 0.544 0.2 0.096 0.504 374 0.2 0.784 0.416 0.12 0.6 0.2 0.088 0.544 375 0.152 0.6 0.28 0.08 0.448 0.136 0.056 0.456 376 0.36 1.088 0.488 0.112 0.856 0.232 0.12 0.816 377 0.168 0.592 0.272 0.072 0.472 0.16 0.072 0.424 87 0.36 1.264 0.6 0.156 1.056 0.392 0.14 0.92 38 0.144 0.536 0.272 0.048 0.448 0.168 0.092 0.4 180 0.12 0.528 0.232 0.064 0.448 0.144 0.068 0.424 361 0.392 1.008 0.52 0.184 0.792 0.272 0.144 0.72 362 0.232 1.04 0.44 0.144 0.872 0.264 0.116 0.768 384 0.136 0.488 0.216 0.052 0.4 0.12 0.068 0.384 21 0.16 0.64 0.328 0.08 0.64 0.176 0.1 0.552 22 0.168 0.68 0.336 0.08 0.592 0.176 0.064 0.544 366 0.232 0.792 0.384 0.128 0.64 0.2 0.124 0.632 367 0.216 0.648 0.336 0.112 0.52 0.16 0.092 0.52 386 0.152 0.552 0.28 0.092 0.456 0.16 0.076 0.448 30 0.32 1.2 0.648 0.108 0.96 0.384 0.156 0.856 170 0.248 0.976 0.456 0.14 0.856 0.296 0.12 0.728 336 0.24 0.776 0.392 0.084 0.696 0.248 0.092 0.64 378 0.28 1.424 0.736 0.172 1.008 0.312 0.148 0.976 329 0.264 0.944 0.44 0.152 0.768 0.256 0.132 0.736 400 0.168 0.648 0.272 0.092 0.528 0.152 0.068 0.48 46 0.176 0.656 0.312 0.076 0.512 0.2 0.096 0.496 74 0.304 1.128 0.472 0.144 0.968 0.296 0.164 0.92 176 0.288 1.104 0.448 0.108 1.016 0.312 0.128 0.984 185 0.192 0.616 0.312 0.092 0.552 0.16 0.1 0.52 187 0.144 0.584 0.288 0.076 0.424 0.168 0.088 0.448 350 0.16 0.456 0.232 0.04 0.44 0.144 0.064 0.376 50 0.24 0.832 0.4 0.1 0.648 0.24 0.116 0.608 76 0.128 0.592 0.28 0.064 0.52 0.16 0.048 0.448 78 0.144 0.512 0.192 0.064 0.416 0.152 0.08 0.416 143 0.272 0.784 0.336 0.136 0.616 0.176 0.1 0.6 181 0.248 0.6 0.304 0.112 0.528 0.208 0.08 0.52 186 0.24 0.712 0.352 0.108 0.528 0.168 0.096 0.512 222 0.216 0.752 0.32 0.128 0.64 0.192 0.108 0.624 223 0.32 0.84 0.44 0.108 0.704 0.24 0.132 0.664 332 0.152 0.544 0.28 0.06 0.464 0.16 0.072 0.44 175 0.216 0.608 0.288 0.12 0.512 0.2 0.108 0.472 179 0.152 0.512 0.264 0.08 0.424 0.16 0.076 0.4 166 0.152 0.584 0.256 0.06 0.464 0.144 0.076 0.448 326 0.144 0.704 0.368 0.088 0.552 0.184 0.084 0.496 55 0.288 0.976 0.48 0.18 0.808 0.304 0.144 0.758

106

Appendix 2. (continued) 155 0.128 0.384 0.192 0.068 0.328 0.152 0.06 0.216 351 0.136 0.544 0.248 0.06 0.464 0.16 0.092 0.408 18 0.12 0.448 0.224 0.048 0.368 0.136 0.06 0.36 257 0.16 0.48 0.208 0.068 0.448 0.168 0.092 0.4 258 0.176 0.376 0.208 0.084 0.36 0.16 0.076 0.28 333 0.128 0.456 0.216 0.056 0.392 0.144 0.064 0.36 47 0.24 0.824 0.416 0.14 0.704 0.248 0.124 0.672 354 0.256 0.784 0.4 0.14 0.664 0.224 0.136 0.632 355 0.176 0.712 0.36 0.088 0.56 0.2 0.092 0.52 15 0.16 0.376 0.216 0.076 0.368 0.2 0.06 0.304 65 0.232 0.552 0.248 0.1 0.44 0.128 0.088 0.432 309 0.2 0.536 0.24 0.068 0.44 0.152 0.088 0.456 62 0.176 0.528 0.24 0.076 0.44 0.144 0.08 0.4 67 0.168 0.496 0.224 0.064 0.448 0.152 0.068 0.424 218 0.128 0.4 0.208 0.048 0.36 0.128 0.06 0.36 219 0.264 0.84 0.44 0.112 0.68 0.256 0.132 0.6 259 0.112 0.32 0.176 0.04 0.4 0.136 0.056 0.368 260 0.144 0.432 0.232 0.068 0.304 0.168 0.152 0.32 261 0.136 0.44 0.224 0.044 0.36 0.152 0.044 0.36 262 0.144 0.456 0.224 0.068 0.368 0.136 0.076 0.384 263 0.144 0.48 0.232 0.072 0.392 0.152 0.183 0.408 264 0.152 0.504 0.256 0.076 0.448 0.16 0.084 0.448 265 0.296 0.872 0.44 0.16 0.728 0.24 0.156 0.664 267 0.152 0.52 0.24 0.044 0.456 0.16 0.044 0.408 268 0.176 0.528 0.232 0.072 0.448 0.16 0.092 0.4 269 0.312 0.896 0.44 0.188 0.744 0.24 0.124 0.728 270 0.144 0.504 0.232 0.06 0.44 0.152 0.084 0.392 271 0.112 0.408 0.192 0.052 0.368 0.128 0.068 0.344 272 0.184 0.592 0.296 0.072 0.472 0.168 0.08 0.464 273 0.184 0.488 0.264 0.072 0.432 0.168 0.088 0.424 274 0.136 0.448 0.208 0.06 0.368 0.136 0.08 0.392 275 0.152 0.504 0.256 0.056 0.408 0.16 0.084 0.448 276 0.16 0.456 0.216 0.08 0.392 0.144 0.076 0.344 277 0.12 0.44 0.208 0.076 0.384 0.144 0.076 0.4 278 0.104 0.336 0.128 0.036 0.4 0.128 0.06 0.328 279 0.144 0.432 0.232 0.052 0.392 0.16 0.072 0.432 280 0.104 0.496 0.216 0.06 0.416 0.128 0.076 0.392 281 0.184 0.56 0.288 0.064 0.496 0.168 0.084 0.48 282 0.28 0.888 0.464 0.152 0.72 0.248 0.14 0.664 283 0.264 0.88 0.44 0.108 0.72 0.232 0.116 0.72 284 0.112 0.48 0.256 0.044 0.384 0.16 0.064 0.392 286 0.152 0.504 0.232 0.072 0.392 0.176 0.068 0.424 288 0.16 0.52 0.264 0.06 0.408 0.16 0.088 0.392

107

Appendix 2. (continued)

290 0.176 0.512 0.264 0.072 0.448 0.16 0.088 0.448 291 0.152 0.544 0.264 0.044 0.448 0.16 0.072 0.456 310 0.16 0.52 0.248 0.052 0.44 0.176 0.064 0.432 360 0.256 0.824 0.408 0.144 0.68 0.24 0.14 0.656 358 0.2 0.8 0.384 0.136 0.704 0.24 0.116 0.648 49 0.184 0.608 0.288 0.076 0.488 0.176 0.092 0.48 61 0.192 0.752 0.384 0.08 0.48 0.208 0.1 0.52 64 0.2 0.624 0.288 0.064 0.52 0.208 0.08 0.504 48 0.2 0.592 0.28 0.088 0.496 0.208 0.104 0.48 52 0.216 0.648 0.336 0.12 0.496 0.152 0.092 0.496 178 0.192 0.664 0.32 0.076 0.536 0.184 0.084 0.472 1 0.256 0.896 0.392 0.096 0.688 0.2 0.12 0.664 4 0.176 0.744 0.328 0.08 0.624 0.192 0.104 0.59 6 0.2 0.928 0.464 0.144 0.736 0.264 0.108 0.656 7 0.168 0.968 0.496 0.104 0.72 0.272 0.124 0.672 8 0.176 1.088 0.584 0.16 0.808 0.32 0.156 0.8 36 0.2 0.752 0.328 0.108 0.728 0.248 0.124 0.656 42 0.144 0.496 0.232 0.06 0.408 0.16 0.072 0.384 51 0.128 0.56 0.256 0.06 0.488 0.176 0.068 0.384 330 0.256 0.776 0.384 0.128 0.576 0.2 0.108 0.536 13 0.16 0.64 0.32 0.116 0.52 0.192 0.18 0.48 165 0.152 0.504 0.264 0.08 0.448 0.168 0.076 0.36 80 0.24 0.752 0.4 0.08 0.632 0.208 0.12 0.544 56 0.192 0.624 0.272 0.072 0.496 0.192 0.072 0.456 357 0.2 0.64 0.32 0.092 0.552 0.192 0.092 0.48 73 0.312 1.28 0.616 0.184 0.944 0.336 0.132 0.896 334 0.16 0.592 0.256 0.076 0.504 0.16 0.084 0.448 389 0.184 0.608 0.376 0.08 0.52 0.184 0.092 0.504 390 0.128 0.512 0.248 0.056 0.448 0.16 0.068 0.448 391 0.232 0.632 0.336 0.12 0.512 0.192 0.104 0.44 335 0.216 0.84 0.416 0.112 0.688 0.224 0.1 0.632 379 0.272 0.856 0.4 0.128 0.688 0.224 0.124 0.656 392 0.296 0.92 0.496 0.152 0.752 0.28 0.12 0.68 29 0.256 0.528 0.344 0.108 0.472 0.288 0.152 0.392 153 0.304 0.744 0.416 0.168 0.6 0.24 0.14 0.56 154 0.216 0.656 0.328 0.108 0.544 0.176 0.1 0.48 341 0.144 0.472 0.184 0.02 0.408 0.12 0.08 0.376 342 0.128 0.344 0.24 0.072 0.296 0.16 0.076 0.304 343 0.168 0.568 0.28 0.088 0.48 0.16 0.084 0.44 344 0.136 0.448 0.216 0.06 0.416 0.16 0.064 0.384 345 0.112 0.488 0.184 0.064 0.448 0.136 0.06 0.424 346 0.128 0.488 0.16 0.044 0.488 0.152 0.064 0.448 347 0.136 0.616 0.272 0.068 0.504 0.16 0.088 0.48

108

Appendix 2. (continued)

348 0.176 0.408 0.256 0.08 0.368 0.176 0.076 0.328 349 0.2 0.64 0.32 0.076 0.504 0.168 0.092 0.464 393 0.264 0.944 0.496 0.156 0.712 0.232 0.136 0.68 394 0.304 0.944 0.464 0.136 0.736 0.248 0.112 0.696 395 0.288 0.912 0.44 0.136 0.72 0.248 0.144 0.76 53 0.136 0.576 0.256 0.064 0.48 0.16 0.072 0.456 338 0.128 0.52 0.216 0.06 0.472 0.128 0.072 0.416 381 0.2 0.648 0.32 0.076 0.584 0.2 0.1 0.512 396 0.168 0.52 0.248 0.088 0.456 0.16 0.092 0.432 397 0.168 0.568 0.272 0.076 0.472 0.16 0.072 0.424 398 0.168 0.624 0.288 0.072 0.52 0.176 0.076 0.52 399 0.176 0.528 0.248 0.064 0.432 0.136 0.08 0.4 401 0.16 0.632 0.32 0.116 0.496 0.168 0.084 0.456 363 0.272 1.008 0.448 0.136 0.808 0.256 0.14 0.792 364 0.248 0.88 0.392 0.12 0.728 0.272 0.124 0.688 365 0.328 1.16 0.52 0.192 0.936 0.28 0.148 0.872 101 0.168 0.432 0.208 0.076 0.376 0.152 0.088 0.36 383 0.208 0.624 0.24 0.068 0.56 0.176 0.076 0.536 318 0.168 0.552 0.24 0.076 0.544 0.168 0.084 0.504 151 0.16 0.536 0.248 0.08 0.44 0.152 0.068 0.4 33 0.144 0.48 0.208 0.048 0.384 0.128 0.056 0.344 77 0.16 0.488 0.224 0.048 0.432 0.128 0.068 0.392 88 0.176 0.496 0.208 0.084 0.416 0.136 0.06 0.384 146 0.136 0.504 0.224 0.072 0.408 0.136 0.064 0.368 164 0.16 0.448 0.208 0.072 0.392 0.136 0.08 0.352 388 0.184 0.488 0.216 0.072 0.416 0.144 0.072 0.4 149 0.16 0.56 0.232 0.08 0.472 0.144 0.08 0.448 3 0.16 0.56 0.248 0.08 0.488 0.16 0.08 0.44 84 0.064 0.48 0.208 0.083 0.392 0.128 0.068 0.408 91 0.192 0.688 0.36 0.088 0.6 0.216 0.108 0.592 92 0.128 0.488 0.232 0.056 0.416 0.136 0.084 0.416 152 0.192 0.664 0.264 0.084 0.608 0.192 0.1 0.568 232 0.12 0.488 0.2 0.056 0.432 0.136 0.06 0.408 320 0.152 0.544 0.248 0.076 0.504 0.192 0.084 0.416 321 0.176 0.48 0.256 0.092 0.384 0.192 0.092 0.424 200 0.176 0.576 0.256 0.06 0.52 0.168 0.068 0.52 195 0.168 0.488 0.232 0.068 0.4 0.144 0.072 0.424 201 0.208 0.608 0.248 0.072 0.544 0.192 0.096 0.504 324 0.168 0.528 0.264 0.048 0.488 0.16 0.048 0.48 32 0.136 0.632 0.232 0.068 0.488 0.168 0.088 0.488 94 0.152 0.496 0.248 0.08 0.44 0.144 0.08 0.36 157 0.16 0.552 0.232 0.064 0.512 0.16 0.08 0.456 193 0.136 0.488 0.24 0.08 0.408 0.16 0.084 0.392

109

Appendix 2. (continued) 199 0.144 0.592 0.272 0.048 0.504 0.168 0.08 0.432 387 0.184 0.576 0.28 0.08 0.52 0.16 0.092 0.448 95 0.232 0.64 0.288 0.064 0.544 0.176 0.1 0.52 215 0.184 0.592 0.248 0.088 0.52 0.168 0.092 0.472 97 0.152 0.408 0.232 0.092 0.392 0.16 0.084 0.424 190 0.136 0.432 0.192 0.072 0.392 0.12 0.08 0.36 203 0.208 0.648 0.256 0.084 0.6 0.208 0.092 0.536 205 0.112 0.312 0.152 0.06 0.36 0.136 0.08 0.328 207 0.168 0.544 0.24 0.092 0.464 0.152 0.092 0.448 323 0.144 0.496 0.224 0.056 0.432 0.128 0.056 0.408 217 0.144 0.496 0.24 0.048 0.408 0.096 0.052 0.408 98 0.184 0.624 0.304 0.072 0.496 0.184 0.096 0.456 99 0.16 0.592 0.264 0.068 0.48 0.136 0.084 0.44 191 0.16 0.568 0.264 0.08 0.48 0.168 0.084 0.448 54 0.16 0.528 0.256 0.076 0.488 0.168 0.096 0.488 93 0.2 0.608 0.296 0.06 0.504 0.176 0.096 0.488 96 0.128 0.552 0.208 0.064 0.52 0.16 0.076 0.528 100 0.208 0.632 0.296 0.08 0.52 0.176 0.092 0.48 103 0.16 0.592 0.28 0.068 0.592 0.128 0.044 0.448 194 0.2 0.648 0.28 0.084 0.56 0.168 0.096 0.496 196 0.168 0.576 0.28 0.088 0.496 0.176 0.1 0.464 197 0.16 0.528 0.24 0.072 0.44 0.152 0.088 0.416 202 0.16 0.6 0.248 0.072 0.552 0.216 0.084 0.52 204 0.16 0.552 0.28 0.064 0.504 0.2 0.072 0.472 208 0.176 0.584 0.24 0.1 0.528 0.16 0.092 0.48 210 0.136 0.52 0.24 0.056 0.4 0.12 0.064 0.408 212 0.152 0.528 0.248 0.076 0.456 0.168 0.084 0.44 213 0.16 0.6 0.28 0.044 0.512 0.168 0.064 0.464 216 0.168 0.592 0.272 0.056 0.528 0.168 0.08 0.528 307 0.12 0.448 0.216 0.06 0.408 0.128 0.072 0.416 308 0.152 0.552 0.184 0.072 0.56 0.168 0.084 0.52 312 0.136 0.56 0.248 0.064 0.464 0.176 0.072 0.448 313 0.168 0.616 0.272 0.08 0.528 0.176 0.092 0.472 314 0.144 0.52 0.232 0.06 0.392 0.152 0.08 0.408 315 0.216 0.592 0.264 0.084 0.464 0.16 0.084 0.456 316 0.152 0.472 0.232 0.064 0.432 0.136 0.06 0.424 317 0.144 0.52 0.24 0.048 0.4 0.12 0.064 0.408 385 0.176 0.552 0.24 0.088 0.472 0.136 0.084 0.424 228 0.192 0.576 0.224 0.072 0.52 0.152 0.096 0.488 224 0.16 0.592 0.304 0.068 0.488 0.16 0.088 0.456 231 0.144 0.576 0.248 0.06 0.4 0.136 0.072 0.448 188 0.24 0.856 0.456 0.16 0.624 0.2 0.112 0.544 293 0.192 0.648 0.328 0.092 0.528 0.192 0.092 0.504

110

Appendix 2. (continued)

294 0.312 0.888 0.44 0.116 0.72 0.248 0.128 0.68 295 0.32 0.864 0.464 0.14 0.632 0.24 0.12 0.608 5 0.176 0.6 0.288 0.092 0.472 0.16 0.1 0.432 233 0.12 0.56 0.264 0.048 0.472 0.128 0.076 0.424 85 0.2 0.576 0.28 0.068 0.488 0.176 0.096 0.44 89 0.168 0.592 0.256 0.088 0.552 0.168 0.092 0.512 31 0.16 0.616 0.28 0.064 0.528 0.168 0.088 0.48 331 0.192 0.528 0.256 0.092 0.44 0.152 0.088 0.384 296 0.184 0.56 0.264 0.08 0.456 0.136 0.056 0.424 298 0.16 0.512 0.24 0.088 0.496 0.152 0.076 0.408 300 0.16 0.6 0.24 0.096 0.592 0.128 0.088 0.512 9 0.12 0.52 0.272 0.072 0.48 0.16 0.096 0.512 230 0.208 0.752 0.376 0.136 0.584 0.2 0.12 0.536 299 0.192 0.592 0.296 0.1 0.48 0.168 0.088 0.464 302 0.192 0.648 0.328 0.08 0.488 0.152 0.064 0.472 297 0.16 0.608 0.28 0.076 0.496 0.16 0.072 0.448 19 0.16 0.632 0.312 0.068 0.488 0.192 0.08 0.456 20 0.16 0.48 0.192 0.068 0.44 0.136 0.076 0.416 23 0.112 0.448 0.192 0.056 0.44 0.128 0.068 0.432 24 0.144 0.584 0.312 0.083 0.52 0.176 0.088 0.48 301 0.184 0.6 0.272 0.116 0.472 0.176 0.1 0.44

111

Appendix 2. (continued)

Number Style Style Ovary Ovary Ovule Ovule Ovule Length Diameter Length Width Length Width Number (cm) (cm) (cm) (cm) (cm) (cm) 167 0.272 0.016 0.176 0.048 0.04 0.028 2 70 0.272 0.018 0.2 0.052 0.04 0.032 2 306 0.168 0.006 0.156 0.04 0.036 0.024 2 424 0.184 0.006 0.204 0.14 0.04 0.028 2 425 0.176 0.008 0.172 0.04 0.048 0.032 2 319 0.216 0.016 0.192 0.044 0.04 0.028 2 69 0.224 0.012 0.224 0.08 0.72 0.068 2 229 0.248 0.006 0.192 0.04 0.044 0.032 2 26 0.256 0.004 0.232 0.048 0.044 0.036 2 41 0.168 0.006 0.192 0.064 0.06 0.044 2 292 0.28 0.016 0.196 0.044 0.044 0.032 2 322 0.248 0.018 0.16 0.048 0.04 0.028 2 303 0.208 0.01 0.192 0.064 0.056 0.048 2 304 0.248 0.012 0.208 0.06 0.052 0.036 2 305 0.224 0.012 0.244 0.052 0.064 0.044 2 380 0.24 0.006 0.2 0.04 0.028 0.024 2 225 0.256 0.018 0.212 0.068 0.06 0.044 2 359 0.2 0.01 0.244 0.08 0.088 0.072 2 226 0.296 0.01 0.204 0.064 0.064 0.056 2 327 0.368 0.022 0.236 0.048 0.048 0.032 2 328 0.296 0.014 0.136 0.032 0.036 0.016 2 369 0.44 0.012 0.268 0.096 0.096 0.08 2 220 0.408 0.02 0.244 0.06 0.048 0.036 2 352 0.28 0.01 0.236 0.08 0.076 0.06 2 353 0.48 0.02 0.24 0.056 0.048 0.036 2 368 0.432 0.012 0.276 0.076 0.068 0.052 2 382 0.552 0.016 0.284 0.1 0.092 0.064 2 325 0.456 0.012 0.24 0.076 0.064 0.056 2 337 0.192 0.012 0.216 0.056 0.06 0.044 2 356 0.36 0.01 0.284 0.056 0.048 0.044 2 75 0.184 0.006 0.156 0.048 0.044 0.02 2 182 0.184 0.008 0.18 0.048 0.044 0.032 2 189 0.168 0.008 0.164 0.04 0.036 0.028 2 12 0.2 0.008 NA NA NA NA NA 311 0.264 0.006 0.204 0.076 0.088 0.068 2 171 0.472 0.008 0.284 0.076 0.076 0.056 2 339 0.2 0.006 0.18 0.064 0.04 0.044 2 370 0.448 0.014 0.224 0.048 0.044 0.028 2 371 0.44 0.014 0.204 0.04 0.04 0.024 2 372 0.32 0.016 0.228 0.068 0.076 0.048 2

112

Appendix 2. (continued) 373 0.312 0.014 0.176 0.06 0.044 0.032 2 374 0.296 0.008 0.244 0.068 0.068 0.04 2 375 0.264 0.008 0.204 0.072 0.076 0.06 2 376 0.512 0.018 0.3 0.072 0.072 0.056 2 377 0.216 0.01 0.228 0.076 0.116 0.064 2 87 0.608 0.024 0.328 0.056 0.048 0.024 2 38 0.208 0.008 0.176 0.052 0.048 0.044 2 180 0.256 0.008 0.172 0.032 0.036 0.028 2 361 0.464 0.014 0.256 0.056 0.032 0.032 2 362 0.496 0.028 0.288 0.06 0.048 0.036 2 384 0.216 0.008 0.18 0.06 0.052 0.04 2 21 0.328 0.006 0.292 0.08 0.04 0.04 2 22 0.32 0.004 0.24 0.072 0.06 0.044 2 366 0.424 0.018 0.208 0.04 0.052 0.032 2 367 0.296 0.01 0.228 0.084 0.092 0.06 2 386 0.216 0.006 0.208 0.052 0.048 0.04 2 30 0.504 0.008 0.328 0.076 0.048 0.052 2 170 0.456 0.014 0.288 0.084 0.06 0.06 2 336 0.368 0.012 0.252 0.084 0.068 0.056 2 378 0.616 0.018 0.376 0.088 0.068 0.06 2 329 0.472 0.018 0.24 0.068 0.048 0.04 2 400 0.24 0.008 0.228 0.056 0.044 0.04 2 46 0.248 0.008 0.28 0.06 0.048 0.048 2 74 0.552 0.01 0.368 0.084 0.068 0.064 2 176 0.52 0.01 0.4 0.1 0.092 0.08 2 185 0.24 0.008 0.256 0.084 0.064 0.06 2 187 0.248 0.01 0.16 0.044 0.012 0.008 2 350 0.2 0.006 0.164 0.056 0.056 0.052 2 50 0.36 0.014 0.26 0.056 0.072 0.044 2 76 0.224 0.006 0.228 0.072 0.052 0.044 2 78 0.208 0.008 0.188 0.044 0.044 0.028 2 143 0.36 0.006 0.228 0.068 0.048 0.04 2 181 0.248 0.01 0.26 0.064 0.052 0.04 2 186 0.304 0.008 0.22 0.04 0.044 0.024 2 222 0.432 0.012 0.248 0.06 0.056 0.04 2 223 0.464 0.018 0.236 0.064 0.052 0.044 2 332 0.208 0.01 0.208 0.088 0.072 0.068 2 175 0.208 0.008 0.2 0.06 0.044 0.04 2 179 0.16 0.006 0.24 0.1 0.092 0.068 2 166 0.24 0.006 0.18 0.056 0.052 0.044 2 326 0.28 0.01 0.224 0.076 0.064 0.052 2 55 0.44 0.016 0.232 0.064 0.052 0.044 2 155 0.192 0.01 0.116 0.04 0.032 0.024 2

113

Appendix 2. (continued) 351 0.216 0.016 0.164 0.048 0.048 0.04 2 18 0.192 0.006 0.168 0.068 0.052 0.048 2 257 0.232 0.012 0.18 0.044 0.04 0.028 2 258 0.112 0.008 0.1 0.028 0.032 0.02 2 333 0.2 0.006 0.16 0.04 0.036 0.028 2 47 0.408 0.018 0.264 0.084 0.096 0.048 2 354 0.456 0.008 0.192 0.028 0.04 0.02 2 355 0.248 0.01 0.22 0.064 0.052 0.048 2 15 0.24 0.008 0.148 0.04 0.036 0.02 2 65 0.176 0.008 0.244 0.088 0.076 0.068 2 309 0.248 0.008 0.216 0.068 0.052 0.04 2 62 0.208 0.008 0.184 0.056 0.044 0.044 2 67 0.184 0.008 0.232 0.1 0.084 0.036 2 218 0.192 0.008 0.176 0.06 0.048 0.044 2 219 0.352 0.014 0.244 0.1 0.08 0.064 2 259 0.176 0.012 0.112 0.032 0.036 0.024 2 260 0.144 0.004 0.16 0.044 0.052 0.04 2 261 0.2 0.01 0.132 0.052 0.06 0.048 2 262 0.176 0.008 0.156 0.04 0.036 0.02 2 263 0.28 0.006 0.168 0.036 0.048 0.032 2 264 0.248 0.006 0.176 0.044 0.056 0.036 2 265 0.424 0.012 0.236 0.052 0.044 0.032 2 267 0.232 0.01 0.172 0.08 0.064 0.056 2 268 0.216 0.006 0.184 0.06 0.06 0.048 2 269 0.464 0.016 0.3 0.056 0.048 0.036 2 270 0.208 0.008 0.184 0.068 0.072 0.048 2 271 0.184 0.006 0.144 0.068 0.064 0.044 2 272 0.24 0.008 0.22 0.056 0.056 0.048 2 273 0.192 0.01 0.212 0.096 0.084 0.076 2 274 0.208 0.008 0.152 0.04 0.036 0.028 2 275 0.2 0.01 0.224 0.068 0.076 0.056 2 276 0.216 0.012 0.156 0.048 0.04 0.028 2 277 0.2 0.006 0.168 0.084 0.072 0.068 2 278 0.2 0.01 0.188 0.076 0.06 0.04 2 279 0.24 0.01 0.184 0.072 0.056 0.052 2 280 0.224 0.01 0.2 0.088 0.1 0.08 2 281 0.24 0.012 0.224 0.06 0.06 0.044 2 282 0.408 0.02 0.2 0.052 0.044 0.036 2 283 0.4 0.012 0.268 0.068 0.06 0.044 2 284 0.192 0.01 0.208 0.08 0.06 0.06 2 286 0.232 0.008 0.188 0.064 0.06 0.052 2 288 0.208 0.01 0.2 0.08 0.096 0.076 2 290 0.24 0.012 0.196 0.032 0.04 0.024 2

114

Appendix 2. (continued) 291 0.24 0.012 0.192 0.044 0.056 0.04 2 310 0.248 0.008 0.176 0.076 0.076 0.064 2 360 0.368 0.02 0.248 0.052 0.044 0.032 2 358 0.4 0.01 0.276 0.084 0.084 0.076 2 49 0.216 0.008 0.244 0.056 0.052 0.044 2 61 0.288 0.006 0.264 0.08 0.084 0.072 2 64 0.216 0.006 0.264 0.052 0.052 0.04 2 48 0.248 0.012 0.18 0.056 0.04 0.028 2 52 0.264 0.016 0.224 0.052 0.044 0.028 2 178 0.264 0.006 0.228 0.072 0.064 0.052 2 1 0.416 0.008 0.264 0.068 0.056 0.04 2 4 0.344 0.012 0.22 0.076 0.064 0.048 2 6 0.392 0.018 0.24 0.048 0.056 0.04 2 7 0.6 0.008 NA NA NA NA NA 8 0.4 0.014 0.34 0.064 0.04 0.032 2 36 0.36 0.006 0.244 0.048 0.04 0.024 2 42 0.184 0.008 0.212 0.06 0.06 0.04 2 51 0.232 0.01 0.208 0.068 0.056 0.052 2 330 0.328 0.012 0.196 0.04 0.044 0.032 2 13 0.24 0.006 0.24 0.044 0.048 0.028 2 165 0.176 0.006 0.176 0.04 0.04 0.032 2 80 0.36 0.014 0.2 0.068 0.04 0.028 2 56 0.272 0.008 0.188 0.064 0.056 0.04 2 357 0.216 0.008 0.212 0.096 0.076 0.072 2 73 0.536 0.014 0.408 0.068 0.064 0.048 2 334 0.2 0.006 0.196 0.08 0.06 0.04 2 389 0.224 0.008 0.284 0.12 0.14 0.088 2 390 0.216 0.008 0.2 0.056 0.044 0.044 2 391 0.24 0.014 0.2 0.068 0.06 0.044 2 335 0.432 0.014 0.22 0.056 0.056 0.04 2 379 0.392 0.014 0.272 0.084 0.052 0.052 2 392 0.408 0.01 0.296 0.044 0.028 0.028 1 29 0.28 0.008 0.148 0.048 0.036 0.028 2 153 0.424 0.018 0.192 0.064 0.044 0.036 2 154 0.288 0.012 0.196 0.064 0.076 0.056 2 341 0.2 0.012 0.144 0.032 0.032 0.024 2 342 0.176 0.012 0.152 0.04 0.032 0.028 2 343 0.208 0.016 0.236 0.088 0.072 0.056 2 344 0.216 0.006 0.168 0.052 0.044 0.036 2 345 0.264 0.008 0.172 0.044 0.056 0.04 2 346 0.248 0.012 0.22 0.088 0.08 0.072 2 347 0.28 0.012 0.184 0.048 0.032 0.028 2 348 0.176 0.012 0.172 0.036 0.036 0.024 2

115

Appendix 2. (continued)

349 0.24 0.012 0.196 0.052 0.052 0.04 2 393 0.408 0.02 0.256 0.048 0.04 0.032 2 394 0.496 0.008 0.208 0.044 0.056 0.032 2 395 0.448 0.012 0.3 0.104 0.092 0.08 2 53 0.224 0.008 0.252 0.088 0.076 0.064 2 338 0.224 0.01 0.168 0.052 0.044 0.032 2 381 0.28 0.008 0.216 0.044 0.044 0.032 2 396 0.232 0.008 0.176 0.06 0.052 0.036 2 397 0.224 0.008 0.18 0.064 0.048 0.048 2 398 0.2 0.006 NA 0.068 0.044 0.04 2 399 0.2 0.016 0.204 0.04 0.036 0.036 2 401 0.256 0.01 0.18 0.072 0.06 0.06 2 363 0.48 0.014 0.292 0.048 0.048 0.036 2 364 0.424 0.008 0.236 0.048 0.056 0.032 2 365 0.536 0.02 0.312 0.056 0.048 0.032 2 101 0.216 0.02 0.112 0.028 0.032 0.024 2 383 0.28 0.01 0.216 0.064 0.056 0.048 2 318 0.28 0.01 0.216 0.06 0.076 0.052 2 151 0.232 0.016 0.188 0.06 0.068 0.052 2 33 0.184 0.008 0.148 0.044 0.04 0.04 2 77 0.208 0.01 0.18 0.064 0.056 0.048 2 88 0.216 0.006 0.172 0.06 0.06 0.048 2 146 0.216 0.008 0.176 0.08 0.076 0.068 2 164 0.2 0.016 0.148 0.04 0.044 0.028 2 388 0.224 0.01 0.18 0.06 0.048 0.04 2 149 0.256 0.006 0.184 0.052 0.048 0.04 2 3 0.224 0.014 0.156 0.072 0.084 0.048 2 84 0.192 0.006 0.2 0.08 0.064 0.048 2 91 0.296 0.008 0.248 0.096 0.068 0.064 2 92 0.2 0.008 0.148 0.044 0.044 0.036 2 152 0.312 0.01 0.22 0.064 0.048 0.04 2 232 0.208 0.01 0.18 0.072 0.072 0.068 2 320 0.272 0.01 0.172 0.044 0.048 0.032 2 321 0.264 0.012 0.156 0.048 0.04 0.032 2 200 0.304 0.01 0.192 0.092 0.072 0.068 2 195 0.216 0.008 0.2 0.084 0.08 0.064 2 201 0.312 0.006 0.176 0.08 0.056 0.052 2 324 0.264 0.006 0.22 0.064 0.06 0.044 2 32 0.288 0.006 0.2 0.064 0.06 0.048 2 94 0.2 0.012 0.156 0.044 0.04 0.032 2 157 0.264 0.01 0.196 0.056 0.06 0.04 2 193 0.24 0.01 0.164 0.036 0.036 0.024 2 199 0.272 0.01 0.192 0.068 0.06 0.06 2

116

Appendix 2. (continued)

387 0.256 0.012 0.184 0.044 0.044 0.028 2 95 0.328 0.01 0.18 0.068 0.04 0.036 2 215 0.296 0.006 0.188 0.048 0.044 0.028 2 97 0.216 0.02 0.164 0.044 0.028 0.024 2 190 0.184 0.01 0.144 0.04 0.04 0.028 2 203 0.304 0.012 0.184 0.068 0.048 0.036 2 205 0.2 0.006 0.112 0.02 0.032 0.024 2 207 0.256 0.006 0.172 0.048 0.048 0.036 2 323 0.216 0.014 0.176 0.06 0.056 0.048 2 217 0.224 0.006 0.184 0.06 0.048 0.048 2 98 0.256 0.014 0.18 0.052 0.04 0.032 2 99 0.272 0.012 0.18 0.052 0.04 0.031 2 191 0.248 0.006 0.168 0.036 0.04 0.02 2 54 0.216 0.008 0.24 0.076 0.084 0.064 2 93 0.248 0.012 0.224 0.052 0.04 0.036 2 96 0.272 0.008 0.224 0.076 0.076 0.052 2 100 0.256 0.008 0.212 0.06 0.056 0.028 2 103 0.232 0.008 0.232 0.052 0.068 0.036 2 194 0.296 0.01 0.196 0.056 0.044 0.04 2 196 0.232 0.006 0.196 0.048 0.032 0.032 2 197 0.24 0.012 0.164 0.052 0.048 0.032 2 202 0.264 0.012 0.228 0.088 0.092 0.06 2 204 0.224 0.006 0.204 0.088 0.068 0.06 2 208 0.264 0.01 0.204 0.096 0.08 0.076 2 210 0.208 0.006 0.224 0.108 0.088 0.088 2 212 0.216 0.008 0.224 0.104 0.08 0.08 2 213 0.264 0.008 0.212 0.088 0.072 0.068 2 216 0.288 0.008 0.224 0.096 0.1 0.08 2 307 0.224 0.008 0.188 0.064 0.072 0.068 2 308 0.28 0.006 0.196 0.044 0.056 0.04 2 312 0.224 0.008 0.212 0.068 0.08 0.064 2 313 0.28 0.006 0.188 0.048 0.06 0.024 2 314 0.224 0.01 0.184 0.056 0.052 0.048 2 315 0.24 0.014 0.208 0.064 0.048 0.036 2 316 0.232 0.008 0.184 0.064 0.064 0.048 2 317 0.224 0.01 0.196 0.068 0.06 0.048 2 385 0.224 0.012 0.18 0.06 0.056 0.048 2 228 0.272 0.012 0.208 0.06 0.052 0.036 2 224 0.24 0.006 0.196 0.048 0.072 0.048 2 231 0.272 0.006 0.188 0.06 0.068 0.056 2 188 0.384 0.012 0.24 0.06 0.052 0.036 2 293 0.296 0.008 0.208 0.068 0.052 0.044 2 294 0.384 0.016 0.264 0.064 0.052 0.048 2

117

Appendix 2. (continued) 295 0.368 0.018 0.208 0.052 0.036 0.028 2 5 0.256 0.012 0.168 0.032 0.04 0.028 2 233 0.264 0.008 0.176 0.06 0.06 0.04 2 85 0.248 0.014 0.22 0.056 0.044 0.032 2 89 0.288 0.02 0.18 0.036 0.04 0.02 2 31 0.272 0.008 0.192 0.036 0.044 0.032 2 331 0.248 0.01 0.16 0.036 0.036 0.024 2 296 0.168 0.016 0.172 0.044 0.04 0.028 2 298 0.248 0.014 0.16 0.04 0.036 0.028 2 300 0.288 0.012 0.24 0.1 0.076 0.072 2 9 0.208 0.006 NA NA NA NA NA 230 0.304 0.01 0.248 0.048 0.04 0.024 2 299 0.248 0.01 0.208 0.048 0.044 0.036 2 302 0.248 0.012 0.224 0.072 0.088 0.06 2 297 0.24 0.006 NA 0.048 NA 0.032 2 19 0.248 0.008 0.228 0.06 0.064 0.044 2 20 0.232 0.006 0.208 0.044 0.04 0.032 2 23 0.192 0.004 0.208 0.056 0.06 0.02 2 24 0.232 0.004 0.232 0.056 0.064 0.044 2 301 0.216 0.008 0.2 0.056 0.048 0.04 2

118

Appendix 3. R Code used for selected statistical analyses and the production of distribution maps.

#Multivariate Data

Data = read.csv("C:\\Users\\Benjamin\\Desktop\\Master's Thesis\\Data\\Measurements March 21\\Variegatumonly.csv")

#PCA

#2D PCA rownames(Data)<-Data[,1]

PC <- princomp(Data[,c(9,12:23,25,27:33,35,36,38:41)], cor=TRUE,center=TRUE, scale=TRUE,) lambda <- PC$sdev*sqrt(nrow(PC$scores))

#plot for all 5 plot (t(t(PC$scores)/lambda),pch=as.numeric(Data[,5])

,col=as.numeric(Data[,5])) legend("bottomleft",c("'T. geminiflorum'","'T. subsalinum'","T. appendiculatum","T. polyodon","T. variegatum"),

pch = c(1,2,3,4,5),col=c("black","red","green","blue","cyan"), horiz = FALSE, cex = 1)

#Plot for all 3 plot (t(t(PC$scores)/lambda),pch=as.numeric(Data[,4])

,col=as.numeric(Data[,4])) legend("bottomleft",c("T. appendiculatum","T. polyodon","T. variegatum"),

pch = c(1,2,3),col=c("black","red","green"), horiz = FALSE, cex = 1)

#PC1 vs PC5 plot(PC$scores[,2],PC$scores[,3],pch=as.numeric(Data[,5])

,col=as.numeric(Data[,5]),xlab="Comp.1"

,ylab="Comp.5") legend("topleft",c("'T. geminiflorum'","'T. subsalinum'","T. appendiculatum","T. polyodon","T. variegatum"),

pch = c(1,2,3,4,5),col=c("black","red","green","blue","cyan"), horiz = FALSE, cex = 1)

#3D PCA

119

Appendix 3. (continued) library(scatterplot3d) with(mtcars, {

scatterplot3d(PC$scores[,1], PC$scores[,2], PC$scores[,2], # x y and z axis

color=as.numeric(Data[,4]), pch=as.numeric(Data[,4]), # filled blue circles

type="h", # lines to the horizontal plane

main="",

xlab="Comp. 1",

ylab="Comp. 2",

zlab="Comp. 3")

}) legend("topleft",c("T. appendiculatum","T. polyodon","T. variegatum"),

pch = c(1,2,3),col=c("black","red","green"), horiz = FALSE, cex = 1)

#Beanplots library(beanplot)

#GEMINIFLORUM beanplot(Banner.Petal.Length~March.Vincent.ID, data = Data,

xlab = "", ylab = "cm",

main = "Banner Petal Length", cex.axis = 1,