Abstract a Morphological Analysis of the Trifolium

ABSTRACT

A MORPHOLOGICAL ANALYSIS OF THE TRIFOLIUM AMABILE KUNTH SPECIES COMPLEX IN NORTH AMERICA

by Tia Ahlquist

The Trifolium amabile Kunth species complex (Fabaceae) is a group of New World of clovers ranging from the mountains of southern Arizona of the United Sates, throughout the mountainous regions of Central America, and continuing southward to northern-central Argentina. As currently circumscribed in Zohary and Heller’s monograph (1984), this group is comprised of a singular species, T. amabile, and five varieties. Specimens from herbaria worldwide have been utilized for a morphometric examination to determine specific boundaries for the North American members of this complex. In this paper nine species, including four new species, are recognized as distinct and described. These species include: Trifolium amabile, T. blanquitum sp. nov., T. cognatum, T. goniocarpum, T. hickeyi sp. nov., T. laciae sp. nov., T. lozani, T. mexicanum, and T. sonoranensis sp. nov. A MORPHOLOGICAL ANALYSIS OF THE TRIFOLIUM AMABILE KUNTH SPECIES COMPLEX IN NORTH AMERICA

A Thesis

Submitted to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Science Department of Botany by Tia Kay Ahlquist Miami University Oxford, Ohio 2012

Advisor ______Michael A. Vincent Reader______R. James Hickey Reader______Richard C. Moore

© Tia K. Ahlquist 2012 Table of Contents Introduction……………………………………………………………………………………1 Materials and Methods………………………………………………………………………6 Results…………………………………………………………………………………………14 Discussion…………………………………………………………………………………….47 Taxonomic Treatment………………………………………………………………………48 Literature Cited………………………………………………………………………………99 Appendix 1………………………………………………………………………………….101 Appendix 2………………………………………………………………………………….104 Appendix 3………………………………………………………………………………….110 Appendix 4………………………………………………………………………………….128 Appendix 5………………………………………………………………………………….141 Appendix 6………………………………………………………………………………….153

iii List of Tables 1. List of published names……………………………………………………………..5 2. List of herbaria………………………………………………………………………..8 3. P values resulting from a Student’s t-Test for T. amabile………………….26 4. P values resulting from a Student’s t-Test for T. blanquitum……………..28 5. P values resulting from a Student’s t-Test for T. cognatum………………..30 6. P values resulting from a Student’s t-Test for T. goniocarpum……………32 7. P values resulting from a Student’s t-Test for T. hickeyi………………..…34 8. P values resulting from a Student’s t-Test for T. laciae…………………….36 9. P values resulting from a Student’s t-Test for T. lozani………….…………38 10. P values resulting from a Student’s t-Test for T. mexicanum……………..40 11. P values resulting from a Student’s t-Test for T. sonoranensis…………..42

iv List of Figures 1. Illustration of Trifolium amabile………………………………………………..….4 2. Diagram of selected vegetative characters………………………………………9 3. Diagram of selected inflorescence characters…………………………………10 4. Diagram of selected floral characters…………………………………………..11 5. Diagram of selected fruit characters……………………………………………12 6. Diagram of pubescence characters……………………………………………..13 7. Preliminary results from NTSYSpc analyses………………………………….16 8. Results from a PCA and SAHN clustering analysis of the largest group….. …………………………………………………………………………………………..17 9. Box and whisker plot of peduncle length………………………………………19 10. Box and whisker plot of wing petal lamina width (widest)…………………20 11. Box and whisker plot of the characters used for the fifth couplet……….21 12. Box and whisker plot of the characters used for the sixth couplet………22 13. Box and whisker plot of the characters used for the seventh couplet…..23 14. Box and whisker plot of the characters used for the eighth couplet…….24 15. Differences in terminal leaflet shapes and sizes……………………………..44 16. Line diagram comparing the mean petal shapes and sizes………………..45 17. Photograph of the Trifolium amabile type specimen…………………………57 18. Trifolium amabile…………………………………………………………………….58 19. Distribution of Trifolium amabile………………………………………………...59 20. Trifolium blanquitum……………………………………………………...... 63 21. Distribution of Trifolium blanquitum, T. cognatum, and T. lozani…………64 22. Trifolium cognatum………………………………………………………………….68 23. Trifolium goniocarpum………………………………………………………………73 24. Distribution of Trifolium goniocarpum…………………………………………..74 25. Trifolium hickeyi………………………………………………………………….....79 26. Distribution of Trifolium hickeyi and T. mexicanum…………………………80 27. Trifolium laciae……………………………………………………………………….84 28. Distribution of Trifolium laciae and T. sonoranensis………………………..85

v 29. Trifolium lozani……………………………………………………………...... ….89 30. Trifolium mexicanum………………………………………………………………..94 31. Trifolium sonoranensis…………………………………………………...... 98

vi Dedication I would like to dedicate this thesis to my mother for all of her love and support.

vii Acknowledgements First I would like to thank my family for all of their support. I would also like to thank my advisor and committee for all of their help and advice they gave me. Finally I would like to thank my friends and fellow graduate students for their advice and support.

viii Introduction The legume family, Fabaceae, is the third largest family of angiosperms after the Orchidaceae and Asteraceae, with more than 727 genera and 19,000 species (Lewis et al., 2005), and second in terms of economic importance (Judd et al., 2008), after the Poaceae. Despite variation in floral structure, Fabaceae is held together by the presence of its unique fruit, the legume (Lewis et al., 2005). Other unifying features include a unilocular superior ovary, parietal placentation, compound leaves (or simple leaves derived from ancestrally compound leaves), pulvini, and stipules (Lewis et al., 2005; Wojciechowski et al., 2006). Traditionally there have been three subfamilies within the Fabaceae, the Caesalpinioideae, the Mimosoideae, and the Papilionoideae, the latter being by far the largest of the three with 476 genera and over 13,800 species (Wojciechowski, 2006). However, recent molecular phylogenetic analyses have shown that while the Fabaceae is a monophyletic unit, the caesalpinioids are paraphyletic, and the mimosoids (excluding the genus Dinizia) and papilionoids are each monophyletic (Wojciechowski et al., 2004). The Papilionoideae are characterized by having a papilionaceous flower, that is with a banner petal, two wing petals, and two keel petals, in which the banner petal is the outermost in bud (Judd et al., 2008). Within this subfamily are many agriculturally important taxa including Medicago (alfalfa), Glycine (soybean), Arachis (peanut), and sixteen or more species of Trifolium (clover), which are important food and fodder crops (Ellison et al., 2006; Lewis et al., 2005). There are 36 tribes within the family, with 28 in the Papilionoideae (Lewis et al., 2005). The tribe Trifolieae contains six genera and over 480 species, more than half of which (about 250 species) belong to the genus Trifolium (Zohary and Heller, 1984; Lewis et al., 2005); some of these have been recognized since the times of Theophrastus, Dioscorides, and Pliny (Zohary and Heller, 1984). Other genera in this tribe include Parochetus, Ononis, Melilotus, Trigonella, and Medicago (Lewis et al., 2005). There are three centers of

1 diversity of Trifolium, two in the Old World and one in the New World (Ellison et al., 2006). The diversity in the Old World is concentrated around the Mediterranean basin and the eastern African highlands, and diversity in the New World is centered in western North America (Ellison et al., 2006). Trifolium species are mostly sun-tolerant and occur in a variety of different habitats from prairies and meadows to alpine peaks (Ellison et al., 2006). Despite its large size and economic importance, relatively little comprehensive work has been done on the genus, especially the New World taxa (Zohary and Heller, 1984). According to Zohary and Heller (1984), the genus Trifolium is divisible into eight sections, but most of these have been shown not to be monophyletic (Ellison, et al., 2006). Only two of Zohary and Heller’s sections, Lotoidea and Involucrarium, occur natively in the New World. Recent molecular analyses of cpDNA loci matK and trnL, and nrDNA locus ITS have shown that Zohary and Heller’s (1984) section Lotoidea is polyphyletic (Ellison, et al., 2006). All New World Trifolium formerly placed in section Lotoidea are now included in a more broadly circumscribed, monophyletic, section Involucrarium (Steele and Wojciechowski, 2003; Ellison et al., 2006). Section Trifoliastrum, containing species such as Trifolium repens L., has been shown to be sister to section Involucrarium (Steele and Wojciechowski, 2003; Ellison et al., 2006). Within section Involucrarium, Trifolium amabile Kunth is sister to the rest of the section excluding T. breweri S. Watson (Ellison et al., 2006). The Trifolium amabile species complex is a very widespread group, ranging from the mountains of southern Arizona, through the mountainous regions of Central America, and continuing southward to the province of Córdoba in north-central Argentina. In addition to its extensive geographical distribution, members of this species complex vary greatly in their size, habit, and morphological features. Trifolium amabile (Fig. 1) was described by Kunth in 1824, from material collected in Mexico by Humboldt and Bonpland. Since then, over thirty names have been published for members of this complex (Table 1). In Zohary and Heller’s (1984) monograph, the most recent treatment

2 of this group, the authors recognize five varieties of T. amabile (based on pubescence characters, calyx lobe shapes, leaflet shape, leaflet size, leaflet apex shape, and inflorescence size): var. amabile, var. hemsleyi (Lojac.) Heller & Zoh., var. pentlandii Ball, var. mexicanum (Hemsl.) Heller & Zoh., and var. longifoliolum Hemsl. However, it has been noted that this treatment is outdated and inadequate. For example, McVaugh (1987) called for a revision of the Mexican species. He especially commented on the differences between T. amabile and T. goniocarpum Lojac., included in var. longifoliolum. Even the monographers themselves noted that the distinctions among the varieties are not clear and difficult to distinguish (Zohary and Heller, 1984). The Trifolium amabile species complex has puzzled taxonomists for some time and this research will provide a much-needed evaluation of the North American members of this group. A thorough morphological analysis should provide a better understanding of this group by elucidating some of the complex taxonomic problems in this group. My primary goal is to present a better understanding of the North American members of the T. amabile species complex and contribute to the limited body of knowledge about New World clover species by performing morphological analyses to determine taxon limits. South American members of this group are being revised separately by Justin Hendy. The following hypothesis and questions will be addressed: Hypothesis: The Trifolium amabile Kunth species complex in North America represents a single unique species. Questions to be addressed: 1. If T. amabile is supported as a single species, what are the unique morphological features that distinguish this species from other species of Trifolium? 2. If T. amabile is not a supported as a single species, which sets of characters serve to delimit the various species identified?

Figure 1. Illustration of Trifolium amabile Kunth in the protologue (Nov. Gen. Sp. 6: 503 (1824)).

4 Table 1. List of published names for the Trifolium amabile species complex. Names in bold-faced type are recognized by Zohary and Heller (1984). Names Published Place of Publication Type Trifolium amabile Kunth Nov. Gen. Sp. 6: 503 (1824). Humboldt & Bonpland s.n. (P) T. amabile var. amabile T. humboldtii Spreng., Syst. 3: 213 (1826). Humboldt s.n. (B,W) Lupinaster amabilis (H., B. & Symb. Bot. 1: 47 (1830). K.) Presl T. pauciflorum Willd. ex Steud. Nom. ed. 2, 2: 707 (1841). Humboldt s.n. (B,W) (=T. nom.illeg. humboldtii) T. mathewsii A. Gray Bot. U. St. Expl. Exped. 1: 398 Matthews s.n. (K) (1854). T. schiedeanum S. Wats. Proc. Am. Acad. 17: 339 Schiede & Deppe 14173 (1882). (B,W) T. lozani House Coult. Bot. Gaz. 41: 342 Pringle 9512 (US) (1906). T. bolivianum Kennedy Muhlenbergia 7: 97 (1911). Bang 2819 (MICH) T. amabile var. hemsleyi Gen. Trifolium 127 (1984). (Lojac.) Heller & Zoh. T. hemsleyi Lojac. Nuov. Giorn. Bot. Ital. 15: 143, Palmer & Parry 136 t. IV, f. 1 (1883). (PAL, K) T. macrorrhizum Ulbrich Fedde Repert. 2: 2 (1906). Weberbauer 3213 (B) T. amabile var. pentlandii Jour. Linn. Soc. Bot. 22: 35 J. Ball s.n. (K) Ball (1885). T. chiclense Ball Jour. Linn. Soc. Bot. 22: 35 J. Ball s.n. (K) (different (1885). specimen than above) T. peruvianum Vogel Nov. Act. Nat. Car. 19: Suppl. Vogel? (BHU or HAL or 1: 12 (1843). K or CGE) T. peruvianum var. chiclense Meyen, Obs. Bot. 12. (1843). See T. chiclense (Hull) Macbr. T. weberbaueri Ulbrich Feddes Repert. 2: 2 (1906). Weberbauer 404, 3965 (B) T. amabile var. mexicanum Gen. ~Trifolium~ 128 (1984). (Hemsl.) Heller & Zoh. T. mexicanum Hemsl. Biol. Centr. Am. Bot. 1: 233 Parry & Palmer 137, (1879). Botteri 704, Graham, Coulter (K) T. potosanum Lojac. Nouv. Giorn. Bot. Ital. 15: 144 Parry & Palmer 137 (1883). (PAL, K) T. cognatum House Coult. Bot. Gaz. 41: 345 Pringle 6933 (US) (1906). T. nelsonii House Coult. Bot. Gaz. 41: 344 E. W. Nelson 1016 (US) (1906). T. amabile var. Biol. Centr. Am. Bot. 1: 232 Parry & Palmer 134, longifoliolum Hemsl. (1879). Botteri 703 (K) T. rhombeum S. Schau. Linnaea 20: 740 (1847). Aschenb. 164 (BR, K, REG) T. goniocarpum Lojac. Nuov. Giorn. Bot. Ital. 15: 145 Parry & Palmer 134? (1883) p.p. (PAL, K) T. longifoliolum (Hemsl.) House Coult. Bot. Gaz. 41: 342 (1906) T. schneideri Standl. Field Mus. Pub. Bot. 22:83. Schneider 942 (F) (1940). 5 Materials and Methods Specimens spanning the morphological diversity and North American range of this species complex were obtained from herbaria worldwide, a total of 948 specimens from 24 herbaria (Table 2). Exemplar specimens, representing the morphological diversity and geographic range of this group, were selected for measurements; a complete list of specimens measured is included in Appendix 1. Both qualitative and quantitative characters (Appendix 2) were chosen for analysis; selected characters are illustrated in Figs. 2-5. Pubescence characters were scored based on their density (Fig. 6). Qualitative descriptions of shape were based on the Systematics Association Committee for Descriptive Biological Terminology (1962) and Harris and Harris (2001). Linear measurements of the macroscopic structures were made from herbarium specimens in millimeters using a Schaedler Precision Rule and a Manostat caliper. Measurements of the smaller structures were made using a Wild MC3 dissecting scope and a calibrated ocular micrometer. Angle measurements were made using a Bausch & Lomb Measuring Magnifier Protractor Scale. Flowers and fruits were removed from the specimens and rehydrated using a 1% v/v dilution of Aerosol OT solution in distilled water. The dilution was made from a stock solution containing 10% Aerosol OT, 20% ethanol, and 70% distilled water (v/v/v). Quantitative data is listed in Appendix 3 and qualitative data is listed in Appendix 4. Analyses were performed using Numerical Taxonomy System (NTSYSpc) (Rohlf, 2000) to first standardize the data, and then perform a similarity matrix, using a sequential, agglomerative, hierarchical, non-overlapping (SAHN) clustering analysis, and a principal components analysis (PCA). These analyses identified preliminary species groups. As more specimens were measured, the data were combined with existing data and the same tests were run in NTSYSpc. These groups were subjected to refinement and further testing in R (R Development Core Team, 2010) to calculate the minimum, maximum, quartiles, median, mean, standard deviation, variance, as well as Shapiro-Wilk Normality Tests, and two-sided Student's t-Tests between 6 pairings of the groups of specimens. In addition, the “car” (Fox et Weisberg, 2010) and “lattice” (Sarkar, 2010) packages in R were used to create comparative box and whisker plots between pairings of the groups. These tests were used to analyze how well each group held together and how much they differed from each other. Outlier specimens differing in several characters from the rest of their group were removed and added to a group whose data were more similar. Groups that had few to no significantly different characters were merged to become one. After any change was made to the groups, the tests in R were run again. This process was repeated until the groups were composed of specimens whose data were analogous, and had significant differences from the other groups that could be used to key out each individual group. Using the statistically significant characters between each group pairing, data sets were made and final PCAs were run in R for each group pairing. Both two- and three-dimensional plots were constructed with the first two and three principal components, respectively. The “scatterplot3d” (Ligges and Mächler, 2011) package was used to make the three-dimensional plots. These results were congruent with the previous results from the t-tests of nine different groups which will be presented here as nine different species. Figures and illustrations were created in Photoshop (Adobe, 2010b) and Illustrator (Adobe, 2010a). Maps were created in ArcGIS (ESRI, 2010) and Illustrator.

7 Table 2. List of herbaria, and number of specimens loaned for study. Acronyms courtesy of Index Herbariorum (Thiers [continuously updated]). Acronym Institution Number of Specimens A Harvard University 8 AAU University of Aarhus 2 ARIZ University of Arizona 41 ASU Arizona State University 14 B Botanischer Garten und Botanisches Museum 7 Berlin-Dahlem, Zentraleinrichtung der Freien Universität Berlin C University of Copenhagen 15 DAV University of California 7 F Field Museum of Natural History 59 GH Harvard University 107 ILL University of Illinois 8 MEXU Universidad Nacional Autónoma de México 66 MICH University of Michigan 123 MO Missouri Botanical Garden 20 MU Miami University 6 NY New York Botanical Garden 160 P Muséum National d'Histoire Naturelle 58 POM Pomona College 14 RSA Rancho Santa Ana Botanic Garden 15 SI Museo Botánico (SI) 3 TEX University of Texas at Austin 4 UAT Universidad Autónoma de Tamaulipas 1 US Smithsonian Institution 134 VT University of Vermont 27 XAL Instituto de Ecología, A.C. 49

8 Terminal Leaflet

Petiolule

Petiole Stipules

Figure 2. Diagram of selected vegetative characters used for analysis. All measurements made in mm. 9

Figure 3. Diagram of selected inflorescence characters used for analysis. All measurements made in mm. 10

Figure 4. Diagram of selected floral characters used for analysis. All measurements made in mm. 11

Figure 5. Diagram of selected fruit characters used for analysis. All measurements made in mm.

Figure 6. Diagram of pubescence characters. A: sparse= ≥1mm between hairs, very sporadic; B: moderate= <1mm between hairs, uniformly spaced, able to see the underlying structure beneath the hairs; C. dense= so many hairs that the underlying structure is obscured.

13 Results This examination of specimens has shown that the T. amabile species complex is composed of nine morphologically distinct species. Five of these species have already been described in the literature: T. amabile Kunth, T. cognatum House, T. goniocarpum Lojac., T. lozani House, and T. mexicanum Hemsl. The remaining four species will be described as new species: T. blanquitum T.K. Ahlquist et Vincent sp. nov., T. hickeyi T.K. Ahlquist et Vincent sp. nov., T. laciae T.K. Ahlquist et Vincent sp. nov., and T. sonoranensis T.K. Ahlquist et Vincent sp. nov. Initial analyses in NTSYSpc showed the potential of multiple species within this complex. Both a PCA and SAHN clustering analysis showed the same preliminary groups of specimens (Fig. 7). As more specimens were measured, they were added to the existing data set. Due to the increasing size of the data set, it was split into many smaller data sets that were easier to manage. PCA and SAHN clustering analyses were run again on these smaller data sets, which provided more support for multiple species (Appendix 5). The largest species group, which was later split, showed a very widespread distribution (Fig. 8). These datasets were imported into R for further refinement. In R, various descriptive statistics and plots were run to help better define the groups and identify possible incongruences. After many rounds of statistics, the groups were composed of like specimens with as few anomalies as possible. Box and whisker plots were especially helpful in finding characters that made each group distinct, and are shown in Figures 9-14. These plots were also used to make the key. In addition, the t-tests run between pairs of groups to find the significantly different characters between them can be found in Tables 3-11. In addition to the quantitative characters, qualitative characters also differ between the species. Vegetative characters, especially leaflet size and shape (Fig. 15), can be useful in distinguishing the

14 different species. Reproductive characters, such as petal shape and size (Figure 16), also vary between the species. After the groups were refined trough t-tests, a final run of principal component analyses were run in R. These results (Appendix 6) coincided with the t-test results, providing further evidence that the groups were, in fact, separate species. In the subsequent pages, these species will be described in detail.

Figure 7. Preliminary results from NTSYSpc analyses. A: A three dimensional PCA projection plot showing six preliminary groups (designated by color and symbol). The axes are the principal components showing the most variation; B: A dendrogram resulting from a SAHN clustering analyses with the same six preliminary groups.

Figure 8a. Results from a PCA of the largest group.

Figure 8b. Results from a SAHN clustering analysis of the largest group.

Figure 9. Box and whisker plot of peduncle length used for the first couplet in the key.

Figure 10. Box and whisker plot of wing petal lamina width (widest) used for the fourth couplet in the key.

Figure 11. Box and whisker plot of the characters; A: keel petal lamina length, and B: inflorescence width, used for the fifth couplet in the key.

Figure 12. Box and whisker plot of the characters; A: internode length, B: terminal leaflet length, and C: terminal leaflet width, used for the sixth couplet in the key. 22

Figure 13. Box and whisker plot of the characters; A: sepal lobe length, B: keel petal claw length, and C: connate filament length, used for the seventh couplet in the key.

Figure 14a-c. Box and whisker plot of the characters; A: sepal length, B: banner petal length, and C: wing petal claw length, used for the eighth couplet in the key.

Figure 14d-f. Box and whisker plot of the characters; D: keel petal length, E: keel petal claw length, and F: ovary length, used for the eighth couplet in the key.

25 Table 3. P values resulting from a Student’s t-Test for T. amabile when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis StemL 0.225 0.2271 0.4677 0.001843 0.5474 0.0259 0.04261 0.9753 StemD 0.2287 0.0444 0.0168 9.54E-05 0.0992 0.5564 9.9E-05 0.5802 InNodL 0.861 0.1909 0.5515 2.29E-05 0.1972 0.0367 0.0015 0.2934 StipL 0.0238 0.5785 0.0776 1.11E-05 0.0110 0.0069 0.00204 0.1346 StipW 0.3406 0.2052 0.9291 5.66E-06 0.0903 0.5102 0.01126 0.2412 StipApx 0.0199 0.2653 0.0011 0.05546 0.3814 0.1707 0.03019 0.1783 Ang PetL 0.8301 0.477 0.6739 0.001668 0.3813 0.1916 0.2894 0.7374 PetD 0.3712 0.1276 0.0134 8.60E-07 0.1056 0.9638 0.00018 0.0177 LfltBasA 0.9079 0.0865 4.8E-05 0.03487 0.28 0.5612 0.1175 0.9262 ng TermLflt 0.7722 0.0299 0.00012 9.43E-07 0.0005 0.891 6.7E-06 0.0047 L TermLflt 0.299 0.9037 0.2425 5.78E-05 0.0062 0.2014 0.00031 0.0218 W PetlleL 0.9874 0.1689 0.06911 0.01192 0.0012 0.1064 2.4E-05 0.0610 PetlleD 0.0362 0.1762 0.2845 0.000626 0.1697 0.9281 0.00435 0.0114 InflL 0.2282 0.0608 0.1961 1.83E-08 0.6903 0.7937 3.1E-08 0.5807 InflW 0.0355 1.2E-05 0.4872 1.97E-12 0.0710 0.5671 5.8E-09 0.0379 FlNo 0.3207 0.0251 0.01066 1.61E-08 0.0021 0.0128 3.6E-07 0.1304 PeduncL 0.6888 0.019 0.0181 1.29E-09 0.2044 0.7462 0.00128 0.0051 Pedunc 0.0005 0.0501 0.0189 3.20E-07 0.0587 0.3214 2.6E-05 0.0279 D PedicL 0.6134 0.0095 0.1208 0.006092 0.0889 0.939 0.00012 0.0455 PedicD 3.9E-05 0.7102 0.221 0.165 0.2125 0.0250 0.4186 0.2596 SepL 0.0158 0.0335 0.2814 3.77E-07 0.1048 0.0106 8.6E-06 0.0034 SepCir 0.2836 0.5557 0.2274 0.7947 0.5049 0.2397 0.597 0.3382 SepLobL 0.02039 0.04364 0.1216 1.32E-06 0.0683 0.0323 2.2E-06 0.0036 SepLob 0.5646 0.3039 0.00583 0.005484 0.071 0.3395 0.3614 0.037 W SepLobA 1.6E-07 0.01823 0.1212 0.001194 0.5048 0.6578 0.00713 0.0698 pxAng BPetL 0.02568 0.07103 0.01604 1.22E-10 0.4808 0.0049 4.3E-08 0.0369 BPetW 0.00995 0.05488 0.00016 3.42E-08 0.8959 0.1201 6.3E-07 0.775 WPetL 0.00271 0.02771 9.9E-05 6.67E-10 0.36 0.0031 1.0E-07 0.0972 WPetClw 0.02245 0.5656 0.258 0.0159 0.0387 0.0104 1.8E-05 0.0001 L WPetClw 0.4528 0.4758 0.1406 0.001626 0.9402 0.4528 0.2013 0.9698 W WPetLa 0.00081 0.00715 2.0E-06 3.53E-10 0.574 0.0018 3.7E-07 0.8235 mL

26 Table 3 continued.

aciae T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. l T. lozani T. mexicanum T. sonoranensis WPetLa 0.02946 0.03666 1.8E-08 1.92E-09 0.0866 0.1135 1.0E-08 0.3845 mWw WPetLa 0.4292 0.00328 3.4E-05 9.13E-10 0.305 0.0010 1.6E-08 0.0023 mWn WPetLa 0.01982 0.2657 0.00509 0.00163 0.0053 0.0139 0.6182 0.6642 mArcAp xAng KPetL 0.0243 0.01266 1.0E-05 9.95E-11 0.3943 0.0018 1.1E-13 0.0022 KPetCl 0.0312 0.6458 0.2467 0.000413 0.0893 0.0202 1.3E-07 6.3E-06 wL KPetCl 0.478 0.07356 0.03546 0.001342 0.0755 0.1873 0.7013 0.7041 wW KPetLa 0.03544 2.7E-05 1.1E-06 1.14E-12 0.8228 0.0015 6.5E-10 0.1499 mL KPetLa 0.8902 0.00191 7.1E-07 2.65E-13 0.6936 0.107 2.8E-10 0.0338 mWw KPetLa 0.4705 0.00134 0.00116 3.88E-08 0.9843 0.0065 0.06704 0.1231 mWn ConFilL 0.00181 0.01789 0.00301 2.45E-10 0.7691 0.0226 1.3E-10 0.0003 ConFil 0.4153 0.00914 0.166 0.000855 0.3028 0.1976 0.1695 0.0226 W ConFilP 0.4618 0.3392 0.6166 0.000310 0.5252 0.1186 0.8074 0.4879 cFus FreFilL 0.00529 0.0444 0.1466 1.10E-05 0.824 0.1719 4.5E-09 0.0016 FreFilD 0.01464 0.524 0.00303 0.5414 0.0344 0.0208 0.3913 0.13 OvL 0.00239 0.07592 0.7833 0.05897 0.1559 0.0040 0.02842 0.3869 OvW 0.4613 0.522 0.7107 0.5198 0.1848 0.0178 0.4283 0.723 StylL 0.02297 6.6E-06 0.6698 2.49E-10 0.3059 0.7633 7.2E-11 0.0003 StylD 0.1334 0.00621 0.2745 3.48E-06 0.5176 5.8E-05 0.00810 0.2515 % Sig. 48.98% 42.86% 48.98% 87.76% 16.33% 44.90% 75.51% 42.86%

27 Table 4. P values resulting from a Student’s t-Test for T. blanquitum when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis StemL 0.225 0.535 0.12 0.00693 0.3554 0.6844 0.0307 0.2573 StemD 0.2287 0.01484 0.0062 5.7E-05 0.0322 0.613 0.0017 0.1424 InNodL 0.861 0.4536 0.6536 0.01285 0.6386 0.3197 0.0068 0.6629 StipL 0.02379 0.01501 0.0024 9.9E-07 0.001 0.9468 9E-05 0.0088 StipW 0.3406 0.06817 0.3968 1.1E-05 0.0266 0.8883 0.0033 0.9424 StipApx 0.01999 0.09286 0.0021 0.00433 0.0143 0.6377 0.0020 0.0048 Ang PetL 0.8301 0.4299 0.866 0.00159 0.3458 0.4985 0.271 0.6673 PetD 0.3712 0.07375 0.0074 2.8E-06 0.0586 0.4045 0.0007 0.0094 LfltBas 0.9079 0.3668 0.0677 0.1696 0.5233 0.5847 0.3023 0.8721 Ang TermLfl 0.7722 0.08282 0.0008 1.7E-06 0.0007 0.8513 0.0004 0.0189 tL TermLfl 0.299 0.3592 0.7704 0.00064 0.0382 0.9245 0.0011 0.0290 tW PetlleL 0.9874 0.2747 0.219 0.01706 0.0242 0.3239 0.0301 0.0887 PetlleD 0.03627 0.00852 0.0085 6.5E-05 0.0169 0.1954 0.0002 0.0011 InflL 0.2282 0.02492 0.0631 8.9E-06 0.235 0.2508 7.9E-05 0.1441 InflW 0.03552 1.6E-05 0.0094 6.9E-10 0.0099 0.3554 9.4E-09 0.0017 FlNo 0.3207 0.01321 0.0054 9.9E-08 0.0055 0.0062 8.4E-07 0.0345 Pedunc 0.6888 0.016 0.0321 2.4E-09 0.1745 0.838 0.02067 0.0435 L Pedunc 0.00046 0.01224 1.4E-05 2.4E-08 0.0075 0.0618 1.4E-07 0.0002 D PedicL 0.6134 0.00635 0.09742 0.00667 0.0733 0.8271 0.0004 0.0341 PedicD 3.9E-05 0.03726 0.00082 6.8E-06 0.0103 0.2825 0.00037 0.1177 SepL 0.01575 0.00160 0.01173 9.4E-08 0.0197 0.6699 2.5E-06 0.0002 SepCir 0.2836 0.01401 0.3054 1.4E-05 0.0334 0.4582 0.00054 0.3662 SepLob 0.02039 0.00215 0.00371 6.0E-07 0.0109 0.4456 7.1E-06 0.0004 L SepLob 0.5646 0.315 0.02159 0.0652 0.4012 0.2222 0.8019 0.0506 W SepLob 1.6E-07 0.00084 9.3E-06 1.2E-09 0.0993 0.0039 1.1E-07 0.0004 ApxAng BPetL 0.02568 0.01035 0.4261 1.6E-06 0.1751 0.3058 5.6E-05 0.0028 BPetW 0.00995 0.00156 0.6005 1.1E-07 0.1268 0.3265 3.1E-06 0.0066 WPetL 0.00271 0.00157 0.6795 1.9E-07 0.0572 0.6096 7.6E-06 0.0009 WPetCl 0.02245 0.05369 0.08722 0.00034 0.4491 0.2822 5.9E-05 0.0002 wL WPetCl 0.4528 0.3739 0.1921 0.00074 0.704 NA 0.03233 0.689 wW

28 Table 4 continued.

T. amabile T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis WPetLa 0.00081 0.00044 0.6823 1.7E-09 0.0198 0.9634 7.0E-07 0.0017 mL WPetLa 0.02946 0.01228 0.01677 9.7E-09 0.5796 0.4554 1.9E-06 0.0094 mWw WPetLa 0.4292 0.00135 0.03291 3.2E-06 0.1635 0.1248 0.00017 0.0112 mWn WPetLa 0.01982 0.01862 0.6716 0.9278 0.7926 0.7245 0.03503 0.0149 mArcAp xAng KPetL 0.0243 0.00088 0.7703 8.4E-06 0.1126 0.3713 9.6E-05 0.0007 KPetCl 0.0312 0.1381 0.1336 6.7E-05 0.9384 0.3492 2.1E-05 0.0002 wL KPetCl 0.478 0.04974 0.4972 0.00336 0.2907 0.5796 0.3626 0.3949 wW KPetLa 0.03544 0.00031 0.6643 5.1E-05 0.0459 0.5596 0.00026 0.008 mL KPetLa 0.8902 0.00132 0.00767 9.2E-07 0.6598 0.2029 8.2E-05 0.0877 mWw KPetLa 0.4705 0.00523 0.00476 2.8E-05 0.6234 0.009 0.2578 0.2692 mWn ConFilL 0.00181 0.00355 0.07081 2.2E-10 0.0881 0.5281 1.1E-09 9E-06 ConFil 0.4153 0.016 0.9287 0.03163 0.1645 0.7309 0.1091 0.0366 W ConFilP 0.4618 0.9795 0.6524 0.01066 0.7443 0.3953 0.5536 0.6865 cFus FreFilL 0.00529 0.00592 0.01857 4.2E-07 0.0828 0.3825 3E-06 0.0002 FreFilD 0.01464 0.1778 0.05096 0.1271 0.1817 0.1778 0.0217 0.7734 OvL 0.00239 0.00674 0.03142 0.00018 0.0212 0.5827 0.0003 0.0248 OvW 0.4613 0.3077 0.7974 0.2546 0.1187 0.1299 0.2345 0.4582 StylL 0.02297 1.9E-06 0.2751 7.4E-11 0.625 0.1054 8.6E-10 3.7E-05 StylD 0.1334 0.00243 0.134 3.2E-05 0.6833 0.1778 0.00323 0.1787 % Sig. 48.98% 65.31% 42.86% 89.80% 36.73% 6.25% 83.67% 61.22%

29 Table 5. P values resulting from a Student’s t-Test for T. cognatum when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis StemL 0.2271 0.535 0.097 0.0002 0.5378 0.1374 0.0043 0.3695 StemD 0.04435 0.01484 0.8593 0.0309 0.4501 0.0328 0.7117 0.0660 InNodL 0.1909 0.4536 0.4575 4.9E-06 0.0227 0.4433 0.0006 0.0510 StipL 0.5785 0.01501 0.1673 3.5E-05 0.0302 0.0104 0.0256 0.2499 StipW 0.2052 0.06817 0.1906 0.00473 0.8205 0.1125 0.3948 0.0597 StipApx 0.2653 0.09286 0.0039 0.019 0.1315 0.4106 0.0096 0.0488 Ang PetL 0.477 0.4299 0.3525 0.00535 0.735 0.1836 0.7478 0.8788 PetD 0.1276 0.07375 0.3209 0.03204 1 0.1331 0.6057 0.889 LfltBasA 0.08655 0.3668 0.0573 0.00254 0.7608 0.1496 0.0079 0.2691 ng TermLflt 0.02995 0.08282 0.0061 8.8E-06 0.0035 0.0391 0.0016 0.2426 L TermLflt 0.9037 0.3592 0.3662 0.00021 0.043 0.29 0.0007 0.0410 W PetlleL 0.1689 0.2747 0.976 0.06478 0.1481 0.0366 0.1837 0.347 PetlleD 0.1762 0.00852 0.6602 0.00348 0.6684 0.4933 0.1088 0.0727 InflL 0.06075 0.02492 0.1174 0.1022 0.1023 0.0813 0.3431 0.0843 InflW 1.2E-05 1.6E-05 6.2E-05 0.01594 0.0183 0.0006 0.5561 0.0034 FlNo 0.02507 0.01321 0.1345 0.04133 0.1452 0.9256 0.1379 0.0819 PeduncL 0.01893 0.016 0.3757 6.3E-07 0.1498 0.0164 0.3349 0.1828 Pedunc 0.05014 0.01224 0.1435 0.01644 0.4797 0.3128 0.915 0.1857 D PedicL 0.0095 0.00635 0.04442 0.4641 0.6044 0.0270 0.85 0.2666 PedicD 0.7102 0.03726 0.3874 0.7695 0.4861 0.0950 0.5171 0.2995 SepL 0.03346 0.00160 0.4921 0.00036 0.5667 0.0011 0.0229 0.3786 SepCir 0.5557 0.01401 0.00323 0.05819 0.6726 0.0075 0.6885 0.0428 SepLobL 0.04364 0.00215 0.8128 0.00086 0.3695 0.0037 0.0121 0.4299 SepLob 0.3039 0.315 0.2646 0.3632 0.3322 0.2905 0.3207 0.2731 W SepLobA 0.01823 0.00084 0.2244 0.87 0.0557 0.0667 0.8455 0.585 pxAng BPetL 0.07103 0.01035 0.01917 0.04482 0.0425 0.0044 0.4377 0.2565 BPetW 0.05488 0.00156 0.00101 0.0012 0.1538 0.008 0.0203 0.0730 WPetL 0.02771 0.00157 0.00269 0.05423 0.0128 0.001 0.519 0.0842 WPetClw 0.5656 0.05369 0.2474 0.2948 0.0998 0.0173 0.0601 0.0969 L WPetClw 0.4758 0.3739 0.2555 0.1976 0.5439 0.3739 0.7976 0.481 W

30 Table 5 continued.

T. amabile T. blanquitum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis WPetLa 0.00715 0.00044 0.00052 0.0282 0.0047 0.0004 0.7968 0.0076 mL WPetLa 0.03666 0.01228 0.00576 0.1697 0.0159 0.0174 0.9677 0.0536 mWw WPetLa 0.00328 0.00135 0.00033 0.1066 0.0126 0.0008 0.68 0.0268 mWn WPetLa 0.2657 0.01862 0.02494 0.01472 0.0134 0.0129 0.1812 0.4021 mArcAp xAng KPetL 0.01266 0.00088 0.00175 0.02823 0.0059 0.0003 0.1078 0.3196 KPetClw 0.6458 0.1381 0.3901 0.1472 0.1452 0.06 0.0624 0.1318 L KPetClw 0.07356 0.04974 0.01015 0.2102 0.0095 0.0203 0.1434 0.231 W KPetLam 2.7E-05 0.00031 9.1E-06 0.00395 0.0001 1.0E-05 0.452 0.0023 L KPetLam 0.00191 0.00132 0.00031 0.01569 0.004 0.00038 1 0.0070 Ww KPetLam 0.00134 0.00523 0.00042 0.1434 0.0116 0.0005 0.0223 0.1071 Wn ConFilL 0.01789 0.00355 0.00814 0.1056 0.0132 0.00153 0.2846 0.3263 ConFilW 0.00914 0.016 0.00167 0.407 0.4028 0.00588 0.1914 0.46 ConFilPc 0.3392 0.9795 0.5437 0.00177 0.6591 0.3581 0.4358 0.5717 Fus FreFilL 0.0444 0.00592 0.03092 0.4054 0.0859 0.01272 0.6652 0.2508 FreFilD 0.524 0.1778 0.05319 0.3754 0.0605 0.07189 0.9096 0.2115 OvL 0.07592 0.00674 0.1661 0.6007 0.9523 0.00411 0.8884 0.4273 OvW 0.522 0.3077 0.4129 0.8889 0.418 0.07258 0.918 0.8404 StylL 6.6E-06 1.9E-06 0.02213 0.00017 0.0002 1.9E-05 0.0872 0.0329 StylD 0.00621 0.00243 0.7805 0.02068 0.0371 0.00085 0.5163 0.4478 % Sig. 42.86% 65.31% 42.86% 59.18% 38.78% 59.18% 22.45% 18.37%

31 Table 6. P values resulting from a Student’s t-Test for T. goniocarpum when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis StemL 0.4677 0.12 0.097 0.02572 0.2296 0.0127 0.2599 0.5771 StemD 0.01676 0.0062 0.8593 0.04024 0.3494 0.0176 0.8801 0.0300 InNodL 0.5515 0.6536 0.4575 9.9E-06 0.0698 0.1289 0.001 0.1341 StipL 0.07755 0.0024 0.1673 0.00094 0.4659 0.0018 0.7754 0.8977 StipW 0.9291 0.3968 0.1906 1.0E-05 0.0851 0.5631 0.0119 0.315 StipApx 0.00114 0.0021 0.0039 0.1225 0.3002 0.0263 0.5062 0.3982 Ang PetL 0.6739 0.866 0.3525 0.00126 0.2976 0.622 0.2153 0.594 PetD 0.01342 0.0074 0.3209 0.2684 0.305 0.0146 0.4502 0.3072 LfltBasA 4.8E-05 0.0677 0.0573 7.2E-06 0.0820 0.0304 2.1E-05 0.0373 ng TermLflt 0.00012 0.0008 0.0061 0.00146 0.5848 0.0002 0.7918 0.0599 L TermLflt 0.2425 0.7704 0.3662 1.2E-05 0.0014 0.7985 6.3E-05 0.0035 W PetlleL 0.06911 0.219 0.976 0.0592 0.0818 0.0057 0.09107 0.3288 PetlleD 0.2845 0.0085 0.6602 0.00184 0.4475 0.6507 0.03749 0.0369 InflL 0.1961 0.0631 0.1174 3.1E-07 0.7368 0.5752 2.4E-06 0.5832 InflW 0.4872 0.0093 6.2E-05 1.9E-09 0.155 0.2931 3.7E-07 0.1181 FlNo 0.01066 0.0054 0.1345 1.7E-06 0.8489 0.1001 3.9E-05 0.4787 PeduncL 0.01805 0.0321 0.3757 5.7E-09 0.4443 0.0122 0.99 0.6122 Pedunc 0.01894 1.4E-05 0.1435 2.3E-06 0.2803 0.8096 0.00314 0.6792 D PedicL 0.1208 0.09742 0.04442 0.1248 0.1631 0.3469 0.0082 0.3311 PedicD 0.221 0.00082 0.3874 0.01963 0.0997 0.0967 0.4682 0.218 SepL 0.2814 0.01173 0.4921 0.00024 0.3312 0.007 0.01269 0.1706 SepCir 0.2274 0.3054 0.00323 3.7E-07 0.0085 0.8407 3.9E-05 0.0776 SepLobL 0.1216 0.00371 0.8128 0.00095 0.318 0.0076 0.01376 0.3648 SepLob 0.00583 0.02159 0.2646 2.4E-05 0.0005 0.0729 0.00357 0.5281 W SepLobA 0.1212 9.3E-06 0.2244 0.1751 0.166 0.3626 0.2553 0.5579 pxAng BPetL 0.01604 0.4261 0.01917 1.6E-11 0.3642 0.0716 1.6E-09 0.0003 BPetW 0.00016 0.6005 0.00101 5.5E-11 0.0800 0.1367 1.5E-11 0.0001 WPetL 9.9E-05 0.6795 0.00269 1.5E-12 0.0704 0.3563 9.7E-12 1.7E-06 WPetClw 0.258 0.08722 0.2474 0.00072 0.1844 0.0314 1.2E-07 4.5E-06 L WPetClw 0.1406 0.1921 0.2555 0.00027 0.4087 0.1921 0.01176 0.3141 W

32 Table 6 continued.

T. amabile T. blanquitum T. cognatum T. hickeyi T. laciae T. lozani T. mexicanum T. sonoranensis WPetLa 2E-06 0.6823 0.00052 7.6E-13 0.0105 0.745 6.4E-12 1.5E-05 mL WPetLa 1.8E-08 0.01677 0.00576 5.0E-11 0.0077 0.0050 1.1E-12 2.1E-05 mWw WPetLa 3.4E-05 0.03291 0.00033 3.4E-12 0.0077 0.1896 2.2E-11 1.3E-06 mWn WPetLa 0.00509 0.6716 0.02494 0.5405 0.4343 0.4268 0.01249 0.00806 mArcAp xAng KPetL 1.0E-05 0.7703 0.00175 9.9E-13 0.0846 0.1296 3.4E-15 1.9E-05 KPetClw 0.2467 0.1336 0.3901 8.2E-05 0.2274 0.0504 9.2E-08 3.7E-06 L KPetClw 0.03546 0.4972 0.01015 5.2E-05 0.5108 1 0.04544 0.1194 W KPetLam 1.1E-06 0.6643 9.1E-06 7.5E-15 0.0012 0.7383 3.3E-14 0.00029 L KPetLam 7.1E-07 0.0077 0.00031 6.8E-15 0.017 0.072 2.1E-13 1.9E-05 Ww KPetLam 0.00116 0.00476 0.00042 1.9E-10 0.1045 0.4375 3.5E-05 0.00627 Wn ConFilL 0.00301 0.07081 0.00814 5.8E-11 0.3448 0.1205 2.1E-11 3.7E-05 ConFilW 0.166 0.9287 0.00167 0.00038 0.1168 0.7421 0.03138 0.00374 ConFilPc 0.6166 0.6524 0.5437 8.7E-05 0.8541 0.1752 0.801 0.8976 Fus FreFilL 0.1466 0.01857 0.03092 4.6E-06 0.5333 0.3291 3.0E-09 0.00041 FreFilD 0.00303 0.05096 0.05319 0.01211 0.9832 0.7522 0.00233 0.07055 OvL 0.7833 0.03142 0.1661 0.2643 0.2539 0.0196 0.1155 0.5938 OvW 0.7107 0.7974 0.4129 0.3986 0.1494 0.1038 0.3378 0.5827 StylL 0.6698 0.2751 0.02213 0.00054 0.4021 0.6133 0.00693 0.1299 StylD 0.2745 0.134 0.7805 0.6406 0.196 0.067 0.6544 0.5701 % Sig. 48.98% 42.86% 42.86% 81.63% 16.33% 28.57% 69.39% 40.82%

33 Table 7. P values resulting from a Student’s t-Test for T. hickeyi when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. laciae T. lozani T. mexicanum T. sonoranensis StemL 0.0018 0.00693 0.0003 0.0257 0.0008 9.5E-05 0.1967 0.0135 StemD 9.5E-05 5.7E-05 0.0309 0.0402 0.0048 0.00017 0.0246 0.0002 InNodL 2.3E-05 0.01285 4.9E-06 9.9E-06 0.0004 2.0E-06 0.3607 0.0009 StipL 1.1E-05 9.9E-07 3.5E-05 0.0009 0.003 1.1E-06 0.0005 0.0012 StipW 5.7E-06 1.0E-05 0.00473 1.0E-05 0.0027 0.00014 0.0060 9.2E-07 StipApx 0.05546 0.00433 0.019 0.1225 0.7994 0.05338 0.5501 0.921 Ang PetL 0.00167 0.00159 0.00535 0.0013 0.0163 0.00079 0.0087 0.00782 PetD 8.6E-07 2.8E-06 0.03204 0.2684 0.0236 1.8E-06 0.0042 0.00516 LfltBasA 0.03487 0.1696 0.00254 7.2E-06 0.0182 0.4467 0.5666 0.2148 ng TermLflt 9.4E-07 1.7E-06 8.8E-06 0.00146 0.0035 8.8E-07 0.002 4.1E-05 L TermLflt 5.8E-05 0.00064 0.00021 1.2E-05 0.0027 4.3E-05 0.7874 0.00866 W PetlleL 0.01192 0.01706 0.06478 0.0592 0.23 0.00391 0.1853 0.2726 PetlleD 0.00063 6.5E-05 0.00348 0.00184 0.0115 0.00433 0.0331 0.09493 InflL 1.8E-08 8.9E-06 0.1022 3.1E-07 0.0006 0.00012 0.109 1.1E-06 InflW 2.0E-12 6.9E-10 0.01594 1.9E-09 0.0013 7.3E-05 0.0312 2.8E-05 FlNo 1.6E-08 9.9E-08 0.04133 1.7E-06 2.6E-06 0.01657 0.3164 2.6E-06 PeduncL 1.3E-09 2.4E-09 6.3E-07 5.7E-09 3.1E-08 1.7E-09 8.4E-09 9.0E-09 Pedunc 3.2E-07 2.4E-08 0.01644 2.3E-06 0.0006 0.00298 0.0002 4.5E-06 D PedicL 0.00609 0.00667 0.4641 0.1248 0.3915 0.06259 0.3159 0.6539 PedicD 0.165 6.8E-06 0.7695 0.01963 0.54 0.00498 0.0306 0.3233 SepL 3.8E-07 9.4E-08 0.00037 0.00024 0.0509 1.1E-07 0.0366 0.00141 SepCir 0.7947 1.4E-05 0.05819 3.7E-07 0.0299 0.00011 0.0331 3.4E-05 SepLobL 1.3E-06 6.0E-07 0.00086 0.00095 0.1112 2.5E-07 0.1232 0.00212 SepLob 0.00548 0.0652 0.3632 2.4E-05 0.1518 0.00156 0.0822 0.00015 W SepLobA 0.00119 1.2E-09 0.87 0.1751 0.0599 0.04738 0.9515 0.6253 pxAng BPetL 1.2E-10 1.6E-06 0.04482 1.6E-11 4.8E-05 1.0E-06 0.0016 3.0E-07 BPetW 3.4E-08 1.1E-07 0.0012 5.5E-11 0.0020 3.8E-06 0.0492 2.4E-07 WPetL 6.7E-10 1.9E-07 0.05423 1.5E-12 4.7E-05 1.5E-06 0.0060 6.7E-08 WPetClw 0.0159 0.00033 0.2948 0.00072 0.0002 0.00059 0.0981 0.2391 L WPetClw 0.00163 0.00074 0.1976 0.00027 0.0162 0.00074 0.0103 0.00352 W WPetLa 3.5E-10 1.7E-09 0.0282 7.6E-13 6.3E-06 1.3E-08 0.0002 2.2E-09 mL

34 Table 7 continued.

tum T. amabile T. blanquitum T. cogna T. goniocarpum T. laciae T. lozani T. mexicanum T. sonoranensis WPetLa 1.9E-09 9.7E-09 0.1697 5.0E-11 1.9E-08 1.6E-08 0.0025 4.7E-08 mWw WPetLa 9.1E-10 3.2E-06 0.1066 3.4E-12 0.0004 1.6E-10 0.004 2.1E-06 mWn WPetLa 0.00163 0.9278 0.01472 0.5405 0.8413 0.7621 0.0040 0.00302 mArcAp xAng KPetL 9.9E-11 8.4E-06 0.02823 9.9E-13 1.6E-05 9.9E-08 0.1116 5.6E-05 KPetClw 0.00041 6.7E-05 0.1472 8.2E-05 0.0011 0.00021 0.2812 0.9422 L KPetClw 0.00134 0.00336 0.2102 5.2E-05 0.0005 0.00115 0.0059 0.02438 W KPetLam 1.1E-12 5.1E-05 0.00395 7.5E-15 1.2E-07 2.0E-07 0.0035 8.5E-06 L KPetLam 2.7E-13 9.2E-07 0.01569 6.8E-15 8.3E-05 2.6E-07 7.5E-05 1.6E-08 Ww KPetLam 3.9E-08 2.8E-05 0.1434 1.9E-10 0.0019 1.3E-09 5.9E-05 0.00986 Wn ConFilL 2.5E-10 2.2E-10 0.1056 5.8E-11 7.8E-05 6.0E-06 0.1898 5.4E-05 ConFilW 0.00086 0.03163 0.407 0.00038 0.6779 0.01137 0.4154 0.9668 ConFilPc 0.00031 0.01066 0.00177 8.7E-05 0.0005 0.00447 0.0002 2.5E-05 Fus FreFilL 1.1E-05 4.2E-07 0.4054 4.6E-06 0.0119 0.00021 0.4315 0.00213 FreFilD 0.5414 0.1271 0.3754 0.01211 0.0615 0.04942 0.2038 0.3623 OvL 0.05897 0.00018 0.6007 0.2643 0.7108 0.00022 0.4834 0.675 OvW 0.5198 0.2546 0.8889 0.3986 0.3299 0.01932 0.7964 0.925 StylL 2.5E-10 7.4E-11 0.00017 0.00054 3.2E-07 2.2E-09 0.0051 7.4E-06 StylD 3.5E-06 3.2E-05 0.02068 0.6406 0.0035 2.6E-08 0.0027 0.6994 % Sig. 87.76% 89.80% 59.18% 81.63% 73.47% 91.84% 57.14% 71.43%

35 Table 8. P values resulting from a Student’s t-Test for T. laciae when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. lozani T. mexicanum T. sonoranensis StemL 0.5474 0.3554 0.5378 0.2296 0.0008 0.0662 0.0154 0.638 StemD 0.0992 0.0322 0.4501 0.3494 0.0048 0.0808 0.1809 0.1655 InNodL 0.1972 0.6386 0.0227 0.0698 0.0004 0.008 0.0045 0.9807 StipL 0.0110 0.001 0.0302 0.4659 0.003 0.0015 0.5093 0.4082 StipW 0.0903 0.0267 0.8205 0.0852 0.0026 0.0585 0.4748 0.0194 StipApx 0.3814 0.0143 0.1315 0.3002 0.7994 0.0847 0.5424 0.7743 Ang PetL 0.3813 0.3458 0.735 0.2976 0.0163 0.1962 0.9476 0.6763 PetD 0.1056 0.0586 1 0.305 0.0236 0.1104 0.5813 0.8824 LfltBasA 0.28 0.5233 0.7608 0.0820 0.0182 0.2315 0.044 0.4107 ng TermLflt 0.0005 0.0008 0.0035 0.5848 0.0036 0.0004 0.7701 0.0191 L TermLflt 0.0062 0.0382 0.043 0.0014 0.0026 0.0095 0.0098 0.7345 W PetlleL 0.0012 0.0242 0.1481 0.0818 0.23 0.0004 0.7425 0.9353 PetlleD 0.1697 0.0169 0.6684 0.4475 0.0115 0.3608 0.3606 0.2054 InflL 0.6903 0.235 0.1023 0.7368 0.0005 0.8878 0.0032 0.9725 InflW 0.0710 0.0099 0.0183 0.155 0.0013 0.0524 0.0084 0.8828 FlNo 0.0021 0.0055 0.1452 0.8489 2.6E-06 0.1098 4.4E-05 0.3544 PeduncL 0.2044 0.1745 0.1498 0.4443 3.1E-08 0.1576 0.3839 0.6424 Pedunc 0.0587 0.0076 0.4797 0.2803 0.00062 0.6262 0.3711 0.4026 D PedicL 0.0889 0.0733 0.6044 0.1631 0.3915 0.0883 0.6601 0.2978 PedicD 0.2125 0.0103 0.4861 0.0997 0.54 0.0222 0.1408 0.3954 SepL 0.1048 0.0197 0.5667 0.3312 0.05091 0.0147 0.3791 0.955 SepCir 0.5049 0.0334 0.6726 0.0085 0.02988 0.0168 0.3896 0.1028 SepLobL 0.0683 0.0109 0.3695 0.318 0.1112 0.0189 0.4462 0.64 SepLob 0.071 0.4012 0.3322 0.0005 0.1518 0.0203 0.5489 0.0024 W SepLobA 0.5048 0.0993 0.0558 0.166 0.05988 0.3815 0.06289 0.1013 pxAng BPetL 0.4808 0.1751 0.0425 0.3642 4.8E-05 0.0385 0.00111 0.0615 BPetW 0.8959 0.1268 0.1538 0.0800 0.00202 0.3537 0.01061 0.7684 WPetL 0.36 0.0572 0.0128 0.0704 4.7E-05 0.0336 0.00079 0.0689 WPetClw 0.0387 0.4491 0.0998 0.1844 0.00018 0.1045 3.2E-06 2.9E-05 L WPetClw 0.9402 0.704 0.5439 0.4087 0.01622 0.704 0.5194 0.9261 W WPetLa 0.574 0.0198 0.0047 0.0105 6.3E-06 0.0213 0.00094 0.6653 mL

36 Table 8 continued.

nensis

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. lozani T. mexicanum T. sonora WPetLa 0.0866 0.5796 0.0159 0.0077 1.9E-08 0.8494 3.6E-06 0.02547 mWw WPetLa 0.305 0.1635 0.0126 0.0077 0.00034 0.0206 0.00881 0.325 mWn WPetLa 0.0053 0.7926 0.0134 0.4343 0.8413 0.8994 0.01064 0.00517 mArcAp xAng KPetL 0.3943 0.1126 0.0059 0.0846 1.6E-05 0.0174 0.00019 0.00601 KPetClw 0.0893 0.9384 0.1452 0.2274 0.00114 0.4591 0.00068 0.00215 L KPetClw 0.0755 0.2907 0.0095 0.5108 0.00054 0.5871 0.05691 0.08196 W KPetLam 0.8228 0.0459 0.0001 0.0012 1.2E-07 0.0035 5.5E-06 0.1217 L KPetLam 0.6936 0.6598 0.004 0.017 8.3E-05 0.1507 0.00208 0.3297 Ww KPetLam 0.9843 0.6234 0.0116 0.1045 0.00194 0.1543 0.2218 0.2023 Wn ConFilL 0.7691 0.0881 0.0132 0.3448 7.8E-05 0.0633 0.00031 0.01032 ConFilW 0.3028 0.1645 0.4028 0.1168 0.6779 0.0975 0.9009 0.7119 ConFilPc 0.5252 0.7443 0.6591 0.8541 0.0005 0.2108 0.6821 0.9252 Fus FreFilL 0.824 0.0828 0.0859 0.5333 0.01189 0.2561 0.02881 0.2405 FreFilD 0.0344 0.1817 0.0605 0.9832 0.06152 0.8 0.0128 0.1601 OvL 0.1559 0.0212 0.9523 0.2539 0.7108 0.0145 0.8561 0.5186 OvW 0.1848 0.1187 0.418 0.1494 0.3299 0.0426 0.4645 0.343 StylL 0.3059 0.625 0.0002 0.4021 3.1E-07 0.4443 2.0E-05 0.00086 StylD 0.5176 0.6833 0.0371 0.196 0.00352 0.208 0.0611 0.2081 % Sig. 16.33% 36.73% 38.78% 16.33% 73.47% 36.73% 46.94% 20.41%

37 Table 9. P values resulting from a Student’s t-Test for T. lozani when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. mexicanum T. sonoranensis StemL 0.02591 0.6844 0.1374 0.0127 9.5E-05 0.0662 0.00099 0.0617 StemD 0.5564 0.613 0.0328 0.0176 0.00017 0.0808 0.0057 0.3742 InNodL 0.03669 0.3197 0.4433 0.1289 2.0E-06 0.008 0.00038 0.0201 StipL 0.00692 0.9468 0.0104 0.0018 1.1E-06 0.0015 4.7E-07 0.0088 StipW 0.5102 0.8883 0.1125 0.5631 0.00014 0.0585 0.01541 0.9159 StipApx 0.1707 0.6377 0.4106 0.0263 0.05338 0.0847 0.03594 0.053 Ang PetL 0.1916 0.4985 0.1836 0.622 0.00079 0.1962 0.08825 0.4191 PetD 0.9638 0.4045 0.1331 0.0146 1.8E-06 0.1104 0.0006 0.021 LfltBasA 0.5612 0.5847 0.1496 0.0304 0.4467 0.2315 0.7037 0.694 ng TermLflt 0.891 0.8513 0.0391 0.0002 8.8E-07 0.0004 2.0E-05 0.0064 L TermLflt 0.2014 0.9245 0.29 0.7985 4.3E-05 0.0095 0.00012 0.0071 W PetlleL 0.1064 0.3239 0.0366 0.0057 0.00391 0.0004 3.8E-05 0.0191 PetlleD 0.9281 0.1954 0.4933 0.6507 0.00433 0.3608 0.1059 0.0642 InflL 0.7937 0.2508 0.0813 0.5752 0.00012 0.8878 0.00096 0.8896 InflW 0.5671 0.3554 0.0006 0.2931 7.3E-05 0.0524 0.00022 0.0416 FlNo 0.01282 0.0062 0.9256 0.1001 0.01657 0.1098 0.07473 0.0558 Pedunc 0.7462 0.838 0.0164 0.0122 1.7E-09 0.1576 0.00111 0.0047 L Pedunc 0.3214 0.0618 0.3128 0.8096 0.00298 0.6262 0.2547 0.946 D PedicL 0.939 0.8271 0.0270 0.3469 0.06259 0.0883 0.01508 0.1297 PedicD 0.02501 0.2825 0.0950 0.0967 0.00498 0.0222 0.04902 0.1494 SepL 0.01058 0.6699 0.0011 0.007 1.1E-07 0.0147 3.1E-06 0.0002 SepCir 0.2397 0.4582 0.0075 0.8407 0.00011 0.0168 0.00115 0.1531 SepLob 0.03234 0.4456 0.0036 0.0076 2.5E-07 0.0189 1.2E-06 0.0003 L SepLob 0.3395 0.2222 0.2905 0.0728 0.00155 0.0203 0.1108 0.2308 W SepLob 0.6578 0.0039 0.0667 0.3626 0.04738 0.3815 0.06584 0.1887 ApxAng BPetL 0.0049 0.3058 0.0044 0.0716 1.0E-06 0.0385 3.0E-05 0.0008 BPetW 0.1201 0.3265 0.008 0.1367 3.8E-06 0.3537 7.3E-05 0.0739 WPetL 0.00313 0.6096 0.001 0.3563 1.5E-06 0.0336 3.2E-05 0.0012 WPetCl 0.01037 0.2822 0.0173 0.0314 0.00058 0.1045 0.00030 0.0006 wL WPetCl 0.4528 NA 0.3739 0.1921 0.00074 0.704 0.03233 0.689 wW

38 Table 9 continued.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. mexicanum T. sonoranensis WPetLa 0.00184 0.9634 0.0004 0.745 1.3E-08 0.0213 3.9E-06 0.0032 mL WPetLa 0.1135 0.4554 0.0174 0.0050 1.6E-08 0.8494 2.7E-06 0.0320 mWw WPetLa 0.00102 0.1248 0.0008 0.1896 1.6E-10 0.0206 2.7E-09 2.7E-05 mWn WPetLa 0.01388 0.7245 0.0129 0.4268 0.7621 0.8994 0.0238 0.0102 mArcAp xAng KPetL 0.00179 0.3713 0.0003 0.1296 9.9E-08 0.0174 4.8E-06 3.7E-05 KPetClw 0.02018 0.3492 0.06 0.0504 0.00021 0.4591 0.00018 0.00066 L KPetClw 0.1873 0.5796 0.0203 1 0.00115 0.5871 0.1434 0.1833 W KPetLa 0.00155 0.5596 1.0E-05 0.7383 2.0E-07 0.0035 2.6E-06 0.00047 mL KPetLa 0.107 0.2029 0.00038 0.072 2.6E-07 0.1507 1.7E-05 0.00785 mWw KPetLa 0.00647 0.009 0.0005 0.4375 1.3E-09 0.1543 0.00014 0.00938 mWn ConFilL 0.02255 0.5281 0.00153 0.1205 6.0E-06 0.0633 2.7E-05 0.00043 ConFilW 0.1976 0.7309 0.00588 0.7421 0.01137 0.0975 0.04441 0.01312 ConFilP 0.1186 0.3953 0.3581 0.1752 0.00447 0.2108 0.1451 0.1835 cFus FreFilL 0.1719 0.3825 0.01272 0.3291 0.00021 0.2561 0.00119 0.01459 FreFilD 0.02079 0.1778 0.07189 0.7522 0.04942 0.8 0.00885 0.1738 OvL 0.00405 0.5827 0.00411 0.0196 0.00022 0.0145 0.00022 0.01618 OvW 0.0178 0.1299 0.07258 0.1038 0.01932 0.0426 0.03281 0.1142 StylL 0.7633 0.1054 1.9E-05 0.6133 2.2E-09 0.4443 1.6E-07 0.00039 StylD 5.8E-05 0.1778 0.00084 0.067 2.6E-08 0.208 3.2E-07 0.134 % Sig. 44.90% 6.25% 59.18% 28.57% 91.84% 36.73% 81.63% 55.10%

39 Table 10. P values resulting from a Student’s t-Test for T. mexicanum when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. sonoranensis StemL 0.04261 0.0307 0.0043 0.2599 0.1967 0.0154 0.001 0.1189 StemD 9.9E-05 0.0017 0.7117 0.8801 0.0246 0.1809 0.0057 0.0006 InNodL 0.0015 0.0068 0.0006 0.0009 0.3607 0.0045 0.0004 0.0049 StipL 0.00204 9.0E-05 0.0256 0.7754 0.0005 0.5093 4.7E-07 0.6715 StipW 0.01126 0.00332 0.3948 0.0119 0.0060 0.4748 0.01541 0.0004 StipApx 0.03019 0.00202 0.0096 0.5062 0.5501 0.5424 0.03594 0.7409 Ang PetL 0.2894 0.271 0.7478 0.2153 0.0087 0.9476 0.08825 0.6879 PetD 0.00012 0.00066 0.6057 0.4502 0.0042 0.5813 0.0006 0.6039 LfltBas 0.1175 0.3023 0.0079 2.1E-05 0.5666 0.044 0.7037 0.3847 Ang TermLfl 6.9E-06 0.00036 0.0016 0.7918 0.0019 0.7701 2.0E-05 0.0255 tL TermLfl 0.00031 0.00106 0.0007 6.3E-05 0.7874 0.0097 0.00012 0.025 tW PetlleL 2.4E-05 0.03006 0.1837 0.09107 0.1853 0.7425 3.78E-05 0.9352 PetlleD 0.00435 0.00022 0.1088 0.03749 0.0331 0.3606 0.1059 0.5863 InflL 3.1E-08 7.9E-05 0.3431 2.4E-06 0.109 0.0032 0.000957 1.3E-05 InflW 5.8E-09 9.4E-09 0.5561 3.7E-07 0.0312 0.0084 0.000224 0.00067 FlNo 3.6E-07 8.4E-07 0.1379 3.9E-05 0.3164 4.4E-05 0.07473 3.7E-05 Pedunc 0.00128 0.02067 0.3349 0.99 8.4E-09 0.3839 0.001105 0.4865 L Pedunc 2.6E-05 1.4E-07 0.915 0.00314 0.0002 0.3711 0.2547 0.01634 D PedicL 0.00012 0.0004 0.85 0.00819 0.3159 0.6601 0.01508 0.1636 PedicD 0.4186 0.00037 0.5171 0.4682 0.0306 0.1408 0.04902 0.2368 SepL 8.6E-06 2.5E-06 0.0229 0.01269 0.0366 0.3791 3.14E-06 0.1116 SepCir 0.597 0.00054 0.6885 3.9E-05 0.0331 0.3896 0.00115 0.00313 SepLob 2.2E-06 71E-06 0.0121 0.01376 0.1232 0.4462 1.22E-06 0.0377 L SepLob 0.3614 0.8019 0.3207 0.00357 0.0822 0.5489 0.1108 0.01465 W SepLob 0.00713 1.1E-07 0.8455 0.2553 0.9515 0.0629 0.06584 0.6895 ApxAng BPetL 4.3E-08 5.6E-05 0.4377 1.6E-09 0.0015 0.0011 2.99E-05 0.00022 BPetW 6.3E-07 3.1E-06 0.0203 1.5E-11 0.0492 0.0106 7.26E-05 4.0E-06 WPetL 1.0E-07 7.6E-06 0.519 9.7E-12 0.0060 0.0008 3.21E-05 7.3E-06 WPetCl 1.8E-05 5.9E-05 0.0601 1.2E-07 0.0981 3.2E-06 0.000304 0.4815 wL WPetCl 0.2013 0.03233 0.7976 0.01176 0.0103 0.5194 0.03233 0.3219 wW

40 Table 10 continued.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. sonoranensis WPetLa 3.7E-07 7.0E-07 0.7968 6.4E-12 0.0002 0.0009 3.88E-06 1.0E-06 mL WPetLa 1.0E-08 1.9E-06 0.9677 1.1E-12 0.0025 3.6E-06 2.72E-06 4.2E-06 mWw WPetLa 1.6E-08 0.00017 0.68 2.2E-11 0.004 0.0088 2.71E-09 0.00103 mWn WPetLa 0.6182 0.03503 0.1812 0.01249 0.0040 0.0106 0.0238 0.3864 mArcAp xAng KPetL 1.1E-13 9.6E-05 0.1078 3.4E-15 0.1116 0.0002 4.83E-06 0.00062 KPetCl 1.3E-07 2.1E-05 0.0624 9.2E-08 0.2812 0.0007 0.000178 0.1712 wL KPetCl 0.7013 0.3626 0.1434 0.04544 0.0059 0.0569 0.1434 0.9342 wW KPetLa 6.5E-10 0.00026 0.452 3.3E-14 0.0035 5.5E-06 2.59E-06 0.0004 mL KPetLa 2.8E-10 8.2E-05 1 2.1E-13 7.5E-05 0.0021 1.74E-05 3.3E-05 mWw KPetLa 0.06704 0.2578 0.0223 3.5E-05 5.9E-05 0.2218 0.000136 0.7632 mWn ConFilL 1.3E-10 1.1E-09 0.2846 2.1E-11 0.1898 0.0003 2.70E-05 0.00060 ConFil 0.1695 0.1091 0.1914 0.03138 0.4154 0.9009 0.04441 0.4959 W ConFilP 0.8074 0.5536 0.4358 0.801 0.0002 0.6821 0.1451 0.6802 cFus FreFilL 4.5E-09 2.9E-06 0.6652 3.0E-09 0.4315 0.0288 0.001188 0.00055 FreFilD 0.3913 0.0217 0.9096 0.00233 0.2038 0.0128 0.00885 0.05709 OvL 0.02842 0.0003 0.8884 0.1155 0.4834 0.8561 0.00022 0.3498 OvW 0.4283 0.2345 0.918 0.3378 0.7964 0.4645 0.03281 0.7644 StylL 7.2E-11 8.6E-10 0.0872 0.00693 0.0051 2.0E-05 1.56E-07 0.00175 StylD 0.00810 0.00323 0.5163 0.6544 0.0027 0.0611 3.16E-07 0.4008 % Sig. 75.51% 83.67% 22.45% 69.39% 57.14% 46.94% 81.63% 48.98%

41 Table 11. P values resulting from a Student’s t-Test for T. sonoranensis when compared with each different species, including the percentage of significant characters for each species. Statistically significant values are in red. α=0.05.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum StemL 0.9753 0.2573 0.3695 0.5771 0.0135 0.638 0.0617 0.1189 StemD 0.5802 0.1424 0.0660 0.0300 0.0002 0.1655 0.3742 0.0006 InNodL 0.2934 0.6629 0.0510 0.1341 0.0009 0.9807 0.0201 0.0049 StipL 0.1346 0.0088 0.2499 0.8977 0.0012 0.4082 0.0088 0.6715 StipW 0.2412 0.9424 0.0597 0.315 9.2E-07 0.0194 0.9159 0.0004 StipApx 0.1783 0.0048 0.0488 0.3982 0.921 0.7743 0.053 0.7409 Ang PetL 0.7374 0.6673 0.8788 0.594 0.00782 0.6763 0.4191 0.6879 PetD 0.01774 0.0094 0.889 0.3072 0.00516 0.8824 0.021 0.6039 LfltBasA 0.9262 0.8721 0.2691 0.0373 0.2148 0.4107 0.694 0.3847 ng TermLflt 0.00473 0.0189 0.2426 0.0599 4.1E-05 0.0191 0.0064 0.0255 L TermLflt 0.02183 0.0290 0.0410 0.0035 0.00866 0.7345 0.0071 0.025 W PetlleL 0.06102 0.0887 0.347 0.3288 0.2726 0.9353 0.0191 0.9352 PetlleD 0.01136 0.0011 0.0727 0.0369 0.09493 0.2054 0.0642 0.5863 InflL 0.5807 0.1441 0.0843 0.5832 1.1E-06 0.9725 0.8896 1.3E-05 InflW 0.03793 0.0017 0.0034 0.1181 2.8E-05 0.8828 0.0416 0.00067 FlNo 0.1304 0.0345 0.0819 0.4787 2.6E-06 0.3544 0.0558 3.7E-05 Pedunc 0.00506 0.0435 0.1828 0.6122 9.0E-09 0.6424 0.0047 0.4865 L Pedunc 0.02785 0.0002 0.1857 0.6792 4.5E-06 0.4026 0.946 0.01634 D PedicL 0.04551 0.0341 0.2666 0.3311 0.6539 0.2978 0.1297 0.1636 PedicD 0.2596 0.1177 0.2995 0.218 0.3233 0.3954 0.1494 0.2368 SepL 0.00338 0.0002 0.3786 0.1706 0.00141 0.955 0.0002 0.1116 SepCir 0.3382 0.3662 0.0428 0.0776 3.4E-05 0.1028 0.1531 0.00313 SepLob 0.00359 0.0004 0.4299 0.3648 0.00212 0.64 0.0003 0.0377 L SepLob 0.03699 0.0506 0.2731 0.5281 0.00015 0.0023 0.2308 0.01465 W SepLob 0.06981 0.0004 0.585 0.5579 0.6253 0.1013 0.1887 0.6895 ApxAng BPetL 0.03687 0.0028 0.2565 0.0003 3.0E-07 0.0614 0.0008 0.00022 BPetW 0.775 0.0066 0.0730 0.0001 2.4E-07 0.7684 0.0739 4.0E-06 WPetL 0.09716 0.0009 0.0842 1.7E-06 6.7E-08 0.0689 0.0014 7.3E-06 WPetCl 0.00012 0.0002 0.0969 4.5E-06 0.2391 2.9E-05 0.0006 0.4815 wL WPetCl 0.9698 0.689 0.481 0.3141 0.00351 0.9261 0.689 0.3219 wW

42 Table 11 continued.

T. amabile T. blanquitum T. cognatum T. goniocarpum T. hickeyi T. laciae T. lozani T. mexicanum WPetLa 0.8235 0.0017 0.0072 1.5E-05 2.2E-09 0.6653 0.0032 1.0E-06 mL WPetLa 0.3845 0.0094 0.0536 2.1E-05 4.7E-08 0.02547 0.0320 4.2E-06 mWw WPetLa 0.00233 0.0112 0.0268 1.3E-06 2.1E-06 0.325 2.7E-05 0.00103 mWn WPetLa 0.6642 0.0149 0.4021 0.00806 0.00302 0.00517 0.01023 0.3864 mArcAp xAng KPetL 0.00218 0.0007 0.3196 1.9E-05 5.6E-05 0.00601 3.7E-05 0.00062 KPetClw 6.3E-06 0.0002 0.1318 3.7E-06 0.9422 0.00215 0.00066 0.1712 L KPetClw 0.7041 0.3949 0.231 0.1194 0.02438 0.08196 0.1833 0.9342 W KPetLa 0.1499 0.008 0.0023 0.00029 8.5E-06 0.1217 0.00047 0.0004 mL KPetLa 0.03379 0.0877 0.0070 1.9E-05 1.6E-08 0.3297 0.00785 3.3E-05 mWw KPetLa 0.1231 0.2692 0.1071 0.00627 0.00986 0.2023 0.00938 0.7632 mWn ConFilL 0.00028 9.0E-06 0.3263 3.7E-05 5.4E-05 0.01032 0.00043 0.00060 ConFilW 0.02257 0.0366 0.46 0.00374 0.9668 0.7119 0.01312 0.4959 ConFilP 0.4879 0.6865 0.5717 0.8976 2.5E-05 0.9252 0.1835 0.6802 cFus FreFilL 0.0016 0.0002 0.2508 0.00041 0.00213 0.2405 0.01459 0.00055 FreFilD 0.13 0.7734 0.2115 0.07055 0.3623 0.1601 0.1738 0.05709 OvL 0.3869 0.0248 0.4273 0.5938 0.675 0.5186 0.01618 0.3498 OvW 0.723 0.4582 0.8404 0.5827 0.925 0.343 0.1142 0.7644 StylL 0.00034 3.7E-05 0.0329 0.1299 7.4E-06 0.00086 0.00039 0.00175 StylD 0.2515 0.1787 0.4478 0.5701 0.6994 0.2081 0.134 0.4008 % Sig. 42.86% 61.22% 18.37% 40.82% 71.43% 20.41% 55.10% 48.98%

Figure 15. Differences in terminal leaflet shapes and sizes between the members of this species complex. A: T. hickeyi, B: T. laciae, C: T. goniocarpum, D: T. mexicanum, E: T. cognatum, F: T. blanquitum, G: T. sonoranensis, H: T. lozani, and I: T. amabile.

Figure 16. Line diagram comparing the mean petal shapes and sizes of the T. amabile species complex.

45 Figure 16 (continued). Line diagram comparing the mean petal shapes and sizes of the T. amabile species complex.

46 Discussion This is the first study to take an in-depth look at the Trifolium amabile species complex. Unlike Zohary and Heller’s monograph (1984), which revised the entire genus of Trifolium, this study only focused only on the North American range of the complex. In doing such a thorough study, I was able to look at a large number of characters to best describe T. amabile and find previously undescribed species within the complex. Previously described species could be studied in detail, to reassure their status as a distinct species. Likewise, specimens that were outside the circumscribed boundaries of T. amabile sensu stricto could be identified as new species. This study concentrated on only the North American portion of the range; the South American portion of the range is being reexamined separately in order to fully understand the Trifolium amabile species complex. Another aspect that this study did not explore was molecular phylogenetics, and a thorough examination of commonly studied loci, particularly the nrDNA locus ITS, could prove helpful in fully understanding this species complex. There have been so many molecular studies involving the genus Trifolium and other related legumes (Ellison, et al., 2006; Steel and Wojciechowski, 2003; Wojciechowski, et al., 2004), it is a wonder that this group has not been studied in depth before. More work is also needed to look at disjunct populations, particularly in Costa Rica, where I had at one point recognized two different morphs, but lacked sufficient evidence to describe them as new species, and included them in T. amabile. T. hickeyi is an interesting species, which shows a strong clinal grade from the northern to the southern end of the range that warrants a closer investigation, especially including molecular work.

47 Taxonomic Treatment

Key to the North American species of the Trifolium amabile species complex 1. Peduncles >76 mm in length; ovaries glabrous..…………....5. Trifolium hickeyi 1. Peduncles <76 mm in length; ovaries pubescent, especially distally…………… …………………………………………………..………………………..……….……..2 2. Leaf texture thick and leathery; sepals serrate, becoming curled in fruit; petals yellow; 3 ovules per ovary and 3 seeds per fruit…...…… …………………….………………………………………………….6. T. laciae 2. Leaf texture thin and delicate; sepals entire, not curled in fruit; petals not yellow; 2 ovules per ovary and 2 seeds per fruit…………………… ……………………….……………………………………………………………3 3. Terminal leaflet lamina moderately to densely pubescent abaxially, at least distally; fruits inflated………………………… …………………..……………………………....……2. T. blanquitum 3. Terminal leaflet lamina glabrous abaxially; fruits laterally compressed…………………………………………………………….4 4. Wing petals not dilated distally, the margins parallel, <0.8 mm at the widest point……………..….4. T. goniocarpum 4. Wing petals dilated distally, the margins not parallel, >0.8 mm at the widest point……..………………..……………..5 5. Keel petal lamina length >2.8 mm; inflorescences >16 mm in diameter………………………………….6 5. Keel petal lamina length <2.8 mm; inflorescences <16 mm in diameter………………………………….7 6. Terminal leaflets <15 mm in length and <8 mm in width; internodes <20 mm in length; flowers pinkish purple to brownish purple; plants prostrate or creeping…….… ….……………………………….3. T. cognatum

48 6. Terminal leaflets >15 mm in length and >8 mm in width; internodes >20mm in length; flowers white or cream; plants prostrate to ascending …………….…….…… …………………………………8. T. mexicanum 7. Sepal lobes >3 mm in length; keel petal claws >1.7 mm in length; connate filaments >3.6 mm in length…………….9. T. sonoranensis 7. Sepal lobes <3 mm in length; keel petal claws <1.7mm in length; connate filaments <3.6 mm in length…………………………………..8 8. Stems densely pubescent; sepals <3 mm in length; banner petals <4.2 mm or length; wing petal claws <1.1 mm in length; keel petals <3.1 mm in length; keel petal claws <1.3 mm in length; ovaries <1.3 mm in length .……………………………….7. T. lozani 8. Stems glabrous to moderately pubescent; sepals >3 mm in length; banner petals >4.2 mm in length; wing petal claws >1.1 mm in length; keel petals >3.1 mm in length; keel petal claws >1.3 mm in length; ovaries >1.3 mm in length…………… ………………………..……1. T. amabile

49 1. Trifolium amabile Kunth, Figs. 17,18. Trifolium amabile Kunth, Nov. Gen. Sp. (quarto ed.) 6: 503, pl. 593. 1823[1824]; Lupinaster amabilis (Kunth) C. Presl, Symb. Bot. 1(3): 47.

1830.—TYPE: MÉXICO. Humboldt & Bonpland (holotype: B (destroyed),

photos GH!, NY!). NEOTYPE (to be designated): MÉXICO. Morelos: banks, Tres Marías, 19.05º, -99.24º, 2743m, Pringle, C. G. 10278 (neotype: GH!; isoneotypes: ARIZ!, ASU!, DAV!, MICH!, RSA!, VT!). Trifolium schiedeanum S. Wats., Proc. Amer. Acad. Arts 17: 339. 1882.—

TYPE: MÉXICO. Coahuila: Lirios, 45 miles east of Saltillo, 25.433, - 100.961, 1556m. Jul 1880. E. Palmer 201 (holotype: GH!; isotypes: NY!, VT!).

Plants perennial; roots large, woody, 2--10 mm in diameter; stems prostrate, ascending, 5--49+ cm in length, 0.5--2 mm in diameter, internodes glabrous--moderately pubescent, nodes sparsely--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular-- triangular, 4--12×1--7.5 mm, glabrous adaxially, glabrous--moderately pubescent abaxially, margins entire--weakly serrate, sparsely pubescent, apices acute; petioles 4--45×0.2--0.5 mm, sparsely--densely pubescent; petiolules 0.3--1.2×≤0.5 mm, sparsely--densely pubescent; leaflets obovate-- obcordate, glabrous adaxially, sparsely--densely pubescent along midribs abaxially, sparsely--moderately pubescent abaxially on outer halves of lateral leaflet, bases acute, margins weakly serrate--serrate, often with alternating short, straight teeth alternating with larger, antrorsely curved teeth, sparsely pubescent, apices rounded--emarginate, weakly mucronulate, terminal leaflets 3--15×2.5--2.5 mm; peduncles 9.5--50.5×<1 mm, sparsely--densely pubescent; inflorescences axillary, racemose-umbellate, 4.5--15.5×0.9--15 mm, flowers 3--25, each subtended by a linear triangular bract, in 1--3 whorls, terminal sterile projection sometimes present; pedicels 0.5--2.5×<0.5 mm, sparsely-- densely pubescent; calyx 2.5--5 mm long, tubes 1.9--3.1 mm in circumference, glabrous--densely pubescent, lobes narrowly triangular, 1.4--4×<1 mm,

50 glabrous--moderately pubescent abaxially, margins entire, sparsely--densely pubescent, apices attenuate; petals pink--purple; banner petals obovate, 3-- 7.5×2.2--5.6 mm; wing petals 2.8--7.2 mm long, asymmetrically clawed, claws 0.7--2.7×≤0.3 mm, laminae spatulate, 2--4.5 mm long, 0.8--2 mm wide at widest point, <1 mm wide at narrowest point, bases shortly auriculate, auricles acute--rounded, margins entire, apices rounded; keel petals 2.7--4.3 mm long, asymmetrically clawed, claws 0.9--1.6×<0.5 mm, laminae spatulate, 1.8--3.5 mm long, 0.9--1.4 mm wide at widest point, 0.3--1.2 mm wide at narrowest point, bases obtuse, margins entire, apices rounded; stamens diadelphous, connate filaments 2.2--3.6×0.9--1.5 mm, fused for 40--72% of their length, free filaments 1--2.7×≤0.2 mm; pistils sessile, ovaries 0.9--3.1×0.4--1.5 mm, sparsely--densely pubescent distally and occasionally along suture, styles 0.6-- 1.3×≤0.2 mm, glabrous, ovules 2; legumes laterally compressed, 3--5.4×1.7-- 2.8 mm; seeds 2, globular--mitten-shaped, brownish orange--reddish brown, occasionally dark-speckled, 1.2--1.9×1--1.5 mm.

Representative specimens

COSTA RICA. Alajuela-Heredia: along the cart-road from Vara Blanca (between Poás and Barva Volcanoes) to La Concordia, 10.1º, -84.1º, 1950m, 23 Jul 1923, W. R. Maxon 8440 & A. D. Harvey (US); Finca La Giralda, above Heredia, in very fertile pasture with kikuyu grass, CR-L25, 10.13º, -84.16º, 2000m, 1 Sep 1959, Semple s. n. (US). Cartago: Volcán Irazú, 9.9º, -83.8º, 2133m, 24 Jun 1874, O. Kuntze 2317 (NY); along the Río Reventado, north of Cartago, "trébol", moist meadow, prostrate, banner red-purple, other petals whitish, 9.83º, -83.9º, 1460m, 26 Feb 1926, P. C. Standley 49409 & J. Valerio (US). San José: Potreros between Guayabillos and Cabeza de Vaca, 9.967º, - 83.892º, 2350m, 04 Nov 1929, C. W. Dodge 4931 & W. S. Thomas (F); prairies et bords des chemins à San José, 9.9º, -84.1º, 1100m, Nov 1894, H. Pittier 1536 (US); San Francisco de Guadalupe, [Bin forage, mais and fen…--illegible], 9.93º, -84.09º, 1150m, Oct 1894, H. Pittier s. n. (US); vicinity of Santa María de

51 Dota, in potrero, prostrate or ascending, flowers purple, abundant, 9.92º, -84º, 1800m, 14 Dec 1925, P. C. Standley 42429 (F, US); chemin prés de la savane de San José, [prairies of San José], 9.9º, -84.1º, 1150m, Nov 1892, A. Tonduz 1536 & W. Barbey (P, US).

GUATEMALA. Chimaltenango: San Rafael, 14.878º, -90.817º, 1900m, 7 Jul 1936, B. B. Lewis 309 (F, P). El Quiché: Nebaj, in clearing, 12km north of the village on Cotzal Road, "Chunama" herb, flowers pinkish-white, 15.4194, - 91.1641, 2000m, 01 Jul 1964, E. Contreras 5152 (MEXU). Guatemala: La Cumbre, Socorro, 14.5º, -90.2º, 1860m, 8 Nov 1930, M. L. Rodriguez 2901 (P). Huehuetenango: Estación Ovino, Aldea San Nicolás, Chiantla, 15.4º, -91.5º, 3130m, 10 Oct 1975, C. J. Rushin s. n. (F). Jalapa: between Jalapa and Montaña Miramundo, prostrate, corolla lilac, in turf in llanos on top of ridge, 14.633º, -89.989º, 1500m, 07 Dec 1939, J. A. Steyermark 32898 (F). Quezaltenango: Volcán Santa Maria; near summit of mountain, in open somewhat weedy and disturbed meadow above above timberline, 14.75º, - 91.55º, 3770m, 12 Aug 1960, J. H. Beaman 4122 (NY). Sacatepéquez: San Lucas Sacatepéquez, hierba común, hojas digitadamente trifoliadas, inflorescencias en cabezuelas, 14.612º, -90.655º, 2088m, 9 Aug 2003, G. Ponce 032 & P. García (MO). San Marcos: Volcán Tacaná, vicinity of Sibinal, open, dry white clay slopes, 15.147º, -92.054º, 2589m, 18 Feb 1940, J. A. Steyermark 35952 (F). Sololá: Volcán Santa Clara, on summit, 14.65º, -91.28º, 3000m, 5 Jun 1942, J. A. Steyermark 47011 (F, NY). Totonicapán: open pine woodland, 11 mi SE of Totonicapán, along roadside, not common here, brs. prostrate, fIs. pinkish, 14.78º, -91.27º, 3048m, 9 Aug 1962, G. L. Webster 12863 et al. (DAV, F); ravines in Sierra Madre Mountains near Villa Las Cruces, about 8km southwest of Totonicapán, flowers pink, pendent, growing in ravine, 14.8º, -91.4º, 2800m, 13 Dec 1963, L. O. Williams 22952 et al. (F, NY, US).

MÉXICO. Chiapas: dorfanger von [illegible] Cina?a?lan, 16.196º, -93.136º, 808m, 4 Mar 1896, Seler 2092 (GH); aergahhang oberh, Huitzlan [illegible], 16.195722º, -93.135730º, 808m, 10 Mar 1896, Seler 2143 (GH). Chihuahua:

52 2km southeast of Cocheño on road to Mena Tarahumara, pine-oak woods, 28.308333º, -108.208333º, 2200m, 28 Aug 1986, P. S. Martin et al. s. n. (ARIZ). Coahuila: Lirios, 45 miles east of Saltillo, 25.433000º, -100.961493º, 1556m, Jul 1880, E. Palmer 201 (GH, NY, VT). Distrito Federal: Villa de Obregon, 19.5º, -99.13º, 2270m, 18 Jul 1935, G. L. Fisher 35236 (ARIZ, NY, P, US). Durango: Mpio. de El Salto, loc. 11km del Entronque de la Brecha a Sn. Miguel de Cruces con Carr. Durango-Mazatlán, hierba rastrera de flores rosadas, encase, veg. bosque de pino-encino, suelo amarillo forestall, 24.042º, - 104.654º, 1900m, 06 Jul 1982, P. Tenorio L. & C. Romero de T. 842 (MEXU). Guanajuato: Puerto Jondo, 26km de Tarimoro, entre los cerros La Bufa y La Pucha, ladera de cerro, potrero, planta herbácea rastrera de 5cm de alto, flor morada, muy abundante, 20.44º, -100.61º, 1900m, 14 Oct 1974, D. Flores 174 (XAL). Guerrero: La Hoya, 18.7º, -100.7º, 700m, Jul 1841, Liebmann 4932 (C). Hidalgo: San Vicente, 20.15º, -98.31º, 2377m, 16 Aug 1937, G. L. Fisher 3757 (US, NY). Jalisco: northern slopes of the Nevado de Colima, west of summit of the northern ridge, near junction of the old pack road to Zapotlán with Atenquique-Jazmín road, steep mountain sides in pine forest, abundant, prostrate, flowers white, 19.56º, -103.6º, 2200m, 15 Oct 1952, R. McVaugh 13526 (MICH, US). México: Paso de Cortes, slopes of Popocatepetl and Iztaccihuatl, 1.4 m W of Paso de Cortes near km. 21, pine forest with grass tussocks, standard pink, keel and wings whitish but tinged with pink, common along road, 19.120º, -98.766º, 3300m, 6 Jul 1972, M. F. Denton 1750 (MICH). Michoacán: 9 km northeast of Patzucuaro, collected within 10 meters of the road in disturbed area, 19.601º, -101.525º, 2144m, 9 July 1981, J. LaSalle 810709-18 et al. (ARIZ). Morelos: banks, Tres Marías, 19º, -99.2º, 2743m, 11 Aug 1906, C. G. Pringle 10278 (ARIZ, ASU, DAV, MICH, RSA, VT, GH); Cuernavaca, Moore lawn, strong taproot, 18.93º, -99.23º, 1500m, Jun 1953, G. B. Ross 36 (US). Nayarit: Acaponeta, in pine and oak forest at La Cinenga on ridge about 10 miles northwest of Mesa del Nayar, flowers white, 22.793º, - 105.193º, 1187m, 30 Jul 1970, D. H. Norris & D. J. Taranto 14554 (MICH).

53 Nuevo León: Sierra Madre Oriental, Mt. "El Infernillo", Pablillo, southeast of Galeana, rocky summit, prostrate herb, petals violet, 24.6º, -99.7º, 3000m, 29 Jun 1934, F. W. Pennell 17119 (US). Oaxaca: 110km by road south of Teotitlán on road to Oaxaca, oak forest, decumbent from taproot, flowers pink and white, 16.2º, -96.6º, 2180m, 11 Oct 1983, W. R. Anderson 13009 (MICH, NY). Puebla: Laguna de San Baltasar, 19º, -98.2º, 2135m, 7 Jul 1907, G. Arsène 1308 (US). San Luis Potosí: Catorce, Sierra de Catorce, Sierra Madre Oriental, grassy, andesitic slopes, SE of Catorce, herb, petals white becoming purple, 23.6º, - 100.87º, 2900m, 24 Jul 1934, F. W. Pennell 17530 (GH, NY, US). Tamaulipas: southern Tamaulipas, top of Sierra Madre Oriental (Sierra de Guatemala), rugged, wooded, karstic limestone plateau with sink holes and big boulders, ca 6km NW of Rancho del Cielo (Harrison Ranch of Martin & Harrell), (ca 4km NW of Julio) on road to La Joya de Salas, ca 12km NW of Gomez Farias, 108km SSW of Ciudad Victoria, very open, sunny secondary growth Pinus-Quercus woodlands (canopy 8m), with rugged white limestone outcrops, brught red soil (hence local name ("Tierra Colorado"), at fork of road to Las Canoas, common, flowers dirty pink and white, 23.2º, -99.283333º, 2000m, 12 Aug 1991, H. H. Iltis & B. Simon 30702 (MU). Tlaxcala: Nanacamilpa, agua Buena, bosque de pino, hierba rastrera, hojas trifoliadas, flores blancos, 19.491º, -98.537º, 2750m, 24 Oct 1986, A. Fontes C. s. n. (XAL). Veracruz: Lomogrande, Mt. Orizaba, flowers white in heads 3/8" diameter, touched pink, prostrate habit to 6" across, growing on open slopes and edges of cultivation, 19º, -97.2º, 3100m, 29 Apr 1938, E. K. Balls 4401 (MICH, US); antes de Acultzingo limite de los edos de Puebla y Veracruz, bosque de Quercus, ruderal junto al pavimento, abundante, herb-rastrera, perenne, flor rosa, 18.715º, -97.309º, 2300m, 28 Jul 1971, L. Nevling I. 2105 A. Gomez-Pompa (GH). Zacatecas: Sierra Madre Oriental, Mt. "El Temeroso", north of Aranzazu, andesitic gravelly slopes, herb, petals pale purple, 24.65º, -101.47º, 3000m, 19 Jul 1934, F. W. Pennell 17470 (GH, MICH, NY, US).

54 Distribution and Habitat Trifolium amabile is by far the most widely distributed species in the T. amabile species complex ranging throughout México, Guatemala, and Costa Rica (Fig. 19). This species is most often collected in sunny, moist, open areas of secondary pine and oak forests on gentle to steep slopes and ravines. It has also been collected in meadows and pastures, including cultivated cornfields. Other habitats include disturbed areas along riverbanks and roadsides. While these are the most common habitats, other locales include xeric hillsides, mountain and volcano summits, old lava flows, and areas of karst topography and limestone outcrops. T. amabile has been found growing on a variety of substrates. Loam to clay-loam soils are the most common along with gravelly and sandy soils which are brown, red, or black in color. Other types of soils include andisols and andesitic soils. It has been collected at elevations from under 100 m to 5000 m. In South America, T. amabile occurs in the countries of Colombia, Ecuador, Perú, Bolivia, and Argentina. It grows in the same variety of habitats as the North American specimens.

Comments Trifolium amabile is widespread throughout North America, and possibly continues on throughout South America as well. It is the only North American species to extend past Guatemala, as it also occurs in Costa Rica. This disjunct population growing in Costa Rica, at first look, appeared to have some differences from the T. amabile specimens from México and Guatemala, but these were never confirmed in my data. Nonetheless, this group warrants a closer look, and a possibly a molecular phylogenetic analysis. Trifolium amabile is most similar to T. blanquitum, but can be easily distinguished by lacking characters such as abaxial terminal leaf pubescence and inflated fruits. Instead, the abaxial surface of the terminal leaflet is glabrous and the fruits are laterally compressed in T. amabile. Another similar

55 species is T. lozani, however, T. amabile is less densely pubescent and has larger flowers than T. lozani.

Figure 17. Photograph of the Trifolium amabile type specimen. Originally housed at the Berlin herbarium (B), it is no longer extant. Photo from Harvard University (GH).

Figure 18. Trifolium amabile. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 58

Figure 19. Distribution of Trifolium amabile in North America. 59 2. Trifolium blanquitum T.K. Ahlquist et Vincent sp. nov., Fig. 20.

TYPE: MÉXICO. Nuevo León: Pablillo on Hwy 68, on grazed grassy slopes of road grade, abundant and short because of dryness and grazing, plants a little larger in damp soil patches, flowers white. 24.611º, -99.993º, 2000m, 20 Aug 1975, J. M. Gillett 17092 et A. Delgado (holotype: NY!).

Plants perennial; roots woody, 3--6.7 mm in diameter; stems prostrate or creeping, 5.5--36+ cm in length, 0.4--1.2 mm in diameter, internodes moderately--densely pubescent, nodes moderately--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular-- triangular, 4.3--8.8×1.5--3.2 mm, glabrous--sparsely pubescent adaxially, sparsely--moderately pubescent abaxially, margins entire--weakly serrate, moderately--densely pubescent, apices acute; petioles 3.5--21×0.2--0.5 mm, sparsely--moderately pubescent; petiolules <1×≤0.3 mm, sparsely--densely pubescent; leaflets obcordate--rhombic, glabrous adaxially, moderately-- densely pubescent along midribs abaxially, moderately--densely pubescent abaxially on distal portion of terminal leaflet and outer halves of lateral leaflets, bases acute, margins weakly serrate--serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely--moderately pubescent, apices emarginate--acute, weakly mucronulate, terminal leaflets 4.2--12.5×2.8--7.7 mm; peduncles 13--40×≤0.3 mm, moderately--densely pubescent; inflorescences axillary, racemose-umbellate, 5.2--12×8.4--11.5 mm, flowers 5--13, each subtended by a linear triangular bract, in 1--2 whorls; pedicels 1--2×≤0.1 mm, sparsely--moderately pubescent; calyx 2.5-- 3.4 mm long, tubes 2.1--2.7 mm in circumference, sparsely--densely pubescent, lobes narrowly triangular, 1.5--2.5×≤0.6 mm, sparsely--densely pubescent abaxially, margins entire, moderately--densely pubescent, apices attenuate; petals white--pinkish; banner petals elliptic--obovate, 3.3--4.9×2.1- -3 mm; wing petals 2.9--4 mm long, asymmetrically clawed, claws 0.8-- 1.2×≤0.2 mm, laminae spatulate, 2--2.8 mm long, 0.8--1.1 mm wide at widest point, ≤0.8 mm wide at narrowest point, bases shortly auriculate, auricles 60 acute, margins entire, apices rounded; keel petals 2.3--3.4 mm long, asymmetrically clawed, claws 1--1.4×<0.5 mm, laminae spatulate--elliptic, 1.3- -2.2 mm long, 0.9--1.2 mm wide at widest point, ≤1 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 2.2--2.7×0.9--1.4 mm, fused for 40--55% of their length, free filaments 0.9--1.6×≤0.1 mm; pistils sessile, ovaries 1--1.5×<1 mm, densely pubescent distally, styles ≤1×<0.2 mm, glabrous, ovules 2; legumes inflated, 2.3--3.8×1.8--2.4 mm; seeds 2, globular--mitten-shaped, brownish yellow-- light brown, occasionally dark-speckled, 1.2--2.1×0.9--1.5 mm.

Paratypes

MÉXICO. Hidalgo: Las Emes, 20km al norte de Ixmiquilpan, hacia la Pechuga, planta herbácea, apenas saliendo del suelo abundante, flores blancas, habitat: vegetación baja, inerme, 20.667º, -99.216º, 2200m, 18 Nov 1981, R. Hernández Magaña. 6666 (MEXU, XAL); Mpio. de Zacualtipan. Loc. Zoquizoquipan, 5km al norte de Zacualtipan, planta herbácea, rastrera, abundante, forrajera, flores lila, hab. bosque perturbado de encinos y pinos principalmente, suelo rojizo, profundo, 20.6484º, -98.7101º, 2200m, 21 Dec 1981, R. Hernández Magaña. 6927 (MEXU). Puebla: in grass beside railroad, 1km northwest of Huachinango, Puebla, prostrate, petals pale pink, 20.191916º, -98.028480º, 1450m, 18 Mar 1949, R. T. Clausen 7465 (MEXU). Veracruz: pastizal ariba de Santiago, Veracruz, pastizal, suelo volcanico, abundancia regular, herb-restrara, perenne, flor blanca-rosa, 19.7º, -97.24º, 1980m, 20 Jul 1971, L. I. Nevling 1859 & Gómez-Pompa (GH).

Distribution and Habitat Trifolium blanquitum is found primarily in the Sierra Madre Oriental mountain range in the Mexican states of Hidalgo, Nuevo León, Puebla, and Veracruz (Fig. 21). It has been collected in grassy pastures and roadsides from 1450 m to 2200 m in elevation.

Comments The epithet for this new species comes from the feminine diminutive form of the Spanish word for white, “blanca”. It is in reference to the small, white flowers of this species. Trifolium blanquitum is distinctive in possessing two unique character conditions: terminal leaflets which are moderately to densely pubescent and inflated fruits. Overall, T. blanquitum is one of the most pubescent members of this species complex.

Figure 20. Trifolium blanquitum. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 63

Figure 21. Distribution of Trifolium blanquitum, T. cognatum, and T. lozani. 64 3. Trifolium cognatum House, Fig. 22.

Trifolium cognatum House, Coult. Bot. Gaz. 41: 345. 1906. TYPE: MÉXICO.— Hidalgo: bare hills above Pachuca. 20.16º, -98.71º, 2743m, 30 Jul 1898, C. G. Pringle 6933 (holotype: US; isotypes: GH!, MICH!, NY!, P!, POM!, VT!).

Plants perennial; roots large, woody, 1.9--11.7 mm in diameter; stems prostrate--creeping, 11--25+ cm in length, 1--2 mm in diameter, internodes and nodes sparsely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular--triangular, 7.3--12×2.5--4.8 mm, glabrous adaxially, glabrous abaxially, margins entire, glabrous--sparsely pubescent, apices acute; petioles 9.5--32×<1 mm, glabrous--sparsely pubescent; petiolules 0.5--1×<0.5 mm, sparsely pubescent; leaflets obovate--rhombic, glabrous adaxially, sparsely pubescent along midribs abaxially, glabrous-- sparsely pubescent abaxially on outer halves of lateral leaflets, bases acute, margins serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely pubescent, apices rounded--acute, weakly mucronulate, terminal leaflets 9--15×5--8 mm; peduncles 27--70×≤1 mm, moderately pubescent; inflorescences axillary, racemose-umbellate, 9-- 20.5×16--20 mm, flowers 12--30, each subtended by a linear triangular bract, in 2--4 whorls; pedicels 1.8--3.2×<0.5 mm, sparsely pubescent; calyx 3.6--4.5 mm long, tubes 2.4--3.3 mm in circumference, glabrous--sparsely pubescent, lobes narrowly triangular, 2.5--3.3×≤0.7 mm, glabrous--sparsely pubescent abaxially, margins entire, sparsely--moderately pubescent, apices attenuate; petals pinkish purple--brownish purple; banner petals obovate--elliptic, 4.7-- 7.6×3.4--4.5 mm; wing petals 4.5--6.7 mm long, asymmetrically clawed, claws 0.9--1.7×≤0.4 mm, laminae spatulate, 3.6--5.1 mm long, 1.4--2.7 mm wide at widest point, ≤1.2 mm wide at narrowest point, bases shortly auriculate, auricles acute, margins entire, apices rounded; keel petals 3.7--5 mm long, asymmetrically clawed, claws 0.9--1.7×<0.5 mm, laminae spatulate, 2.8--3.3 mm long, 1.3--1.7 mm wide at widest point, 1--1.3 mm wide at narrowest 65 point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 2.9--4.5×1.3--1.6 mm, fused for 45---55% of their length, free filaments 1.6--3×≤0.2 mm; pistils sessile, ovaries 1.4--2.5×0.5--1.1 mm, glabrous--moderately pubescent distally, styles 1.7--2×≤0.2 mm, glabrous, ovules 2; legumes laterally compressed, 5.3--5.7×3--3.3 mm; seeds 2, mitten- shaped, burnt orange--olive brown, occasionally dark-speckled, 1.2--1.7×1.2-- 1.5 mm.

Representative specimens

MÉXICO. México: Cerro Ahumada, cerca del Rancho Nuevo, 4km al NE de Huehuetoca, ladera andesítica con vegetación de matorral xerófilo, planta rastrera, flores morado-rosadas, 19.872º, -99.219º, 2350m, 11 Aug 1971, Rzedowski 28414 (ARIZ). Hidalgo: El Bordo, 4km al N de Pachuca, encinar perturbado, planta rastrera, flores moradas, orilla de arroyo, 20.151389º, - 98.739444º, 2600m, 27 Aug 1967, Rzedowski 24287 (MICH); Cerro Ventoso, entre Pachuca y Real del Monte, bosque abierto de Juniperus y Quercus, planta rastrera, flores moradas, 20.134º, -98.698º, 2700m, 29 Aug 1965, Rzedowski 20602 (MICH); 4km al NE de Pachuca, sobre la carretera a Real del Monte, matorral xerófilo, planta herbácea perenne, flores moradas, 20.134º, -98.698º, 2650m, 14 Sep 1975, Rzedowski 33557 (MICH).

Distribution and Habitat Trifolium cognatum is found primarily in the Sierra Madre Oriental mountain range in the Mexican states of México and Hidalgo (Fig. 21). It has been collected in hilly regions with disturbed pine-oak forests between 2350 m and 2750 m in elevation.

Comments Trifolium cognatum is most similar to T. mexicanum, but T. cognatum differs in being more sparsely pubescent, smaller leaved, lacking a sterile

66 projection at the apex of the inflorescence, larger fruits, and most distinctly is purple flowered.

Figure 22. Trifolium cognatum. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary. 68 4. Trifolium goniocarpum Lojac., Fig. 23.

Trifolium goniocarpum Lojac., Nuov. Giorn. Bot. Ital. 15: 145. 1883.—TYPE:

MÉXICO. San Luis de Potosí, E. Palmer & C. C. Parry 134. (lectotype (to be designated): K; isolectotypes: GH!, NY!). Trifolium amabile Kunth var. longifoliolum Hemsl., Biol. Centr. Am. Bot. 1: 232. 1879.; Trifolium longifoliolum (Hemsl.) House (as “longifolium”),

Coult. Bot. Gaz. 41: 342. 1906.—TYPE: MÉXICO. San Luis Potosi, E. Palmer & C.C. Parry 134 (lectotype (to be designated): K; isolectotypes:

GH!, NY!); syntype: MÉXICO. Orizaba, Botteri 703 (K, NY!, P!).

Plants perennial; roots large, woody, 3.3--10 mm in diameter; stems creeping, 13--53+ cm in length, 0.7--3.1 mm in diameter, internodes sparsely-- moderately pubescent, nodes sparsely--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular, 4.8--19×1--4.5 mm, glabrous adaxially and abaxially, margins entire--weakly serrate, glabrous-- sparsely pubescent, apices acute; petioles 1.3--29×≤2 mm, sparsely-- moderately pubescent; petiolules 0.4--1.1×≤0.5 mm, sparsely--moderately pubescent; leaflets obcordate--elliptic, glabrous adaxially, sparsely-- moderately pubescent along midribs abaxially, glabrous--sparsely pubescent abaxially on outer halves of lateral leaflets, bases acute, margins weakly serrate--serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, glabrous--sparsely pubescent, apices rounded-- emarginated, weakly mucronulate, terminal leaflets 8.9--28.5×2.6--9 mm; peduncles 30--77×<1 mm, sparsely--densely pubescent; inflorescences axillary, racemose-umbellate, 6--14×8--16.5 mm, flowers 11--23, each subtended by a linear triangular bract, in 1--3 whorls; pedicels 0.9--3×<0.5 mm, sparsely--densely pubescent; calyx 2.5--5.2 mm long, tubes 1.7--3.4 mm in circumference, glabrous--sparsely pubescent, lobes narrowly triangular-- linear triangular, 1.8--4.4×≤0.5 mm, sparsely--moderately pubescent abaxially, margins entire, sparsely--moderately pubescent, apices acuminate; petals

69 white; banner petals obovate, 3.6--5.4×2--3 mm; wing petals 3--4.5 mm long, asymmetrically clawed, claws 1--1.4×≤0.2 mm, laminae spatulate, 1.9--3 mm long, 0.5--0.8 mm wide at widest point, ≤0.7 mm wide at narrowest point, bases shortly auriculate--truncate, if auriculate, auricles acute, margins entire, apices rounded; keel petals 2.8--3.5 mm long, asymmetrically clawed, claws 1.1--1.6×<0.5 mm, laminae elliptic--spatulate, 1.5--2 mm long, 0.7--1 mm wide at widest point, 0.6--0.8 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 2.4--3×0.8-- 1.5 mm, fused for 44---60% of their length, free filaments 1.5--1.9×<0.2 mm; pistils sessile, ovaries 1.2--3.6×0.4--1.3 mm, sparsely--densely pubescent distally and occasionally along suture, styles 0.3--1.3×≤0.2 mm, glabrous, ovules 2; legumes laterally compressed, 4.2--5.8×2.3--3 mm; seeds 2, globular--mitten-shaped, brownish orange--dark brown, occasionally dark- speckled, 1.5--2×1.1--1.6 mm.

Representative specimens

MÉXICO. Aguascalientes: shrub-covered, nearly treeless mountain sides ca 20km east of Rincón de Romos, road to Asientos, between Cerro Altamira and Cerro San Juan, limestone areas with Ceanothus, Ephedra, Mimosa, areas of rhyolitic rock with Opuntia, Mimosa, Eupatorium, Erigonum, in flat land 3km east of Asientos; seepage area near a small watercourse, abundant, 22.2303º, - 102.0568º, 2100m, 4 Sep 1967, R. McVaugh 23781 (MICH). Chihuahua: plains near Guerrero, 28.56º, -107.49º, 2000m, 8 Sep 1887, C. G. Pringle 1208 (GH, MICH, NY, RSA, US, VT). Distrito Federal: Valley of Mexico, 19.25º, -98.99º, 2316m, 28 Sep 1909, C. G. Pringle 15645 (GH, MICH, US, VT). Durango: city of Durango, 24.02º, -104.6º, 1900m, 1 Aug 1898, E. W. Nelson 4586 (GH, US). Guanajuato: San Luis de la Paz, Cerro El Quijay, camino hacia Pozos, matorral arbustivo, ladera de cerro, planta herbácea rastrera, flores blancas, escasa, 21.25º, -100.507º, 1900m, 31 Jul 1991, E. Ventura 9380 & E. López (XAL). Hidalgo: Tula, 20.052259º, -99.344429º, 2048m, Jul 1905, J. N. Rose et al.

70 8307 (GH). Jalisco: Guadalajara, 20.67º, -103.34º, 1550m, Jul 1886, E. Palmer 236 (GH, MICH, NY, P). México: Lecheria, railroad station, 19.611º, - 99.186º, 1917m, 2 Jul 1904, C. G. Pringle 13264 (ARIZ, ASU, GH, MICH, VT). Michoacán: Morelia, Loma del Zapota, 16.7º, -101.2º, 1950m, Sep 1911, G. Arsène 6011 (GH, US). Morelos: Cuernavaca, 18.934º, -99.232º, 1700m, 13 Aug 1906, C. G. Pringle 10282 (ARIZ, ASU, DAV, GH, MICH, RSA, VT, XAL). Nayarit: collected in the Sierra Madre, near Santa Teresa, Territorio de Tepic, 22.5º, -104.7º, 2100m, 9 Aug 1897, J. N. Rose s. n. (US). Oaxaca: Miahuatlán, San Juan Mixtepec, edge. PMZ #103, trifoliate leaves, 16.301667º, - 96.3000278º, 1897m, 26 Oct 1996, E. Hunn OAX-416 (MO, XAL). Puebla- Veracruz: Carretera 140, ruderal, secundaria a los lados de la Carretera, abundante, hierba, perenne, arrosetada rastrera, flor blanca, 19.5º, -97.4º, 2250m, 12 Jul 1971, L. I. Nevling 1611 & F. Chiang (GH). Sonora: 6.6km west of Yécora on Mex. 16, pine-oak forest, annual, 28.361667º, -108.985º, 1760m, 5 Sep 1996, A. Burquez M. 96-844 et al. (NY). Tlaxcala: Jardín Botánico de Tizatlán, bosque secundario de Alnus, ruderal, arenoso, abundante, hierba, 7cm de tamaño, annual, fruto verde, flor blanca-rosada, hojas con manchas en forma de "V", 19.3308º, -98.2168º, 2300m, 18 Aug 1988, B. Acosta P. & P. Rizo B. 2267 (XAL). Zacatecas: 9 miles northwest of Sombrerete on eastern bajada of Sierra Papanton, oak-juniper grassland, clay loams from igneus or sedimentary rocks, 23.657º, -103.663º, 2400m, 26 Sep 1948, H. S. Gentry 8487 (ARIZ, MICH, POM).

Distribution and Habitat Trifolium goniocarpum is fairly widespread throughout México, growing in the states of Aguascalientes, Chihuahua, Distrito Federal, Durango, Guanajuato, Hidalgo, Jalisco, México, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, San Luis Potosí, Sonora, Tlaxcala, Veracruz, and Zacatecas (Fig. 24). It has been collected in a variety of habitats including secondary pine-oak forests, along roadsides and other disturbed areas, hillsides, along streams or damp

71 depressions in grasslands, and riparian areas. In these locations the soil varies from very sandy and rocky to moist clay loams. Trifolium goniocarpum is found across a broad range in elevation from 1100 m to 4500 m. Comments The most distinguishing characters of Trifolium goniocarpum are the narrow wing and keel petals in the flowers and the long and narrow leaflets. T. goniocarpum is most similar to T. laciae, however, they differ in aforementioned characters in addition to sepal circumference and lobe width. T. goniocarpum also has relatively small inflorescences, which sets it apart from most other members of this species complex.

Figure 23. Trifolium goniocarpum. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 73

Figure 24. Distribution of Trifolium goniocarpum. 74 5. Trifolium hickeyi T.K. Ahlquist et Vincent sp. nov., Fig. 25.

TYPE: MÉXICO. Michoacán: Morelia, Rincón prise d'eau. 16.698º, - 101.187º, 1950m, 14 Apr 1910, G. Arsène 5483 (holotype: ILL!, isotypes: GH!, NY!).

Plants perennial; roots large, woody, 3.3--11.8 mm in diameter; stems prostrate--decumbent, 12--78+ cm in length, 1--4.5 mm in diameter, internodes and nodes glabrous--moderately pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular--triangular, 19.5-- 37.5×2--8.5 mm, glabrous adaxially and abaxially, margins entire--weakly serrate, glabrous--sparsely pubescent, apices acute; petioles 5.5--107×≤1.5 mm, glabrous--sparsely pubescent; petiolules 0.5--4×≤1.2 mm, glabrous-- moderately pubescent; leaflets lanceolate--elliptic, occasionally rhombic, glabrous adaxially, glabrous--moderately pubescent along midribs abaxially, glabrous--sparsely pubescent abaxially on outer halves of lateral leaflets, bases acute, margins serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, glabrous--sparsely pubescent, apices rounded--acute, weakly mucronulate, terminal leaflets 13.5--46×5.5--18.5 mm; peduncles 76-- 211×≤2 mm, glabrous--moderately pubescent; inflorescences axillary, racemose-umbellate, 9.5--26×16--26 mm, flowers 17--50, each subtended by a linear triangular bract, in 2--5 whorls; pedicels 0.7--3.6×<0.5 mm, glabrous-- sparsely pubescent; calyx 3.9--7.4 mm long, tubes 2.6--4.1 mm in circumference, glabrous--sparsely pubescent, lobes narrowly triangular, 3-- 6×<1 mm, glabrous--sparsely pubescent abaxially, margins entire, sparsely-- moderately pubescent, apices acuminate; petals white--yellowish white; banner petals obovate, 6--9.6×3.8--6.3 mm; wing petals 5.5--8.2 mm long, asymmetrically clawed, claws 1.1--2.1×≤0.5 mm, laminae spatulate, 4.2--6.2 mm long, 1.7--2.9 mm wide at widest point, 0.9--1.4 mm wide at narrowest point, bases shortly auriculate--truncate, auricles acute, margins entire, apices rounded; keel petals 4.5--6.6 mm long, asymmetrically clawed, claws 1.4-- 2.4×≤0.6 mm, laminae spatulate, 3--4.2 mm long, 1.5--2 mm wide at widest 75 point, 0.9--1.5 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 3.7--5.6×1.1--1.6 mm, fused for 50---70% of their length, free filaments 2.1--4.4×≤0.2 mm; pistils sessile, ovaries 1.3--3.5×0.4--1.6 mm, glabrous, styles 1.7--2.9×≤0.3 mm, glabrous, ovules 2; legumes laterally compressed, 3.3--5.2×1.9--2.7 mm; seeds 2, mitten-shaped, light brown--dark brown, occasionally dark-speckled, 1.7--2.2×1.2--1.6 mm.

Paratypes

GUATEMALA. Chimaltenango: Chichavac, flowers greenish white, 14.8º, - 90.98º, 2550m, Nov 1930, A. F. Skutch 88 (MICH, US). Huehuetenango: Vicinity of San Mateo Ixtatán, Sierra de los Cuchumatanes, flowers dull whitish, 15.8º, -91.4º, 3100m, 31 Jul 1942, J. A. Steyermark 49895 (F, NY, US). San Marcos: San Pedro Sacatepéquez, Aldea Chamác, hierba prostrada, muy abundante, 14.969º, -91.756º, 2200m, 26 Aug 1992, M. Véliz 92.2418 (MO).

MÉXICO. Chiapas: Laguna Jusnajav, 16 km. northeast of Comitán, 16.380º, -92.038º, 1758m, 29 Aug 1945, E. J. Alexander 1265 (MICH, NY). Guanajuato: Municipio de Acámbaro, La Cieneguilla, zoan de cultivo, terreno plano, hierba abundante, 21.075º, -100.175º, 1775m, 18 Aug 1985, A. Rubio 69 (XAL). Jalisco: Sierra del Tigre, 2 miles northeast of Mazamitla, gently sloping wet meadow-like areas in pine forest zone, abundant, mostly past flower, flowers cream color, 19.883º, -102.983º, 2100m, 23 Sep 1952, R. McVaugh 13186 (MICH). México: Mpio. de Tepotzoltan, cerca de la Presa de Concepción, flor blanquecina; orilla de arroyo, 19.717º, -99.223º, 2400m, 7 Aug 1966, Rzedowski 22885 (ASU, MICH). Michoacán: Morelia, 19.7º, -101.2º, 1950m, 1 Aug 1912, G. Arsène 8323 (B, NY, P); Mpio. de Villa Morelos, alrededores de El Fresno, hab. orilla de una acequia, planta herbácea perenne, flores de color crema, 20.026º, -102.882º, 2250m, 30 Aug 1987, Rzedowski 44278 (MICH). Oaxaca: Ixlán de Juárez, east-facing slopes, mountains along

76 Route 175, 12 km north of Ixlán de Juárez on road to Valle Nacional, in forest of pine and deciduous trees, abundant, open sun, gravelly soil, flowers white, 17.439º, -96.474º, 2500m, 26 Jul 1956, R. M. King 2043 (MICH); Mpio. Santiago Juxtlahuaca, loc. Senda hacia la torre de microondas de El Manzanal, entrada por Santa Rosa-San Miguel Cuevas, bosque de Pinus-Quercus, Suelo café rocoso, hierba de 60 a 70cm de alto, flor blanca, planta de regular abundancia, 17.225333º, -98.054611º, 2225m, 12 Sep 1996, J. I. Calzada 21326 (MO); Tlacolula, a 16 km. al N-NE de Díaz Ordaz, o sea a 6 km. al S de Cuajimaloya, Distr. de Tlacolula, herbacea perenne, arrocetada, flores blancas con rosa, veg: alterada de encinar, 17.070º, -96.4º, 2670m, 7 Aug 1977, M. Sousa 7822 et al. (MICH, NY); 110km by road south of Teotitlán on road to Oaxaca, oak forest, decumbent from taproot, flowers yellowish green, 16.2º, - 96.6º, 2180m, 11 Oct 1983, W. R. Anderson 13019 (MICH); Cumbre de Ixtepec, 16.57º, -95.1º, 60m, Sep 1942, Liebmann 4928 (C!, US!). Puebla-Oaxaca: San Luis Tultitanapa, Cerro Verde, near Oaxaca, 18.1º, -97.3º, 1400m, Jul 1908, C. A. Purpus 3228 (GH, NY, US). Puebla-Veracruz: Orizaba, 19º, -97.2º, 4500m, 1856, Botteri 704 (NY, P).

Distribution and Habitat Trifolium hickeyi is found primarily in southern México, excluding the Yucatan Peninsula. It is known from Chiapas, Jalisco, México, Michoacán, Oaxaca, Puebla, and Veracruz, and in Guatemala (Fig. 26). It has been collected in oak-pine forests on grassy slopes, with gravelly to rocky soil between 1400 m and 4500 m in elevation.

Comments Trifolium hickeyi is identified by a number of characters well outside of the range of the other species, including both vegetative and reproductive characters. One such character of note is the very long peduncles, measuring over 76 mm in length and glabrous ovaries. Within T. hickeyi there is a clinal

77 grade from north to south evident in leaflet morphology. The specimens at the northern end of the range had rhombic leaflets remarkable similar to those of T. mexicanum, however, all other characters are consistent with the southern specimens of T. hickeyi. Further research is definitely needed here to assess this difference in morphology. T. hickeyi is named in honor of R. James Hickey, professor and chair of the Department of Botany at Miami University.

Figure 25. Trifolium hickeyi. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 79

Figure 26. Distribution of Trifolium hickeyi and T. mexicanum. 80 6. Trifolium laciae T.K. Ahlquist et Vincent sp. nov., Fig. 27.

TYPE: MÉXICO. Chihuahua: Mesa, west of Hop Valley, Sierra Madre Mountains. 27.2688º, -106.793º, 2133m, 17 Sep 1903, M. E. Jones 7501 (holotype: POM! 27959; isotypes: MICH!, POM! 27942 & 27943, US!).

Plants perennial; roots large, woody, 4.3--7.9 mm in diameter; stems 15--30+ cm in length, 1.1--1.9 mm in diameter, internodes sparsely-- moderately pubescent, nodes moderately--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular--triangular, 10-- 16.5×2.7--5 mm, glabrous adaxially and abaxially, margins entire--weakly serrate, sparsely--moderately pubescent, apices acute; petioles 6--19×<1 mm, moderately--densely pubescent; petiolules ≤1×≤0.5 mm, sparsely--densely pubescent; leaflets obovate--rhombic, glabrous adaxially, sparsely--moderately pubescent along midribs abaxially, sparsely--moderately pubescent abaxially on outer halves of lateral leaflets, bases acute, margins weakly serrate--serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely--moderately pubescent, apices rounded--acute, weakly mucronulate, terminal leaflets 14--21.5×7.5--9 mm; peduncles 13--50×<1 mm, moderately-- densely pubescent; inflorescences axillary, racemose-umbellate, 6.5-- 15.5×11.5--18 mm, flowers 10--20, each subtended by a linear triangular bract, in 1--3 whorls, terminal sterile projection present; pedicels 1.6-- 4.5×<0.5 mm, sparsely--moderately pubescent; calyx 3.3--5.7 mm long, tubes 2.5--3.5 mm in circumference, sparsely--densely pubescent, lobes narrowly triangular, 2.6--4.6×≤0.6 mm, sparsely--densely pubescent abaxially, margins usually serrate, sparsely--densely pubescent, apices acuminate; petals yellow; banner petals obovate, 3.6--5.7×2.4--4.2 mm; wing petals 3.4--5 mm long, asymmetrically clawed, claws 0.9--1.2×≤0.3 mm, laminae spatulate, 2.6--3.7 mm long, 0.8--1.2 mm wide at widest point, <1 mm wide at narrowest point, bases shortly auriculate, auricles acute, margins entire, apices rounded; keel petals 2.8--3.8 mm long, asymmetrically clawed, claws 0.8--1.4×<0.5 mm, laminae spatulate, 2--2.5 mm long, 0.9--1.3 mm wide at widest point, 0.6--1 81 mm wide at narrowest point, bases truncate, margins entire, apices rounded-- obtuse; stamens diadelphous, connate filaments 2.2--3.4×1--1.6 mm, fused for 47---55% of their length, free filaments 1.2--2.3×≤0.1 mm; pistils sessile, ovaries 1.4--2.7×0.5--1.3 mm, sparsely--densely pubescent distally and occasionally along suture, styles 0.6--1.1×≤0.2 mm, glabrous, ovules 3; legumes laterally compressed, 5--5.3×2.6--2.9 mm; seeds 3, globular--mitten- shaped, brownish orange--dark brown, occasionally dark-speckled, 1.4-- 1.8×1.2--1.5 mm.

Paratypes

MÉXICO. Chihuahua: Collected near Colonia García in the Sierra Madres, 29.97º, -108.34º, 2286m, 27 Jul 1899, C. H. T. Townsend 177 & C. M. Barber (GH, ILL, NY, P, POM, US, VT); just northeast of La Zaragosa, about 19.4 miles southwest of San Buenaventura, cornfields at base of hill with Mexican pine- oak woodland, perennial from rootstock, flat area near cornfield, under Pinus emgelmannii, 29.620º, -107.899º, 2500m, 22 Oct 1984, T. R. Van Devender 84- 523 et al. (ARIZ); Madera, Laguna de Babícora, Arroyo El Jara, bosque de pino- encino, herb, floras amarillas, frecuente, 29.3º, -107.8º, 2300m, 10 Sep 1994, G. Quintana 3602 & E. Estrada (NY); Mesa, west of Hop Valley, Sierra Madre Mountains, 27.2688º, -106.793º, 2133m, 17 Sep 1903, M. E. Jones 7501A (POM). Durango: Otinapa, 24.05º, -105º, 2367m, Jul 1906, E. Palmer 417 (GH, NY);

Distribution and Habitat Trifolium laciae is found in the Sierra Madre Occidental mountain range, primarily in the Mexican states of Chihuahua and Durango (Fig. 28). It has been collected in pine-oak woodlands and occasionally near lakes between 1750 m and 2400 m in elevation.

Comments Trifolium laciae has several characters that distinguish it from other members of this species complex, most of which are floral and reproductive characters. While other species may have a sterile projection of the inflorescence, it is a prominent character in T. laciae. The serrate sepals, yellow petals, and three ovules per ovary and three seeds per fruit, are not shared with any other members of this species complex. Other observations worth noting not found in any other species in this complex are that the leaves seem to be thicker and have a more leathery texture, and the remnant sepals become curled in fruit. T. laciae is named in honor of my sister, Laci Ahlquist.

83 Figure 27. Trifolium laciae. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 84

Figure 28. Distribution of Trifolium laciae and T. sonoranensis. 85 7. Trifolium lozani House, Fig. 29.

Trifolium lozani House, Bot. Gaz. 41: 342. 1906.—TYPE: MÉXICO. Distrito Federal: Eslava, Valley of México. 19.25º, -98.99º, 2316m, 15 Jun 1901, C. G. Pringle 9512 (holotype: US; isotypes; GH!, VT!).

Plants perennial; roots large, woody, 1.2--5.1 mm in diameter; stems prostrate, 6--26+ cm in length, 0.4--1.3 mm in diameter, internodes and nodes moderately--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules elliptic, 5--7.4×1.5--3.5 mm, glabrous adaxially, moderately pubescent abaxially, margins entire--weakly serrate, sparsely--moderately pubescent, apices acute; petioles 7--14×≤0.5 mm, moderately--densely pubescent;

petiolules ≤0.6×≤0.5 mm, moderately--densely pubescent; leaflets obovate-- elliptic, glabrous adaxially, sparsely--moderately pubescent along midribs abaxially, sparsely--moderately pubescent abaxially on outer halves of lateral leaflets, bases acute, margins weakly serrate--serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely--moderately pubescent, apices rounded, weakly mucronulate, terminal leaflets 6.1--11.2×4- -7.9 mm; peduncles 15--31×<1 mm, moderately--densely pubescent; inflorescences axillary, racemose-umbellate, 4.5--12×7--13 mm, flowers 13-- 28, each subtended by a linear triangular bract, in 1--3 whorls, terminal sterile projection sometimes present; pedicels 0.8--2.4×≤0.2 mm, sparsely--densely pubescent; calyx 2.2--3.3 mm long, tubes 1.9--2.6 mm in circumference, sparsely--densely pubescent, lobes narrowly triangular, 1.7--2.4×≤0.5 mm, moderately--densely pubescent abaxially, margins entire, moderately--densely pubescent, apices attenuate; petals light pink--pink; banner petals obovate, 2.5--4.2×2.1--3.2 mm; wing petals 2.3--3.8 mm long, asymmetrically clawed, claws 0.5--1.1×≤0.2 mm, laminae spatulate, 1.8--2.7 mm long, 0.8--1.2 mm wide at widest point, ≤0.6 mm wide at narrowest point, bases shortly auriculate--truncate, auricles acute, margins entire, apices rounded; keel petals 2.2--3.1 mm long, asymmetrically clawed, claws 0.8--1.3×≤0.3 mm, laminae spatulate, 1.4--2 mm long, 0.8--1.1 mm wide at widest point, 0.7--0.8 86 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 1.6--2.7×0.8--1.2 mm, fused for 37-- 55% of their length, free filaments 0.7--1.9×<0.2 mm; pistils sessile, ovaries 0.7--1.3×0.3--0.6 mm, sparsely--densely pubescent distally, styles 0.8-- 1.2×<0.2 mm, glabrous, ovules 2; legumes laterally compressed, 3.4×2.1--2.8 mm; seeds 2, mitten-shaped, brown, 1.3×1.1 mm.

Representative specimens

MÉXICO. Guanajuato: 5 miles east of San Miguel Allende, 20.916565º, - 100.667001º, 2032m, 18 Aug 1947, L. A. Kenoyer 2425 (A). Hidalgo. Pachuca, open meadow and rocks in fir forest above Pueblo Neuvo and below Parque Nacional El Chico on road from Real del Monte to El Chico, prostrate herb, flowers pinkish, 20.163º, -98.710º, 3000m, 6 Jul 1948, H. E. Moore Jr. 3693 & C. E. Wood Jr. (A). México: at pine woods, open field, Llano Grande, 19.219º, - 98.817º, 2800m, 09 Jul 1950, E. Matuda 19222 (NY). Puebla: Boca del Monte, moist meadows, 18.4º, -97.3º, 1200m, Aug 1908, C. A. Purpus 3053 (GH). Veracruz: Mpio. de Perote, northwestern slopes of Cofre de Perote, 6.5km (by road) SE of town of Perote, along road to television towers on summit of Cofre de Perote, bosque de pinos, forest of Pinus teocote, P. pseudostrobus, Quercus crassifolia, Quercus spp., steep slopes and now dry canyon, prostrate, flowers light pink, 19.4871º, -97.1503º, 2700m, 08 Jul 1980, B. F. Hansen 7682 & M. Nee (POM, NY); Las Vigas de Ramírez, San Juan del Monte, Predio San Juan del Monte, bosque de pino, pradera inundable, herbácea de 5cm, flor rosada, 19.633333º, -97.1º, 2450m, 05 Aug 1985, C. A. Marín 89 (XAL).

Distribution and Habitat Trifolium lozani is found in the Sierra Madre Oriental mountain range and the Valley of Mexico, in the Mexican states of Distrito Federal, Guanajuato, Hidalgo, México, Puebla, San Luis Potosí, and Veracruz (Fig. 21). It has been

87 collected in open meadows and pine forests between 1200 m and 3250 m in elevation. Comments Along with Trifolium blanquitum, T. lozani is one of the most densely pubescent members of this complex. T. lozani differs from T. blanquitum most notably with laterally compressed fruits. T. lozani also has denser inflorescences than T. blanquitum. T. lozani is also somewhat similar to T. amabile, but generally has smaller characters than T. amabile including smaller sepals, ovaries, and banner, wing, and keel petals.

Figure 29. Trifolium lozani. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 89 8. Trifolium mexicanum Hemsl., Fig. 30. Trifolium mexicanum Hemsl., Biol. Centr. Am. Bot. 1: 233. 1879. Trifolium amabile Kunth var. mexicanum (Hemsl.) Heller & Zoh., Gen. Trifolium

128. 1984.—TYPE: MÉXICO. San Luis de Potosí: 1887. E. Palmer & C. C: Parry 137 (holotype: K; isotypes: GH!, NY!, P!).

Trifolium potosanum Lojac., Nouv. Giorn. Bot. Ital. 15: 144. 1883.—TYPE:

MÉXICO. San Luis de Potosí: 1887. E. Palmer & C. C: Parry 137 (holotype: K; isotypes: GH!, NY!, P!).

Trifolium nelsoni House, Bot. Gaz. 41: 344. 1906.—TYPE: MÉXICO. Oaxaca: vicinity of La Parada. 19 Aug 1894. E. W. Nelson 1016 (holotype: US).

Plants perennial; roots large, woody, 3.5--9.2 mm in diameter; stems prostrate--ascending, 14--70+ cm in length, 1--2.5 mm in diameter, internodes sparsely--moderately pubescent, nodes moderately--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular-- triangular, 8.4--17×2.5--8 mm, glabrous adaxially, glabrous--basally moderately pubescent abaxially, margins entire--serrate, sparsely--moderately pubescent, apices acute; petioles 5--60.5×≤1 mm, sparsely--densely pubescent; petiolules 0.6--1.2×<1 mm, moderately--densely pubescent; leaflets rhombic--obovate, glabrous adaxially, sparsely--moderately pubescent along midribs abaxially, sparsely pubescent abaxially on outer halves of lateral leaflets, bases acute, margins serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely pubescent, apices rounded--acute, weakly mucronulate, terminal leaflets 9.7--26.5×5.9--18 mm; peduncles 28--78.5×≤1 mm, moderately--densely pubescent; inflorescences axillary, racemose-umbellate, 10--24×15--24 mm, flowers 20--50, each subtended by a linear triangular bract, in 2--5 whorls, terminal sterile projection sometimes present; pedicels 1--4.2×<0.5 mm, sparsely--moderately pubescent; calyx 3.4--6.1 mm long, tubes 2.3--4 mm in circumference, sparsely--moderately pubescent, lobes narrowly triangular--linear triangular,

90 2.3--4.8×≤0.7 mm, sparsely--moderately pubescent abaxially, margins entire, sparsely--moderately pubescent, apices attenuate; petals white--cream; banner petals obovate, 5.7--8.5×3.5--5.5 mm; wing petals 5.1--7.3 mm long, asymmetrically clawed, claws 1.4--2.2×<0.5 mm, laminae spatulate, 3.7--5.2 mm long, 1.3--2.3 mm wide at widest point, ≤1.2 mm wide at narrowest point, bases shortly auriculate, auricles acute, margins entire, apices rounded; keel petals 4.4--5.9 mm long, asymmetrically clawed, claws 1.6--2.4×<0.5 mm, laminae spatulate, 2.8--3.9 mm long, 1.2--1.7 mm wide at widest point, ≤1.3 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 3.4--5.1×0.4--1.6 mm, fused for 45--- 61% of their length, free filaments 2.1--3.5×≤0.2 mm; pistils sessile, ovaries 1.5--4.5×0.5--2.1 mm, sparsely--densely pubescent distally and occasionally along suture, styles 1.4--2.5×≤0.2 mm, glabrous, ovules 2; legumes laterally compressed, 3.5--3.6×2.1--2.2 mm; seeds 2, globular--mitten-shaped, brownish orange--dark brown, occasionally dark-speckled, 1.2--1.5×1--1.2 mm.

Representative specimens

MÉXICO. Durango: Súchil, San Juan de Michis, rumbo al Cerro "Chihuahuilla", flores de color crema, aprox. 35cm de altura, 23.626º, - 103.929º, 2000m, 14 Aug 1984, F. Chávez 54 (TEX); on upper slopes of Mimbres Canyon, 26 miles west of Durango, Route 40, plants repent, flowers white, 24.337º, -104.306º, 2020m, 24 Jul 1958, D. S. Correll 20122 & I. M. Johnston (ASU, NY); Súchil, al S de San Juan de Michis, bosque de pino- encino, común, 23.43º, -104.12º, 2370m, 06 Aug 1981, S. González 1810 & S. Acevedo (XAL); Mpio. de El Salto, 9km al E de Llano Grande, planta herbácea de 10-20cm de alto, abundante, flores blancas, veg: bosque de pino-encino principalmente, suelo: pardo o casi blanco, 23.77º, -105.347º, 2200m, 12 Jul 1982, R. Hernández M. 7877 et al. (RSA). Guanajuato: Cañada de La Virgen, bosque de encino colorado, ladera de cerro, planta de 60cm de alto. flor roja,

91 abundante, 21.15º, -101.183º, 2500m, 13 Jun 1994, M. Cano Mares s. n. & J. Cano Mares (XAL). Guerrero: Omiltemi, tipo de vegetación es encinar-pinar, flor blanca, hierba perenne postrada, 17.554373º, -99.516250º, 2020m, 3 Aug 1967, M. Sousa 3144 (GH). Hidalgo: Hills, Cuylamaloya station, 20.033º, - 98.533º, 2575m, 9 Aug 1942, C. G. Pringle 10274 (ARIZ, ASU, DAV, GH, MICH, MU, RSA, VT). Jalisco: Bolaños, Las Banderitas, crucero Txpan de Bolaños- Las Banderitas, flora de la Sierra de Bolaños, bosque de Quercus con Alnus, hierba 0.3m, semipostrada, flores blancas, 21.922222º, -103.867778º, 2450m, 4 Jul 1996, J. Calónico-Soto 2488 & G. Flores F. (MO); pine forest 1-2 miles east of Tapalpa, rocky soil on broken hills, abundant, prostrate, flowers white, 19.9455º, -103.7376º, 2100m, 1 Nov 1960, R. McVaugh 20588 (MICH). México: Amecameca, 19.11º, -98.75º, 2773m, 29 Jul 1924, Fisher, G. L. 243 (US); calcareous bluffs, Flor de Maria, large heads, 19.34º, -99.20º, 2317m, 4 Sep 1890, C. G. Pringle 3238 (GH, MICH, MU, NY, P, RSA, VT); partially grazed, moist Quercus forest with epiphytic ferns, steep hillsides with many moist cliffs covered with mosses, Pinus-Quercus zone 6km S of Temascaltepec on Hwy 130, white flowers, 18.95º, -100.083333º, 2000m, 03 Sep 1965, K. Roe 1696 et al. (MICH, NY). México-Puebla: Ixtaccíhuatl, 19.2º, -98.6º, 4600m, Nov 1905, C. A. Purpus 1753 (GH, NY). Michoacán: Mpio. de Morelia, Lado Sureste del Cerro El Águila subiendo por Huatzanguio, bosque de Quercus, hierba con flores blancas, 19.610278º, -101.379167º, 2484m, 16 Nov 2007, E. S. García 388 et al. (MO). Morelos: hacia el Valle del Tepeite, canal Mt. Zempoala, Sta. Maria, 19.0607º, -99.3203º, 3000m, Aug 1932, H. E. Lyonnet 1029 & A. Guerra (MEXU). Nayarit: Mpio. de Acaponeta, in pine and oak forest al La Ciénaga on ridge about 10 miles northwest of Mesa del Nayar, flowers white, 22.328º, - 104.744º, 2600m, 30 Jul 1970, D. H. Norris 14579 & D. J. Taranto (MICH). Nuevo León: Zaragoza, Cerro El Viejo, oak and pine woods, flowers purple, scattered plants, 23.97º, -99.769º, 2465m, 29 Jul 1992, G. B. Hinton 22213 et al. (TEX). Oaxaca: San Miguel, Nochixtlán, Centro, 17.458º, -97.224º, 2500m, 19 Jun 1907, C. Hugo 1872 & C. Conzalli (MEXU). Puebla: Pico de Orizaba,

92 19.03º, -97.27º, 3048m, Botteri 349367 (GH). Querétaro: aprox. 3-4km al ENE de Pinal de Amoles, bosque de pino y encino, ladera de cerro, plant herbácea, flores blanco-verdosas, abundante, 21.153º, -99.596º, 2450m, 25 Oct 1988, E. Carranza 1090 (XAL). San Luis Potosí: F. C. Potosi & Rio Verde, Sierra de Alvarez, Sierra Madre Oriental, 21.9º, -100º, 2150m, 1 Aug 1934, F. W. Pennell 17763 (GH, US). Tamaulipas: southern Tamaulipas, top of Sierra Madre Oriental (Sierra de Guatemala): rugged, wooded, karstic limestone plateau with sink holes and big boulders, ca 6km NW of Rancho del Cielo (Harrison Ranch of Martin & Harrell), (ca 3km NW of Julio) on road to La Joya de Salas, ca 11km NW of Gomez Farias, 108km SSW of Ciudad Victoria, dry, open Pinus- Quercus forest with rich herbaceous cover on top of ridge, 23.2º, -99.266667º, 2060m, 12 Aug 1991, H. H. Iltis 30682 & B. Simon (MU). Veracruz: Huayacocotle, Jarillos, bosque de pino-encino, suelo aroillos, hierba, annual, 0.7m, regular, flor blanca, bracteas rosadas, 20.5º, -98.5º, 2490m, 11 Aug 1984, L. Cabrera-R. 171 & H. Nerave (XAL).

Distribution and Habitat Trifolium mexicanum is widely distributed throughout México in the states of Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tamaulipas, and Veracruz (Fig. 26). It has been collected in a variety of habitats including open pine-oak woodland, hillsides and slopes, grasslands, and along roadsides. In these locations the soil varies from sandy, calcareous, rocky, volcanic, to clay loams. Trifolium mexicanum is found from 1100 m to 4600 m in elevation.

Comments Trifolium mexicanum is a very robust species being most similar to T. hickeyi. The two are most readily differentiated by the moderate to dense pubescence and shorter peduncles of T. mexicanum.

Figure 30. Trifolium mexicanum. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary. 94 9. Trifolium sonoranensis T.K. Ahlquist et Vincent sp. nov., Fig. 31.

TYPE: UNITED STATES. Arizona: Cochise County, Huachuca Mountains, Garden Canyon. 31.49º, -110.32º, 1500m, 3 Sep 1928, G. J. Harrison 5768 & T. H. Kearney (holotype: ARIZ! 92895; isotypes: US! 1435169, 1435168).

Plants perennial; roots woody, 3.5--8 mm in diameter; stems prostrate, 25--38+ cm in length, 0.8--1.5 mm in diameter, internodes sparsely-- moderately pubescent, nodes moderately--densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular, 6.7--16.5×1.8--3.1 mm, glabrous adaxially, glabrous--sparsely pubescent abaxially, margins entire--weakly serrate, sparsely pubescent, apices acute; petioles 2.5--50.5×<1 mm, moderately--densely pubescent; petiolules 0.6--1.6×≤0.5 mm, sparsely-- densely pubescent; leaflets obovate--obcordate, glabrous adaxially, sparsely-- moderately pubescent along midribs abaxially, glabrous--sparsely pubescent abaxially on outer halves of lateral leaflets, bases acute, margins weakly serrate--serrate, often with short, straight teeth alternating with larger, antrorsely curved teeth, sparsely pubescent, apices rounded--emarginate, weakly mucronulate, terminal leaflets 9.5--16.5×6.8--11.5 mm; peduncles 28-- 50×<1 mm, moderately--densely pubescent; inflorescences axillary, racemose- umbellate, 8.5--15.5×9--15.5 mm, flowers 10--20, each subtended by a linear triangular bract, in 1--2 whorls; pedicels 1.2--2.7×<0.5 mm, sparsely--densely pubescent; calyx 3.3--5 mm long, tubes 2--3 mm in circumference, sparsely-- densely pubescent, lobes narrowly triangular, 2.5--4×≤0.5 mm, sparsely-- densely pubescent abaxially, margins entire, sparsely--densely pubescent, apices acuminate; petals white--pink; banner petals obovate, 5.3--6.5×2.8-- 3.8 mm; wing petals 4.5--5.3 mm long, asymmetrically clawed, claws 1.5-- 1.9×≤0.3 mm, laminae spatulate, 2.8--3.5 mm long, 1--1.5 mm wide at widest point, <1 mm wide at narrowest point, bases auriculate, auricles acute, margins entire, apices rounded; keel petals 3.7--4.8 mm long, asymmetrically clawed, claws 1.7--2×<0.5 mm, laminae spatulate, 1--1.3 mm long, 0.8--1.3 95 mm wide at widest point, 0.9--1.3 mm wide at narrowest point, bases truncate, margins entire, apices rounded; stamens diadelphous, connate filaments 3.1-- 4×1.1--1.5 mm, fused for 50---56% of their length, free filaments 1.9--2.5×≤0.1 mm; pistils sessile, ovaries 1.3--3.1×0.4--1.4 mm, moderately--densely pubescent distally and occasionally along suture, styles 1.3--2×≤0.2 mm, glabrous, ovules 2; legumes laterally compressed, 4.3--4.4×2--2.7 mm; seeds 2, globular--mitten-shaped, brownish orange--olive brown, occasionally dark- speckled, 1.2--1.7×1.2--1.5 mm.

Paratypes

MÉXICO. Chihuahua: Mpio. de Temoschi, Nabogame, pine/oak/cypress forest, trailing, flowers pink, 28.5º, -108.5º , 1800m, 24 Aug 1988, J. E. Laferrière, J.E. 1771 (ARIZ). Jalisco: forest of Quercus scattered with Arbutus, 14-18 km southwest of Tequila on Volcán de Tequila, flowers pink, 20.788º, - 103.847º, 2600m, 7 Nov 1974, D. E. Breedlove 39248 (MICH). Sinaloa: Mpio. de Concordia, 1-2 km N of the Mazatlan to Durango road at Loberas, pine-oak forest, perennial, flowers white, 23.47º, -105.85º, 1850m, 2 Oct 1985, B. Bartholomew 2551 et al. (GH, NY). Sonora: el llano on Mesa del Campanero (west of Yécora), uncommon herb, flowers white, 28.341667º, -109.031944º, 2100m, 16 Aug 1998, A. L. Reina G. 98-958 et al. (NY); Yecora, 9.5km west of Maycoba on Mex. 16, oak woodland, uncommon herbaceous perennial on moist roadside, flowers pink, 28.408889º, -108.725º, 1495m, 7 Aug 2000, T. R. Van Devender 2000-406 et al. (ASU, MO, RSA).

UNITED STATES. Arizona: Cochise County, wet soil at streamside in woodland with Juniperus deppeana, Pinus leiophylla, NW 1/4, Sec. 15, T23S R19E, 31.4314º, -110.4007º, 1783m, 11 Aug 1990, J. E. Bowers 3242 & S. P. McLaughlin (ARIZ); Cochise County, Huachuca Mountains, Tanner Canyon, wet places near springs, 31.49º, -110.32º, 1500m, 24 Aug 1910, L. N. Goodding 794 (ARIZ, NY, US); Cochise County, Huachuca Mountains, Garden Canyon, 31.49º, -110.32º, 1500m, 3 Sep 1928, G. J. Harrison 5768 & T. H. Kearney

96 (ARIZ, US); Cochise County, Bear Canyon, at junction of jeep trail with canyon bottom, about 2km northeast of canyon crossing with road from Parker Canyon Lake to Montezuma Pass, common in patches in moist soil, with grasses, on stream bank under Platanus wrightii, 31.3914º, -110.347º, 1770m, 26 Sep 1980, G. Yatskievych 80-719 (ARIZ).

Distribution and Habitat Trifolium sonoranensis is found largely in the Sonoran Desert of North America, but continues on further south. It has been collected in the state of Arizona in the United States, and the states of Chihuahua, Jalisco, Sinaloa, and Sonora in México (Fig. 28). It has been collected in open pine forests in canyons by stream banks or in moist soil from 1495 m to 2600 m in elevation.

Comments While Trifolium sonoranensis has no single identifying feature, it can be distinguished from other members of the complex by suite of character states not found in other species. These most notably include floral characters such as the banner, wing, and keel petals. T. sonoranensis flowers are larger than those of T. amabile and T. lozani, yet smaller than those of T. hickeyi, T. mexicanum, and T. cognatum. The most similar species in appearance to T. sonoranensis is T. laciae. However, these two species can be distinguished by their differing ranges. T. sonoranensis occurs both within and on the western side of the Sierra Madre Occidental mountain range, and T. laciae occurs on the eastern side of the range. Other differences include floral characters and the distinguishing characteristics of T. laciae such as serrate sepals, sterile projection in the inflorescence, and yellow petals, T. sonoranensis has entire sepals, lacks a sterile projection, and has white or pink flowers. T. sonoranensis is named for the Sonoran Desert, to where it is native.

Figure 31. Trifolium sonoranensis. A: habit silhouette; B: fertile branch; C: abaxial leaf surface; D: sepals; E: banner petal; F: wing petals; G: keel petals; H: connate stamens; I: free stamen; J: ovary; K: legume; L: seeds. 98 Literature Cited ADOBE SYSTEMS INCORPORATED. 2010a. Illustrator CS5 version 15.0.2. Adobe Systems Incorporated. San Jose, California, USA.

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100 Appendix 1. Table of specimens measured.

Species Tag # Barcode/ Collector Accession T. amabile ARIZ34 244262 LaSalle, J. 810709-18 et al. T. amabile DAV2 54179 Webster, G. L. 12863 T. amabile F28 892266 Dodge, C. W. 4931 & W. S. Thomas T. amabile F75 1857956 Rushin, C. J. s. n. T. amabile GH17 Nevling, L. I. 2105 & A. Gomez- Pompa T. amabile GH37 Seler 2092 T. amabile GH40 Pringle, C. G. 10278[a] T. amabile GH48 Seler 2143 T. amabile MO34 MO-1302057 Ponce, G. 032 & P. García 04831435 T. amabile NY28 Kuntze, O. 2317 T. amabile P76 P00735568 Tonduz, A. 1536 & W. Barbey T. amabile US109 2537713 Williams, L. O. 22952 et al. T. amabile US114 1794128 Balls, E. K. 4401 T. amabile US67 1182047 Maxon, W. R. 8440 & A. D. Harvey T. amabile US74 365884 Tonduz, A. 1536 T. amabile US76 228420 Pittier, H. 1536 T. amabile US78 578565 Pittier, H. s. n. T. amabile US79 2297974 Semple s. n. T. amabile US87 1252780 Standley, P. C. 42429 T. amabile US89 1306021 Standley, P. C. 49409 & J. Valerio T. amabile US99 2578129A Ross, G. B. 36 T. amabile VT22 Palmer, E. 201 T. amabile XAL48 54450 Flores, D. 174 T. blanquitum GH105 Nevling, L. I. 1859 & A. Gomez- Pompa T. blanquitum MEXU29 366949 Hernández M., R. 6927 T. blanquitum MEXU31 365063 Hernández M., R. 6666 T. blanquitum MEXU43 315903 Clausen, R. T. 7465 T. blanquitum NY248 Gillett, J.M. 17092 & A. Delgado T. cognatum ARIZ39 185297 Rzedowski 28414 T. cognatum GH8 00003503 Pringle, C. G. 6933 T. cognatum MICH92 1183835 Rzedowski 24287 T. cognatum MICH93 1183836 Rzedowski 20602 T. cognatum MICH94 1183837 Rzedowski 33557 T. goniocarpum GH1 Pringle, C. G. 13264 T. goniocarpum GH74 Nevling, L. I. 1611 & F. Chiang T. goniocarpum GH94 Palmer, E. 236

101 Appendix 1 continued.

Species Tag # Barcode/ Collector Accession T. goniocarpum NY142 Parry, C. C. 134 & E. Palmer T. goniocarpum NY30 Burquez M., A. 96-844 et al. T. goniocarpum RSA5 102930 Pringle, C. G. 1208 T. goniocarpum US154 1003918 Arsène, G. 6011 T. goniocarpum US155 2296536 Pringle, C. G. 15645 T. goniocarpum US45 332603 Nelson, E. W. 4586 T. goniocarpum XAL13 Ventura, E. 9380 & E. López T. goniocarpum XAL22 Pringle, C. G. 10282 T. hickeyi ASU10 57390 Rzedowski 22885 T. hickeyi B14 B 10 Arsène, G. 8323 0358257 T. hickeyi C19 Liebmann 4928 T. hickeyi ILL3 Arsène, G. 5483 T. hickeyi MICH1 1183721 King R.M. 2043 T. hickeyi MICH17 1183830 Rzedowski 44278 T. hickeyi MICH21 1183833 Alexander, E.J. 1265 T. hickeyi MICH48 1183745 Anderson, W. R. 13019 T. hickeyi MO29 MO-2055882 Calzada, J. I. 21326 6030486 T. hickeyi MO33 MO-1126487 Véliz, M. 92.2418 5908676 T. hickeyi NY127 Sousa, M. 7822 et al. T. hickeyi NY162 Botteri 704 T. hickeyi US144 1585057 Skutch, A. F. 88 T. hickeyi US145 1949824 Steyermark, J. A. 49895

T. hickeyi US147 840898 Purpus, C. A. 3228

T. laciae ARIZ16 252634 Van Devender, T. R. 84-523 et al. T. laciae NY1 Quintana, G. 3602 & E. Estrada T. laciae NY140 Palmer, E. 417 T. laciae P91 P00735574 Townsend, C. H. T. 177 & C. M. Barber T. laciae POM8 27959 Jones, M. E. 7501A T. lozani A2 Moore Jr. H. E. 3693 & C. E. Wood Jr. T. lozani A6 Kenoyer, L. A. 2425 T. lozani POM2 349690 Hansen, B. F. 7682 & M. Nee T. lozani VT21 Pringle, C. G. 9512

102 Appendix 1 continued.

Species Tag # Barcode/ Collector Accession T. lozani XAL54 Marín C., A. 89 T. mexicanum ASU5 124579 Correll, D. S. 20122 & I. M. Johnston T. mexicanum DAV5 59438 Pringle, C. G. 10274 T. mexicanum GH49 Pennell, F. W. 17763 T. mexicanum GH69 Purpus, C. A. 1753 T. mexicanum MICH50 1183731 McVaugh, R. 20588 T. mexicanum MO24 MO-833368 Calónico-Soto, J. 2488 & G. Flores F. 5875583 T. mexicanum MO28 MO-2007933 García, E. S. 388 et al. 6136015 T. mexicanum MU2 000089678 Pringle, C. G. 3238 47522 T. mexicanum NY158 Correll, D. S. 20122 & I. M. Johnston T. mexicanum NY172 Roe, K. 1696 et al. T. mexicanum RSA4 514999 Hernández M., R. 7877 et al. T. mexicanum TEX3 Chávez, F. 54 T. mexicanum US139 1207362 Fisher, G. L. 243 T. mexicanum XAL14 Cabrera-R., L. 171 & H. Nerave T. mexicanum XAL44 54697 González, S. 1810 & S. Acevedo T. sonoranensis ARIZ1 297681 Bowers, J.E. 3242 & S. P. McLaughlin T. sonoranensis ARIZ2 224476 Yatskievych, G. 80-719 T. sonoranensis ASU1 254530 VanDevender, T. R. 2000-406 et al. T. sonoranensis GH75 Bartholomew, B. 2551 et al. T. sonoranensis NY8 Goodding, L. N. 794 T. sonoranensis NY9 Reina G., A. L. 98-958 et al. T. sonoranensis US191 1435168 Harrison, G. J. 5768 & T. H. Kearney

103 Appendix 2. List of characters measured.

Abbreviation Character Description Habit Habit Qualitative description of the overall habit of the plant. Root Roots Qualitative description of the taproot (woody, large/small, etc.). RootDiam Root Measurement of the taproot at 1cm below the Diameter crown. Stem Stems Qualitative description of the stems (erect, ascending, prostrate, etc.) based on the label data. StemL Stem Length Measurement of the length of the stem from the crown to the apex of the terminal leaf or inflorescence of the longest stem. StemD Stem Measurement of the diameter of the stem at Diameter 1cm above the crown. InNodPub Internode Qualitative description of the pubescence of Pubescence the internodes. InNodL Internode Measurement of the internode between the Length first and second nodes from the crown, or the basal most nodes if crown was absent. NodPub Node Qualitative description of the pubescence of Pubescence the nodes. StipL Stipule Measurement of the length of the stipules Length from the base to the apex. StipW Stipule Width Measurement of the width of the stipules at the widest point. StipShp Stipule Shape Qualitative description of the shape of the stipules. StipAdPub Stipule Qualitative description of the pubescence of Adaxial the adaxial surface of the stipules. Pubescence StipAbPub Stipule Qualitative description of the pubescence of Abaxial the abaxial surface of the stipules. Pubescence StipMar Stipule Qualitative description of the margin of the Margins stipules (entire, serrate). StipApxAng Stipule Apex Measurement of the apex angle of the Angle stipules. PetL Petiole Length Measurement of the length of the petioles from the base to the apex.

104 Appendix 2 continued.

Abbreviation Character Description PetD Petiole Measurement of the diameter of the petioles. Diameter PetPub Petiole Qualitative description of the pubescence of Pubescence the petioles. LfltShp Leaflet Shape Qualitative description of the shape of the leaflets. LfltAdPub Leaflet Qualitative description of the pubescence of Adaxial the adaxial surface of the leaflets. Pubescence LfltAbMdrb Leaflet Qualitative description of the pubescence of Pub Abaxial the abaxial surface of the midrib. Midrib Pubescence LfltAbPub Leaflet Qualitative description of the pubescence of Abaxial the abaxial surface of the leaflets. Pubescence LfltBasAng Leaflet Base Measurement of the base angle of the Angle leaflets. LfltMar Leaflet Qualitative description of the margin of the Margins leaflets. LfltMarPub Leaflet Qualitative description of the pubescence of Margin the margin of the leaflets. Pubescence LfltApxAng Leaflet Apex Measurement of the apex angle of the Angle leaflets. LfltMucr Leaflet Qualitative description of the mucron of the Mucronulate leaflets. TermLfltL Terminal Measurement of the length of the terminal Leaflet Length leaflets from the base to the most distal point of the lamina. TermLfltW Terminal Measurement of the width of the terminal Leaflet Width leaflets at the widest point. PetlleL Petiolule Measurement of the length of petiolule from Length the base to the apex. PetlleD Petiolule Measurement of the diameter of the Diameter petiolules. PetllePub Petiolule Qualitative description of the pubescence of Pubescence the petiolules. Infl Inflorescence Qualitative description of the type of inflorescence.

105 Appendix 2 continued.

Abbreviation Character Description InflL Inflorescence Measurement of the length of the Length inflorescence parallel to the peduncle from the tip of the basalmost to the tip of the distalmost flower. InflW Inflorescence Measurement of the width of the Width inflorescence perpendicular to the peduncle from tip to tip of the lateral flowers. FlNo Flower Count of the number of flowers in an Number inflorescence. Bract Bracts Qualitative description of the bracts subtending the flowers of the inflorescence. Whorl Whorls Count of the number of whorls of flowers comprising the inflorescences. PeduncL Peduncle Measurement of the length of peduncles from Length the base to the apex. PeduncD Peduncle Measurement of the diameter of the Diameter peduncles. PeduncPub Peduncle Qualitative description of the pubescence of Pubescence the peduncles. PedicL Pedicel Measurement of the length of pedicels from Length the base to the apex. PedicD Pedicel Measurement of the diameter of the pedicels. Diameter PedicPub Pedicel Qualitative description of the pubescence of Pubescence the pedicels. SepL Sepal Length Measurement of the length of the sepals from the base of the tube to the apex of the lobes. SepCir Sepal Measurement of the circumference of the Circumferenc opened sepal tube at the base of the lobes. e SepTubPub Sepal Tube Qualitative description of the pubescence of Pubescence the fused sepal tubes. SepLobShp Sepal Lobes Qualitative description of the shape of the Shape sepal lobes. SepLobL Sepal Lobe Measurement of the length of the free sepal Length lobes from the base to the apex. SepLobW Sepal Lobe Measurement of the width of the free sepal Width lobes at the base. SepLobPub Sepal Lobe Qualitative description of the pubescence of Pubescence the free sepal lobes.

106 Appendix 2 continued.

Abbreviation Character Description SepLobMar Sepal Lobe Qualitative description of the margin of the Margins free sepal lobes. SepLobMar Sepal Lobe Qualitative description of the pubescence of Pub Margin the margins of the free sepal lobes. Pubescence SepLobApx Sepal Lobe Measurement of the apex angle of the sepal Ang Apex Angle lobes PetCol Petals Color Qualitative description of the color of the flower petals based on the label data. BPetShp Banner Petal Qualitative description of the shape of the Shape banner petals. BPetL Banner Petal Measurement of the length of the banner Length petal from the base to the apex. BPetW Banner Petal Measurement of the width of the banner Width petal at the widest point. WPetL Wing Petal Measurement of the wing petals from the Length base of the claw to the apex of the lamina. WPetClwL Wing Petal Measurement of the length of the claw of the Claw Length wing petals from the base of the claw to the base of the lamina. WPetClwW Wing Petal Measurement of the width of the claw of the Claw Width wing petals. WPetLam Wing Petal Qualitative description of the shape of the Lamina lamina of the wing petals. WPetLamL Wing Petal Measurement of the length of the lamina of Lamina the wing petals from the base to the apex of Length the lamina. WPetLamWw Wing Petal Measurement of the width of the lamina of Lamina Width the wing petals at the widest point. Wide WPetLamWn Wing Petal Measurement of the width of the lamina of Lamina Width the wing petals at the narrowest point. Narrow WPetLamBas Wing Petal Qualitative description of the shape of the Lamina Base base of the lamina of the wing petal. WPetLamArc Wing Petal Measurement of the apex angle of the auricle ApxAng Lamina of the lamina. Auricle Apex Angle

107 Appendix 2 continued.

Abbreviation Character Description WPetLamMar Wing Petal Qualitative description of the margin of the Lamina lamina of the wing petals. Margin WPetLamApx Wing Petal Qualitative description of the wing petal Lamina Apex lamina apices. KPetL Keel Petal Measurement of the keel petals from the base Length of the claw to the apex of the lamina. KPetClwL Keel Petal Measurement of the length of the claw of the Claw Length keel petals from the base of the claw to the base of the lamina. KPetClwW Keel Petal Measurement of the width of the claw of the Claw Width keel petals. KPetLam Keel Petal Qualitative description of the shape of the Lamina lamina of the keel petals. KPetLamL Keel Petal Measurement of the length of the lamina of Lamina the keel petals from the base to the apex of Length the lamina. KPetLamWw Keel Petal Measurement of the width of the lamina of Lamina Width the keel petals at the widest point. Wide KPetLamWn Keel Petal Measurement of the width of the lamina of Lamina Width the keel petals at the narrowest point. Narrow KPetLamBas Keel Petal Qualitative description of the shape of the Lamina Base base of the lamina of the keel petal. KPetLamMar Keel Petal Qualitative description of the margin of the Lamina lamina of the keel petals. Margin KPetLamApx Keel Petal Qualitative description of the keel petal Lamina Apex lamina apices. ConFilL Connate Measurement of the length of the connate Filaments filaments from the base to the apex of the Length longest filament. ConFilW Connate Measurement of the width of the connate Filaments filaments at the widest point. Width ConFilPcFus Connate The proportion of the connate filaments that Filament is fused. Percent Fusion

108 Appendix 2 continued.

Abbreviation Character Description FreFilL Free Filament Measurement of the length of the free Length filament. FreFilD Free Filament Measurement of the width of the free filament Diameter at the widest point. OvL Ovary Length Measurement of the length of the ovary from the base of the ovary to the base of the style. OvW Ovary Width Measurement of the width of the ovary at the widest point. OvPub Ovary Qualitative description of the pubescence of Pubescence the ovaries. StylL Style Length Measurement of the length of the style from the apex of the ovary to the base of the stigma. StylD Style Measurement of the diameter of the style. Diameter StylPub Style Qualitative description of the pubescence of Pubescence the styles. OvlNo Ovule Count of the number of ovules in the ovary. Number FruitL Fruit Length Measurement of the length of the fruit from the base of the fruit to the base of the style remnant. FruitW Fruit Width Measurement of the width of the fruit at the widest point. SeedpFruit Seeds Per Count of the number of seeds in the fruit. Fruit SeedShp Seed Shape Qualitative description of the shape of the seeds based on Gillett et al., 2001. SeedCol Seed Color Qualitative description of the color of the seeds based on Kornerup and Wanscher, 1978. SeedL Seed Length Measurement of the length of the seeds. SeedW Seed Width Measurement of the width of the seeds.

109 Appendix 3. Table of quantitative data used for analysis.

OTU StemL StemD InNodL StipL StipW StipApxAng PetL PetD LfltBasAng ARIZ34 156 1.9 6 7.5 2.8 45 11 0.5 65 DAV2 80 0.71 13.064 5.6 2.4 30 4.6 0.4 95 F28 160 1.8 16.5 7.5 2.3 45 15 0.2 50 F75 443 1.5 41 11.3 7.4 60 12.78 0.5 60 GH17 135 0.9 22 9.1 4 60 8.5 0.3 65 GH37 50 0.5 5 4.2 1.5 30 6.9 0.2 75 GH40 238.5 0.7 7 7.2 1.8 35 7.8 0.3 55 GH48 230 0.9 19 10.1 3.8 5 7 0.23 60 MO34 486 1 65.5 10.9 2.9 40 20 0.4 69 NY28 378 1.5 35.5 10.2 2.1 40 16 0.4 45 P76 185.5 0.8 6 7 2.7 30 12.9 0.3 45 US109 472.5 1.25 51 10.4 3.1 60 7 0.5 70 US114 212 1 23 7.6 4 75 7 0.3 60 US67 325.5 0.9 16 12 4 30 14 0.2 45 US74 446 0.8 33 11.5 3 30 20 0.4 45 US76 364.5 0.6 12.5 8.5 1.6 30 8.3 0.4 45 US78 242 1 9.5 7.3 1.6 15 21 0.3 40 US79 407 0.5 31.5 23 2 5 45 0.5 65 US87 185 1 11 11.5 1.1 15 32 0.5 60 US89 232 1 7 9.6 3.2 60 12.5 0.4 60 US99 205.5 0.8 14 4 1.2 30 4 0.3 85 VT22 139 1 9 7.7 2.9 45 6 0.5 45 XAL48 123 0.8 7.5 6 6 30 8.5 0.2 75 ARIZ1 324 1.1 16.5 16.5 2.5 10 7 0.9 60 ARIZ2 255.5 0.8 13 12 3 25 19 0.4 45 ASU1 321.5 1.5 39.5 13.348 3.124 15 12.5 0.8 75 GH75 26.9 1 42 15.194 1.846 15 50.5 0.5 65 NY8 283 1 15 6.7 2.3 60 13 0.5 60 NY9 385 1 23 11.644 2.485 28 5 0.7 30 US191 210 1 35.5 7 2.3 30 2.5 0.4 90 GH105 139 0.9 24.5 7 3.2 75 15 0.3 60 MEXU29 365 0.4 55 6 2.8 60 3.5 0.2 45

110 Appendix 3 continued.

OTU StemL StemD InNodL StipL StipW StipApxAng PetL PetD LfltBasAng MEXU31 55 0.4 10 4.3 1.9 75 8.4 0.2 60 MEXU43 58 0.9 7.5 5.8 1.5 45 15 0.3 85 NY248 228 1.2 12 8.8 3 45 21 0.5 45 ARIZ16 306.5 1.1 23 10 2.7 15 15 0.3 60 NY1 150 1.1 38 14 4 45 15.5 0.4 45 NY140 214.9 1.1 24.5 12.5 4 30 6 0.9 45 P91 245 1.5 29 13.5 5 45 19 0.6 45 POM8 250 1.9 18 16.5 3.5 10 41 0.7 70 ARIZ39 227 1.9 19 12 4.8 60 32 0.9 60 GH8 250 2 12 10.8 4.4 40 18 0.8 60 MICH92 219 1.6 8.5 8.9 2.7 45 11.5 0.3 45 MICH93 118 1.2 22 7.3 2.5 30 9.5 0.4 45 MICH94 240 1 10 9.8 4.1 45 13 0.5 45 GH1 527 2.5 22 12.07 3.408 15 29 2 34 GH74 138 1.5 16 4.8 1.08 15 1.3 0.5 40 GH94 528.5 1.8 15 10.3 2.9 15 4 0.7 40 NY142 286 1.1 43.5 18 2 15 22 0.5 37 NY30 246.5 1.5 5.5 15 3 30 5 0.8 40 RSA5 323 1.4 19 19 4.5 20 12 1 55 US154 263.5 1.4 19.5 13 4.5 15 8 0.7 55 US155 271.05 3.1 11 12 3 15 7 0.6 40 US45 245 1.4 12.5 12.5 3 40 7.8 0.6 30 XAL13 190 0.7 9 5.6 1.4 30 5.7 0.3 30 XAL22 186 1.1 18 10 3 10 28.5 0.7 50 ASU10 122 2 30 28 4 25 87.5 1 65 B14 511.5 4.5 45 37.5 6 40 107 1.1 77 C19 308 1.8 76 19 4 13 32 0.8 85 ILL3 400 3.5 54 20 8 5 47 1.2 100 MICH1 463 1.7 44 12.5 6 45 15 1 90 MICH17 446.5 3 86.5 21.5 6.5 10 102 1 70 MICH21 359.5 2 27 19.5 6 25 24 0.7 50 MICH48 427.5 1 46 17 6.5 20 10 0.5 100 MO29 600 2 64 15 5 30 5.5 0.5 45 MO33 394 2.5 110.5 17 6.2 40 35 1.3 55

111 Appendix 3 continued.

OTU StemL StemD InNodL StipL StipW StipApxAng PetL PetD LfltBasA NY127 422 1.5 51 17.5 7 35 9.4 0.9 65 NY162 265 1 42 10.5 2 45 59.5 0.6 75 US144 521.5 2.9 42 27 7.5 25 38.5 1 50 US145 787 2.5 74 25 8.5 15 69 1.5 87 US147 315 2.2 42.5 18 5.5 30 65 1 68 A2 109.5 0.8 15 7.4 3 45 12 0.3 55 A6 95 0.7 5.5 6 1.5 30 7 0.3 75 POM2 165 1.2 10.05 6.5 3.3 90 10 0.4 75 VT21 264 1.3 13 7.3 3.5 60 8 0.5 80 XAL54 67 0.4 15 5 1.5 45 14 0.3 40 ASU5 252 1.3 15 13 4.7 40 7.5 0.5 55 DAV5 334 1.5 100.5 12.5 3 15 19 0.5 70 GH49 223.5 1.5 9 11.36 5.1 15 9 0.8 85 GH69 695 1.7 141 10.5 3 15 8 1 45 MICH50 283 1 45 9 3.8 30 7 0.25 60 MO24 295 1.3 25.5 12 4 30 16 0.5 45 MO28 571.5 1.5 94.5 14 4 14 60.5 0.7 65 MU2 315 2.5 70 14.5 8 30 8.5 1 89 NY158 298 1.5 64 8.4 4.9 75 5 0.5 70 NY172 380 1.1 37 14 3 10 11 0.5 75 RSA4 146 2.2 15 9.5 4.4 30 16 0.7 58 TEX3 439.5 2 137.5 12.5 6 15 46 0.5 85 US139 252.5 2.3 44 14 2.5 10 6 0.8 45 XAL14 385 1.7 146 17 4 15 14.5 0.6 90 XAL44 412 1.3 80 14.5 3 9 48 0.9 90

112 Appendix 3 continued.

W OTU TermLfltL TermLfltW PetlleL PetlleD InflL Infl FlNo PeduncL ARIZ34 7 6.9 0.8 0.3 12.5 12 19 22 DAV2 5.7 4.8 0.46 0.23 4.5 5.7 4 11.9 F28 4.6 4.2 0.5 0.3 10.4 9.6 6 17 F75 13.49 8.094 0.6 0.4 15.5 15 25 38 GH17 12 8.5 0.3 0.2 8.5 15 16 39 GH37 3.3 2.8 0.4 0.2 7.5 12.5 3 9.5 GH40 6.3 5.3 0.598 0.276 5.2 9.9 3 12.354 GH48 5.75 5.2 0.3 0.24 8 13.632 10 23 MO34 15 12.5 1 0.4 5.5 0.9 10 42 NY28 6.2 5.9 0.6 0.3 5 7 20 12.78 P76 5.9 5.6 0.3 0.3 12 12 6 12.5 US109 10.4 7.9 0.6 0.3 10 15 7 50.5 US114 7.2 6.9 0.5 0.3 9 12 15 33 US67 8.5 6.5 0.7 0.5 15.5 14.5 13 17 US74 14 10 1 0.4 10.5 13 12 47 US76 4.5 3.5 0.5 0.4 12.5 12 7 11.5 US78 5.6 4.6 0.5 0.2 5.8 8.9 8 17 US79 11.928 9.6 0.7 0.5 8.5 10.5 8 46.5 US87 12.212 10.2 1.2 0.4 7.3 11.9 7 9.5 US89 6.4 5 0.5 0.3 11 13 6 15 US99 6.4 6 0.6 0.3 8.5 11.5 18 34.5 VT22 12.1 5.5 0.6 0.3 11.5 13 13 37 XAL48 5.7 5.4 0.5 0.2 9 13 16 30 ARIZ1 19 11.5 0.8 0.4 9 14.5 17 42 ARIZ2 13 9 0.7 0.4 9.5 15.5 12 43 ASU1 15.336 9.23 0.6 0.5 9 15.5 20 42 GH75 16.472 8.2 1.3 0.5 8.6 9 18 29.5 NY8 9.65 7.4 0.8 0.5 15.5 16 10 49.5 NY9 12 6.8 1.6 0.6 8.946 13.348 12 28 US191 10.1 7.6 0.69 0.276 9 14 10 36.5 GH105 9.4 7.7 0.7 0.3 5.2 9 9 32 MEXU29 8.3 5 0.6 0.2 8 9 11 15

113 Appendix 3 continued.

OTU TermLfltL TermLfltW PetlleL PetlleD InflL InflW FlNo PeduncL MEXU31 4.2 2.8 0.3 0.2 6 8.4 5 13 MEXU43 4.3 3.4 0.5 0.2 6 9 7 15 NY248 12.5 7.5 0.9 0.3 12 11.5 13 40 ARIZ16 14 7.5 0.7 0.3 6.5 11.5 15 13 NY1 21 8 1 0.3 11 18 16 50 NY140 18 8 1 0.5 15.5 15.5 16 40 P91 21.5 9 1 0.4 9 13 18 40 POM8 21.5 9 1 0.4 8 13 14 33.5 ARIZ39 12.5 8 1 0.4 10.5 16 21 50 GH8 15 8 0.8 0.4 20.5 19.5 30 70 MICH92 11.4 5.8 0.9 0.3 9 17.5 18 53 MICH93 9 5 0.5 0.3 19.5 20 12 27 MICH94 10 5.5 0.6 0.4 17 19 27 50 GH1 28.5 7 0.5 0.4 12 14 14 46.5 GH74 8.9 2.9 0.7 0.3 11.2 10.2 13 3.36 GH94 18.5 7 0.9 0.3 13.5 16.5 13 49 NY142 22.5 7 1.1 0.3 11.5 12.5 13 50 NY30 16 5.5 1 0.3 14 14 14 48 RSA5 22 9 0.6 0.5 7 10 18 46 US154 19.5 6 0.9 0.3 12 12 23 41.5 US155 16 3.5 0.5 0.4 10 11 21 30 US45 18 5.5 1 0.4 9 13 17 77 XAL13 9.6 2.6 0.4 0.3 10.5 11.5 11 42 XAL22 18 6 0.8 0.3 6 8 14 27 ASU10 29 10 1.4 0.6 20 19.5 35 155 B14 46 17.5 0.8 1.2 22 22.5 40 116 C19 16 9.5 1.1 0.4 21 20 30 186 ILL3 37.5 13 1 0.9 24 23 50 147 MICH1 23.5 11 1 0.4 19 21 36 157 MICH17 33 12 1.2 0.7 22.5 23.5 30 133 MICH21 34 8.5 0.7 0.5 21 20 35 156 MICH48 22.5 16.5 4 0.4 23 22 30 131 MO29 13.5 5.5 0.8 0.3 22 22.5 20 76 MO33 37.5 10 1.2 0.5 14 18.5 25 104

114 Appendix 3 continued.

OTU TermLfltL TermLfltW PetlleL PetlleD InflL InflW FlNo PeduncL NY127 24.5 9.5 0.5 0.5 20.5 20.5 32 89 NY162 16.5 9.5 1 0.4 20 19.5 26 123 US144 37 12 1.5 0.7 26 26 30 211 US145 45.5 18.5 1.2 0.7 NA 23 28 152.5 US147 24.5 12.5 0.8 0.5 9.5 16 17 161 A2 8 4.4 0.5 0.3 10.5 9.5 28 26.5 A6 6.2 5.2 0.6 0.3 10 11 18 15 POM2 9 5.5 0.5 0.3 11.5 12.5 25 26 VT21 11.2 7.9 0.5 0.5 12 13 22 31 XAL54 6.1 4 0.3 0.2 4.5 7 13 23 ASU5 16.5 9.5 0.9 0.3 15 15 20 23 DAV5 13 8.5 0.6 0.4 15 18.5 21 34.5 GH49 14.484 9.798 0.92 0.391 10 17 20 23 GH69 26.5 10 1 0.3 24 23.5 50 57 MICH50 19 11 0.8 0.4 18.5 18 26 45 MO24 11.5 7 1 0.4 17.5 18 26 41 MO28 24.5 16 1.2 0.5 15.904 16.756 30 78.5 MU2 22.5 14.5 1 0.6 20.5 21.5 25 40 NY158 10.1 7.8 0.9 0.4 21 22 28 43 NY172 15 9.5 1.1 0.4 16 17 27 43 RSA4 9.7 5.9 0.7 0.3 14.5 16 33 46 TEX3 25.2 16 0.6 0.5 20 21 20 53.5 US139 24 10 1 0.3 20 19 40 34 XAL14 21 16.5 1 0.5 22 24 28 36 XAL44 25.5 18 1 0.7 19 18 25 31.5

115 Appendix 3 continued.

OTU PeduncD PedicL PedicD SepL SepCir SepLobL SepLobW SepLobApxAng BPetL BPetW ARIZ34 0.5 1.5 0.2 3.3 2.5 2.5 0.5 5 5 3.3 DAV2 0.4 0.9 0.18 3.5 2.6 2.4 0.3 5 3.7 2.7 F28 0.3 1.5 0.3 3.9 2.5 2.7 0.5 10 5 2.6 F75 0.7 2 0.15 5 2.5 3.8 0.4 5 5.7 3.8 GH17 0.5 1.1 0.2 3.5 2.9 2.8 0.7 10 5.4 3.9 GH37 0.2 0.7 0.2 3 2.7 2 0.6 10 6.1 3.9 GH40 0.2 1 0.2 2.5 2.2 1.4 0.4 15 4.4 3.1 GH48 0.4 1.38 0.345 3.1 3.2 2.3 0.4 15 7.3 4.5 MO34 0.4 1.5 0.2 4.5 2.9 3.5 0.6 10 5.2 3.4 NY28 0.5 1 0.4 3.3 2.4 2.2 0.5 10 3.1 2.2 P76 0.2 1.3 0.2 3.2 2.5 2.5 0.5 10 5.8 3.3 US109 0.3 1.8 0.3 3.5 2.5 2.5 0.7 15 5.8 4 US114 0.5 1.5 0.2 2.9 2.8 2.2 0.5 5 4.4 5.6 US67 0.5 2 0.2 3.8 2.5 2.9 0.6 5 5.6 2.8 US74 0.3 2.5 0.2 3.5 1.9 2.7 0.5 10 5.2 2.8 US76 0.3 1.6 0.2 3.3 2.4 2.3 0.3 7 4.9 2.6 US78 0.3 1.9 0.2 2.9 2.2 2 0.5 10 5.4 2.9 US79 0.5 1.656 0.184 3.45 2.3 2.5 0.4 10 4.5 2.4 US87 0.4 0.506 0.23 3.6 2.8 2.3 0.4 15 6 3.2 US89 0.5 1.1 0.2 3.85 3.1 2.8 0.65 13 5.5 3.5 US99 0.4 1.9 0.15 3.2 2.3 2.5 0.5 5 4 2.9 VT22 0.5 2.5 0.2 3.8 2.7 2.7 0.5 10 4.9 3.2 XAL48 0.4 1.8 0.3 2.7 1.9 1.8 0.4 10 4.7 2.6 ARIZ1 0.7 2.7 0.3 5 2.3 3.9 0.5 5 5.7 3.5 ARIZ2 0.5 3 0.2 4.3 2.4 3.4 0.5 5 6.1 3.5 ASU1 0.6 2.3 0.2 4.4 2.6 3.1 0.4 5 6 3.8 GH75 0.5 1.265 0.391 4.55 3 3 0.4 10 5.6 3.3 NY8 0.5 1.3 1.5 4.3 2.1 3.4 0.4 3 4.9 2.8 NY9 0.35 2.369 0.184 3.3 2.5 2.5 0.4 10 5.3 3 US191 0.5 2.07 0.345 4.2 2.8 3.1 0.3 10 6.4 3.4 GH105 0.2 1 0.1 3.4 2.1 2.5 0.4 15 3.6 2.45 MEXU29 0.3 1.5 0.1 2.4 2.2 1.5 0.5 15 3.3 2.1

116 Appendix 3 continued.

OTU PeduncD PedicL PedicD SepL SepCir SepLobL SepLobW SepLobApxAng BPetL BPetW MEXU31 0.2 1.2 0.15 2.6 2.7 1.8 0.6 15 4.9 2.9 MEXU43 0.3 1.3 0.1 2.9 2.5 1.6 0.5 15 4.5 3 NY248 0.2 2 0.15 2.85 2.4 2 0.6 15 4.4 2.4 ARIZ16 0.5 2 0.2 3.7 2.8 2.9 0.6 10 3.6 2.6 NY1 0.5 4.5 0.2 5.7 3.5 4.6 0.6 15 5.7 4.2 NY140 0.6 4.5 0.3 4.7 2.9 3.9 0.5 5 5.1 3.4 P91 0.9 1.6 0.4 4.2 2.8 3 0.6 15 5 3.5 POM8 0.5 2.1 0.3 3.3 2.5 2.6 0.5 10 4.7 2.4 ARIZ39 0.6 2.3 0.2 4.5 3.2 3.5 0.7 5 7.3 4.5 GH8 1 3 0.2 3.6 2.4 2.5 0.5 5 7.2 3.4 MICH92 0.9 1.8 0.4 4.5 3.1 3.3 0.7 5 5.6 3.9 MICH93 0.4 2.5 0.2 3.9 3 2.8 0.5 5 4.7 3.4 MICH94 0.6 3.2 0.2 3.7 3.3 2.9 0.7 10 7.6 4 GH1 0.6 0.9 0.3 4.8 3.4 3 0.5 13 5.4 3 GH74 0.4 1.3 0.2 2.5 2 1.8 0.2 10 3.9 2.3 GH94 0.6 3 0.2 4.7 2.2 3.6 0.5 7 5 2.5 NY142 0.4 1.9 0.2 4.5 1.7 3.7 0.4 5 4.2 2.2 NY30 0.4 1.8 0.2 5.2 2.2 4.4 0.3 5 4.6 2.4 RSA5 0.7 2 0.2 4.1 2.3 3.3 0.4 5 4.7 2.6 US154 0.5 2 0.2 3.15 2.2 2.4 0.4 10 4.4 2.8 US155 0.5 1.7 0.2 2.5 2.2 2 0.4 10 3.6 2.2 US45 0.5 2.5 0.2 4 1.9 2.9 0.4 5 5.3 2.9 XAL13 0.4 1.4 0.1 3.3 2 3 0.4 5 4 2.2 XAL22 0.5 1.7 0.2 3 2.1 2.1 0.4 10 3.7 2 ASU10 1 3.6 0.2 4.5 3.5 3 0.6 5 7.1 3.8 B14 1.5 2.3 0.2 6.4 3.3 5 0.5 5 8.1 4.6 C19 1 3.3 0.2 5 3.6 3.5 0.7 10 9 5.4 ILL3 2 3.2 0.3 4.9 3.2 3.7 0.5 5 8.1 4.6 MICH1 1 1.7 0.3 4.7 4.1 3.7 0.9 10 8.7 6.3 MICH17 0.9 1.3 0.3 5.6 3.1 4.5 0.7 5 7.3 4.8 MICH21 1.1 3.3 0.2 5.7 3.6 4.3 0.8 5 7.9 5.3 MICH48 0.8 1 0.2 5.6 3.4 4.7 0.5 5 9.2 5.3 MO29 1 2.4 0.3 3.9 3 3 0.5 10 8 5 MO33 0.852 2.1 0.3 6.5 3.3 5.1 0.5 10 6 3.9

117 Appendix 3 continued.

OTU PeduncD PedicL PedicD SepL SepCir SepLobL SepLobW SepLobApxAng BPetL BPetW NY127 1 1.4 0.2 5 3.3 3.9 0.5 5 7.7 5.3 NY162 0.9 2.9 0.2 4.9 3.3 3.8 0.6 5 8.1 5.3 US144 1.5 2.8 0.3 6.8 3.9 5.2 0.8 5 8.9 5.7 US145 1 2.5 0.4 7.4 4.1 6 0.8 3 9.6 6 US147 1 0.7 0.2 5 2.6 3.4 0.5 5 7.3 4.3 A2 0.5 1 0.2 2.6 1.9 1.8 0.4 10 3.6 2.8 A6 0.3 0.8 0.15 2.9 2.4 2.3 0.5 10 4.1 3.1 POM2 0.65 2 0.1 2.6 2.4 2.1 0.5 5 3.95 3.2 VT21 0.9 2.4 0.2 3.3 2.6 2.4 0.5 10 4.2 3 XAL54 0.3 1.2 0.1 2.2 1.9 1.7 0.4 10 2.5 2.1 ASU5 0.5 1 0.3 3.9 3.1 3.1 0.7 10 6.1 4.2 DAV5 0.5 2.2 0.2 4.4 3.1 3.1 0.6 10 7 4.1 GH49 0.6 1.955 0.23 6.1 4 4.8 0.4 5 6.9 4.5 GH69 0.9 3 0.2 3.5 2.7 4.6 0.5 5 7.45 4.4 MICH50 0.8 2.5 0.2 4.8 2.7 3.5 0.6 3 6.7 3.5 MO24 0.5 2.5 0.2 4.7 3.7 3.4 0.7 10 6.4 4.8 MO28 0.5 2.4 0.24 4.7 3.2 3.7 0.4 5 7.2 4.6 MU2 1 2.4 0.2 6 3.5 4.3 0.7 4 8.5 5.1 NY158 0.8 3.5 0.2 5.4 3 4.5 0.5 5 7.4 5.4 NY172 0.5 1.6 0.2 4.5 2.8 3.8 0.5 5 6.3 4.2 RSA4 0.7 3 0.2 3.4 2.3 2.3 0.7 15 5.7 3.9 TEX3 1 2.3 0.2 5.2 2.8 3.2 0.2 2 7.2 5 US139 0.6 3.1 0.2 5 3.2 4 0.5 5 6.7 4.6 XAL14 0.8 4.2 0.2 4.8 2.9 3.9 0.5 5 8.2 4.6 XAL44 0.6 3.7 0.2 5 3.2 3.8 0.5 5 7 5.5

118 Appendix 3 continued.

OTU WPetL WPetClwL WPetClwW WPetLamL WPetLamWw WPetLamWn WPetLamArcAp xAng KPetL KPetClwL ARIZ34 4.5 1.4 0.2 3.1 0.9 0.6 45 3.6 1.6 DAV2 3.5 1 0.2 2.5 1.2 0.7 15 3.3 1 F28 4.9 1.4 0.2 3.5 0.9 0.7 75 4 1.5 F75 4.9 1.2 0.2 3.7 1.15 0.6 105 3.9 1.4 GH17 4.9 1.1 0.2 3.8 1.6 0.9 75 4 1.2 GH37 4.9 1.4 0.3 3.5 1.6 0.9 60 4.3 1.5 GH40 3.7 1.1 0.1 2.6 1 0.7 42 3.4 1.4 GH48 7.15 2.65 0.2 4.5 2 0.9 47 4.3 1.633 MO34 4.7 1.4 0.2 3.3 1.2 0.7 60 3.7 1.4 NY28 2.85 0.75 0.3 2.1 0.9 0.6 80 2.7 0.9 P76 5.1 1.5 0.2 3.6 1.3 0.7 45 4 1.6 US109 5.2 1.4 0.275 3.8 1.3 0.7 45 4.1 1.6 US114 4.5 1.2 0.2 3.3 1.5 0.7 75 3.7 1.5 US67 4.9 1.6 0.2 3.3 1.1 0.7 50 4 1.6 US74 4.5 1.3 0.2 3.2 1.1 0.6 55 3.4 1.4 US76 4.5 1.3 0.2 3.2 0.9 0.7 50 3.6 1.3 US78 4.7 1.3 0.2 3.4 1.1 0.7 45 3.6 1.5 US79 4.1 1.2 0.2 2.9 1 0.6 60 3.2 1.2 US87 5.3 1.38 0.207 3.92 0.9 0.7 45 3.816 1.62 US89 4.8 1.3 0.3 3.5 1.35 0.65 60 3.9 1.6 US99 3.6 1.3 0.1 2.3 0.9 0.6 60 3.2 1.4 VT22 4.3 1.1 0.2 3.2 1 0.8 75 3.7 1.3 XAL48 3.8 1.2 0.2 2.6 0.8 0.6 75 3.3 1.5 ARIZ1 4.7 1.6 0.2 3.1 1.3 0.85 70 4.3 1.8 ARIZ2 5.3 1.8 0.3 3.5 1.2 0.9 60 4.5 1.9 ASU1 5.3 1.75 0.2 3.55 1.472 0.8 60 4.8 2 GH75 4.98 1.9 0.2 3.08 1.4 0.9 30 4.7 1.7 NY8 4.5 1.7 0.15 2.8 1 0.8 60 4.1 1.7 NY9 4.7 1.5 0.2 3.2 1.1 0.7 60 3.7 1.7 US191 5.1 1.8 0.2 3.3 1.2 0.8 50 4.3 1.8 GH105 3 0.8 0.2 2.2 0.8 0.6 90 2.3 1 MEXU29 2.9 0.9 0.2 2 0.7 0.6 90 2.9 1.3

119 Appendix 3 continued.

OTU WPetL WPetClwL WPetClwW WPetLa WPetLamWw WPetLamWn WPetLamArcAp xAng KPetL KPetClwL MEXU31 4 1.2 0.2 2.8 1 0.7 75 3.4 1.4 MEXU43 3.7 1.1 0.2 2.5 1.1 0.8 75 3.3 1.1 NY248 3.75 1.2 0.2 2.55 1 0.6 60 3.2 1.2 ARIZ16 3.4 0.9 0.2 2.6 0.8 0.6 65 2.8 0.8 NY1 5 1.2 0.2 3.7 1.2 0.9 75 3.8 1.25 NY140 4.5 1.2 0.2 3.3 0.9 0.7 90 3.7 1.4 P91 4.5 1.2 0.3 3.2 1.1 0.8 80 3.7 1.3 POM8 3.9 1.1 0.15 2.8 0.9 0.8 90 3.5 1.2 ARIZ39 6.4 1.7 0.2 4.7 1.7 1 20 5 1.8 GH8 6 1.5 0.2 4.5 1.6 0.9 55 4.8 1.5 MICH92 5.5 1.4 0.4 4.1 1.8 1.2 60 4.8 1.7 MICH93 4.5 0.9 0.2 3.6 1.4 0.9 35 3.7 0.9 MICH94 6.7 1.6 0.2 5.1 2.7 1 65 4.9 1.6 GH1 4 1.3 0.2 2.7 0.75 0.6 65 3.24 1.38 GH74 3 1 0.1 2 0.6 0.5 90 2.8 1.1 GH94 4 1.4 0.15 2.5 0.6 0.5 60 3.5 1.5 NY142 3.6 1 0.2 2.5 0.65 0.4 65 2.9 1.1 NY30 3.7 1.3 0.2 2.4 0.7 0.5 65 3.2 1.4 RSA5 3.6 1.3 0.2 2.4 0.6 0.6 90 3.1 1.5 US154 3.6 1.3 0.2 2.3 0.7 0.7 90 3.1 1.4 US155 3 1 0.2 2 0.6 0.5 90 2.8 1.2 US45 4.5 1.4 0.2 3 0.8 0.6 75 3.5 1.6 XAL13 3.2 1.2 0.2 2 0.6 0.5 75 2.9 1.3 XAL22 3.2 1.3 0.2 1.9 0.5 0.5 60 2.9 1.3 ASU10 5.5 1.4 0.3 4.2 1.7 0.9 75 4.7 1.6 B14 6.8 1.9 0.2 5 1.9 1.1 70 5.7 2 C19 7.7 1.3 0.4 6.2 2.4 1.3 90 5.5 1.6 ILL3 6.7 1.8 0.2 4.9 1.9 1 60 5.4 2 MICH1 7 1.2 0.5 5.8 2.9 1.3 90 4.9 1.5 MICH17 6.7 1.8 0.3 4.8 2 1.1 75 5.7 2 MICH21 6.8 1.9 0.3 4.8 2.1 1.1 75 5.7 2.3 MICH48 7.5 1.9 0.2 5.6 2.8 1.1 50 5.6 1.5 MO29 7 1.5 0.4 5.5 2.4 1.2 120 5.4 1.6 MO33 5.6 1.4 0.3 4.2 1.7 1 80 4.9 1.5

120 Appendix 3 continued.

lwL

OTU WPetL WPetClwL WPetClwW WPetLamL WPetLamWw WPetLamWn WPetLamArcAp xAng KPetL KPetC NY127 6.9 1.2 0.4 5.7 2.2 1.2 90 5.3 1.8 NY162 6.3 1.65 0.2 4.6 2.2 1.1 60 5.2 1.7 US144 7.8 1.9 0.3 5.9 2.5 1.1 90 5.9 2 US145 8.2 2.1 0.4 6.1 2.7 1.4 65 6.6 2.4 US147 6.2 1.1 0.2 5.1 2 0.9 90 4.5 1.4 A2 3.5 1.1 0.2 2.4 0.9 0.6 90 2.8 1.3 A6 3.8 1.1 0.2 2.7 1 0.6 90 3.1 1.3 POM2 3.5 0.8 0.2 2.6 1.2 0.6 90 2.8 1 VT21 3.35 0.9 0.2 2.5 1.1 0.6 75 3 1 XAL54 2.3 0.5 0.2 1.8 0.8 0.5 60 2.2 0.8 ASU5 5.5 1.65 0.3 3.85 1.7 1 30 5.1 1.8 DAV5 6.2 1.9 0.2 4.25 2 1 75 5 2 GH49 6.4 1.9 0.18 4.5 1.8 0.9 75 5.2 2.1 GH69 6.2 1.7 0.2 4.45 2 1 50 5.4 1.9 MICH50 6 2.2 0.25 3.8 1.3 0.8 60 5.2 2.4 MO24 5.6 1.5 0.3 4.1 1.7 1.1 50 5.1 1.7 MO28 6.1 1.8 0.2 4.3 2 1.1 65 5.4 2.1 MU2 7.3 2.1 0.2 5.2 2.3 1.1 60 5.4 1.8 NY158 6 2 0.2 4 1.7 0.9 70 5.1 1.7 NY172 5.8 1.6 0.3 4.2 1.6 0.9 55 4.8 1.8 RSA4 5.1 1.4 0.2 3.7 1.7 0.9 75 4.4 1.6 TEX3 6.7 1.7 0.2 5 2.15 1.2 60 5.2 2 US139 5.8 1.6 0.2 4.2 1.7 0.9 50 5 1.8 XAL14 7.1 1.9 0.2 5.2 1.8 0.9 60 5.9 2.1 XAL44 5.8 1.7 0.3 4.1 2 0.9 80 4.8 1.7

121 Appendix 3 continued.

OTU KPetClwW KPetLamL KPetLamWw KPetLamWn ConFilL ConFilW ConFilPcFus FreFilL ARIZ34 0.3 2 1.1 0.9 3 1.2 56.67 2.2 DAV2 0.3 2.3 1.1 0.3 2.2 1.3 59.09 1 F28 0.4 3.5 1 0.8 2.5 1.3 72 1.8 F75 0.2 2.5 1.2 0.8 3.6 1.2 44.44 2.65 GH17 0.4 2.8 1.4 1 2.7 1.4 44.44 2.1 GH37 0.3 2.8 1.3 1 3 1.3 53.33 1.4 GH40 0.2 2 1.1 0.9 2.7 1 51.85 1.6 GH48 0.3 2.667 1.2 1.2 3.5 1.5 42.86 1.56 MO34 0.3 2.3 1 0.8 2.8 1.2 57.14 1.85 NY28 0.3 1.8 0.9 0.7 3 NA 53.67 NA P76 0.3 2.4 1.1 0.9 3.2 1.3 50 2.05 US109 0.3 2.5 1.1 0.8 3.3 1.2 51.52 1.8 US114 0.3 2.2 1.2 0.9 3 0.9 60 2 US67 0.3 2.4 1 0.9 3.2 1.4 62.5 2.2 US74 0.3 2 0.9 0.8 2.6 1 53.85 1.7 US76 0.2 2.3 0.9 0.9 3 1.1 60 1.8 US78 0.2 2.1 1.05 0.9 3 1 53.55 1.8 US79 0.2 2 1 0.7 2.8 0.966 46.43 1.6 US87 0.36 2.196 0.972 0.756 2.898 1.288 59.52 1.56 US89 0.3 2.3 1.15 0.9 3.2 1.2 46.88 1.8 US99 0.2 1.8 0.9 0.9 2.9 1.1 51.72 2.1 VT22 0.3 2.4 1.1 1 3 1.1 53.33 2 XAL48 0.2 1.8 0.9 0.6 2.6 1 50 1.8 ARIZ1 0.3 2.5 1.2 0.9 3.8 1.4 55.26 2.2 ARIZ2 0.4 2.5 1.2 0.9 3.6 1.5 50 2.5 ASU1 0.3 2.8 1.3 1.1 3.6 1.5 55.56 2.4 GH75 0.3 3 1.3 1.3 4 1.4 50 2.3 NY8 0.15 2.4 1 0.8 3.6 1.1 52.78 2.1 NY9 0.3 2 1.1 0.8 3.1 1.2 51.61 1.9 US191 0.3 2.5 1.2 1 4 1.4 52.5 2.6 GH105 0.2 1.3 1 0.8 2.2 1.1 40.91 1.1 MEXU29 0.2 1.6 0.9 0.8 2.3 1 52.17 1.1

122 Appendix 3 continued.

OTU KPetClwW KPetLamL KPetLamWw KPetLamWn ConFilL ConFilW ConFilPcFus FreFilL MEXU31 0.3 2 1.1 0.9 2.7 1.1 55.56 1.6 MEXU43 0.3 2.2 1.2 0.9 2.6 1.4 53.85 1.5 NY248 0.3 2 1.1 1 2.4 0.9 54.17 0.95 ARIZ16 0.25 2 1 0.6 2.2 1.6 54.55 1.2 NY1 0.2 2.5 1.3 0.9 3 1.5 53.33 2.3 NY140 0.15 2.2 1.1 0.9 2.9 1.05 55.17 1.8 P91 0.3 2.4 1.2 1 3.4 1.3 47.06 NA POM8 0.2 2.3 0.9 0.8 2.9 1.1 51.72 2.3 ARIZ39 0.4 3.2 1.6 1.1 4.5 1.3 55.56 3 GH8 0.3 3.1 1.4 1 4.2 1.4 45.24 2.9 MICH92 0.4 3.3 1.7 1.3 4.2 1.6 52.38 2.6 MICH93 0.3 2.8 1.3 1.1 2.9 1.3 48.28 1.6 MICH94 0.3 3.3 1.5 1.1 4.2 1.5 54.76 3.4 GH1 0.24 1.86 0.9 0.78 2.599 1 50.79 1.656 GH74 0.2 1.7 0.8 0.7 2.76 1.5 50 NA GH94 0.2 2 0.9 0.7 3 1.1 56.67 1.9 NY142 0.2 1.7 0.8 0.7 2.6 1.2 50 1.7 NY30 0.25 1.7 0.8 0.7 2.7 1.1 55.56 1.8 RSA5 0.25 1.7 0.8 0.7 2.7 1.1 59.26 1.7 US154 0.2 1.7 0.9 0.7 2.7 1 51.85 1.8 US155 0.3 1.5 0.8 0.7 2.4 0.8 54.17 1.5 US45 0.2 2 1 0.8 2.9 1.1 51.72 1.8 XAL13 0.3 1.6 0.7 0.6 2.5 1 56 1.7 XAL22 0.3 1.7 0.8 0.6 2.5 1.1 44 1.6 ASU10 0.4 3 1.5 1.2 4.3 1.3 62.79 3 B14 0.2 3.7 1.6 1.1 5 1.3 64 3.9 C19 0.4 3.8 1.9 1.3 4.5 1.45 66.67 2.8 ILL3 0.3 3.2 1.6 1.1 4.3 1.4 69.77 3.4 MICH1 0.6 3.4 1.7 1.1 4.5 1.6 53.33 2.6 MICH17 0.3 3.6 1.8 1.3 4.9 1.4 61.22 3.1 MICH21 0.4 3.5 1.8 1.4 4.9 1.4 61.22 3.5 MICH48 0.4 4 2 1.3 4.4 1.15 50 2.2 MO29 0.4 4 1.8 1.1 5 1.4 56 2.3 MO33 0.3 3.4 1.5 1.2 3.7 1.3 67.57 2.1

123 Appendix 3 continued.

OTU KPetClwW KPetLamL KPetLamWw KPetLamWn ConFilL ConFilW ConFilPcFus FreFilL NY127 0.5 3.7 1.9 1.4 4.3 1.5 60.47 2.7 NY162 0.4 3.4 1.7 1.2 4.8 1.5 70.83 3.2 US144 0.4 3.7 1.9 1.2 5 1.4 66 3.9 US145 0.5 4.2 2 1.5 5.6 1.2 60.71 4.4 US147 0.3 3.3 1.7 0.9 3.7 1.1 59.46 2.1 A2 0.3 1.5 0.9 0.7 2.5 1.1 48 1.5 A6 0.2 1.8 1 0.7 2.7 1.2 55.56 1.9 POM2 0.2 1.8 1.1 0.8 2.3 1 47.83 1.5 VT21 0.2 2 1 0.7 2.4 1.2 37.5 1.8 XAL54 0.3 1.4 0.8 0.7 1.6 0.8 50 0.7 ASU5 0.4 3.3 1.5 0.9 4 1.4 57.5 3 DAV5 0.3 3 1.5 0.9 4.5 0.4 46.67 3.1 GH49 0.1 3.1 1.6 0.8 4.4 1.4 52.27 2.9 GH69 0.3 3.5 1.5 0.8 4.3 1.5 48.84 2.9 MICH50 0.2 2.8 1.4 0.9 4.9 1.1 51.02 3.4 MO24 0.4 3.4 1.7 1.2 4.2 1.6 52.38 2.1 MO28 0.3 3.3 1.7 1.1 4.2 1.2 59.52 2.9 MU2 0.3 3.6 1.5 1.3 5.1 1.5 56.86 3.5 NY158 0.3 3.3 1.6 0.8 4.1 1.3 58.54 2.8 NY172 0.2 3 1.4 1 4.1 1.4 46.34 2.6 RSA4 0.3 2.9 1.4 1 3.4 1.4 61.76 2.4 TEX3 0.35 3.2 1.2 1 4.6 1.3 57.61 2.4 US139 0.3 3.2 1.4 0.8 4 1.4 55 2.7 XAL14 0.3 3.9 1.7 1 4.8 1.2 47.92 3.2 XAL44 0.3 3 1.4 0.7 4.8 1.3 45.83 2.8

124 Appendix 3 continued.

L OTU FreFilD Ov OvW StylL StylD OvlNo ARIZ34 0.1 1.8 0.5 1 0.2 2 DAV2 0.2 0.92 0.46 1.035 0.1161 2 F28 0.15 1.6 0.6 0.7 0.15 2 F75 0.1 1.3 0.55 1.2 0.1 2 GH17 0.1 3.1 1.5 1.1 0.2 2 GH37 0.2 2 0.9 1.1 0.2 2 GH40 0.15 1.6 0.7 1.05 0.1 2 GH48 0.12 1.4 0.5 1.3 0.1 2 MO34 0.1 1.6 0.7 1.2 0.2 2 NY28 NA 1.311 0.4 0.69 0.138 2 P76 0.2 1.4 0.6 0.9 0.1 2 US109 0.05 1.6 0.6 1 0.15 2 US114 0.1 1.5 0.55 1.05 0.2 2 US67 0.15 2.3 0.9 0.8 0.2 2 US74 0.1 1.6 0.6 1 0.15 2 US76 0.1 1.4 0.5 1 0.1 2 US78 0.1 1.4 0.5 1.1 0.1 2 US79 0.1 1.5 0.5 0.805 0.184 2 US87 0.12 2.1 0.7 1 0.1 2 US89 0.15 1.5 0.6 0.9 0.2 2 US99 0.1 2 0.7 0.7 0.1 2 VT22 0.1 1.6 0.6 1.1 0.15 3 XAL48 0.1 1.4 0.5 0.9 0.1 2 ARIZ1 0.1 1.8 0.6 1.5 0.2 2 ARIZ2 0.1 1.7 0.6 1.5 0.2 2 ASU1 0.1 2.1 0.7 1.3 0.2 2 GH75 0.1 1.6 0.5 2 0.2 2 NY8 0.15 1.5 0.45 1.495 0.138 2 NY9 0.069 1.3 0.55 1.38 0.852 2 US191 0.1 3.1 1.4 1.5 0.1 2 GH105 0.1 1.2 0.6 0.7 0.1 2 MEXU29 0.1 1 0.5 0.7 0.15 2

125 Appendix 3 continued.

OTU FreFilD OvL OvW StylL StylD OvlNo MEXU31 0.1 1.2 0.5 1 0.15 2 MEXU43 0.1 1.1 0.5 0.8 0.1 2 NY248 0.1 1.5 0.8 0.8 0.1 2 ARIZ16 0.1 2.7 1.1 0.6 0.1 3 NY1 0.1 2.3 0.9 0.8 0.2 3 NY140 0.05 1.4 0.5 1.1 0.15 3 P91 1.6 0.6 1.1 0.1 3 POM8 0.05 2.5 1.3 0.7 0.1 3 ARIZ39 0.1 2 0.8 2 0.2 2 GH8 0.2 2.5 1.1 1.7 0.15 2 MICH92 0.1 2.3 0.5 1.8 0.2 2 MICH93 0.2 1.4 0.5 1.7 0.2 2 MICH94 0.1 2.4 0.7 1.7 0.2 2 GH1 0.046 3.6 1.3 3.5 0.7 2 GH74 NA 1.5 0.4 0.345 0.1035 2 GH94 0.15 1.7 0.5 1 0.15 2 NY142 0.1 1.5 0.6 0.9 0.2 2 NY30 0.1 1.7 0.7 1 0.1 2 RSA5 0.05 1.5 0.5 0.8 0.15 2 US154 0.05 1.6 0.55 0.7 0.2 2 US155 0.05 1.2 0.5 0.9 0.2 2 US45 0.1 1.6 0.6 1.3 0.1 2 XAL13 0.05 1.4 0.5 0.9 0.2 2 XAL22 0.05 1.5 0.5 0.7 0.15 2 ASU10 0.1 3 1.6 2.2 0.2 1 B14 0.1 1.8 0.5 2.9 0.3 2 C19 0.2 2 0.6 2.3 0.2 2 ILL3 0.1 1.7 0.5 2.5 0.3 2 MICH1 0.1 3.5 1.3 1.8 0.2 2 MICH17 0.1 1.6 0.6 2.6 0.3 2 MICH21 0.2 2 0.7 2.2 0.2 2 MICH48 0.1 1.9 0.5 2.5 0.2 2 MO29 0.1 2 0.7 1.9 0.2 2 MO33 0.12 1.3 0.5 2.4 0.3 2

126 Appendix 3 continued.

OTU FreFilD OvL OvW StylL StylD OvlNo NY127 0.1 1.8 0.7 1.8 0.2 2 NY162 0.1 1.9 0.6 2.5 0.2 2 US144 0.1 1.8 0.6 2.7 0.2 2 US145 0.1 2 0.7 2.8 0.2 2 US147 0.1 1.5 0.4 1.7 0.25 2 A2 0.05 1.2 0.4 1 0.1 2 A6 0.1 1.3 0.6 1.2 0.1 2 POM2 0.1 1.1 0.45 0.9 0.1 2 VT21 0.1 1.3 0.5 0.8 0.1 3 XAL54 0.05 0.7 0.3 0.9 0.1 2 ASU5 0.2 2 0.5 2.1 0.2 2 DAV5 0.2 2.1 0.6 2 0.2 2 GH49 0.2 2.1 0.6 1.725 0.115 2 GH69 0.1 1.8 0.55 2.2 0.1 2 MICH50 0.1 2.1 0.5 2 0.2 2 MO24 0.2 1.6 0.6 1.6 0.2 2 MO28 0.1 2.3 0.6 1.8 0.2 2 MU2 0.1 4.5 2.1 1.8 0.2 2 NY158 0.2 1.7 0.6 1.9 0.2 2 NY172 0.1 1.7 0.6 2.5 0.2 2 RSA4 0.1 1.5 0.5 1.9 0.2 2 TEX3 0.1 3.2 1.4 1.4 0.15 2 US139 0.05 1.8 0.6 1.9 0.15 2 XAL14 0.1 2 0.5 2.3 0.2 2 XAL44 0.2 2 0.8 2.1 0.2 2

127 Appendix 4. Table of qualitative data and additional quantitative data recorded, but not used in analyses.

OTU Habit Root RootDiam Stem InNodPub NodPub StipShp StipAdPub StipAbPub StipMar StipMarPub PetPub LfltShp ARIZ34 P NA NA NA S S NT G G Sr S M Oc DAV2 P NA 5.8 Pr S S T G S E NA D Oc/Oo F28 P NA NA NA M M NT G G E S M Oc F75 P NA NA NA S D T G S E NA M Ot GH17 P W 3.3 C M D T G G Sr S M Oo GH37 P NA 2.5 NA G S NT G G E NA M Oc/Oo GH40 P NA 2 NA S S NT G G E NA D Oc/Oo GH48 P NA 10 NA G M NT G M E NA D Oc/Oo MO34 P NA NA NA S M NT G G E NA S Oc/Oo NY28 P NA NA NA S S NT G S Sr NA S Oc/Oo P76 P W 3.8 NA S S NT G G E S S Oc US109 P NA NA NA S M NT G G E NA M Oc/Oo US114 P NA 4.5 Pr S D T G M E NA D Oc/Oo US67 P NA NA NA M D NT G G E S M Oc US74 P NA NA NA S S NT G G E S S Oc US76 P NA NA NA S M NT G G E S M Oc US78 P NA NA NA S M NT G G Sr S M Oc US79 P NA NA NA S S NT G G E NA M Oc/Oo US87 P NA 5 As S S NT G S E NA S Oc/Oo US89 P NA 3 Pr S M NT G S E NA M Oc/Oo US99 P W 3.6 NA S S NT G G E S S Oo VT22 P LW 4.9 NA M M NT G G E S M Oo XAL48 P W 3 C S S T G G E S S Oo/Oc ARIZ1 P NA NA NA S M NT G S Sr S M Oo/Rh ARIZ2 P W 5.2 NA M D NT G G E S M Oc ASU1 P NA NA NA S M NT G G E NA D Tl GH75 P NA 3.5 NA S M NT G S E NA D Oc/Oo NY8 P NA 4 NA M M NT G G E NA M Oc/Oo NY9 P NA NA NA S M NT G G Sr NA M Oc/Oo US191 P NA 8 NA M D NT G G Sr NA M Oc/Oo GH105 P W 3 NA M D T S S Sr M S Oc MEXU29 P NA NA C M M T G S E M S Oc/Oo/Rh MEXU31 P NA NA NA M D T G S E D M Oo/Rh MEXU43 P W 4.4 NA D D NT G M E D M Oo/El

128 Appendix 4 continued.

abit OTU H Root RootDiam Stem InNodPub NodPub StipShp StipAdPub StipAbPub StipMar StipMarPub PetPub LfltShp NY248 P W 6.7 NA M D NT G M Sr M S Oo ARIZ16 P LW 7.5 NA S M NT G G Sr M M Rh NY1 P W 4.8 NA M M NT G G E S M Oo NY140 P W 4.3 NA M D NT G G Sr S D Oo/El P91 P W 6.1 NA M D NT G G Sr S M Oo/El POM8 P W 7.9 NA M M NT G G Sr S M El ARIZ39 P LW 10.5 C S S NT G G E S S Oo/Rh GH8 P LW 8 Pr S S T G G E G G Oo MICH92 P W 2.9 C S S NT G G E S S Oo/Rh MICH93 P LW 11.7 C S S NT G G E S S Oo MICH94 P LW 1.9 NA S S T G G E S S Ot GH1 P NA NA NA S S NT G S E NA S Oc/Oo GH74 P NA 10 C S M NT G G E NA M Oc/Oo GH94 P W 7.1 NA S S NT G G Sr S S El NY142 P NA NA NA S M NT G G E G S Ol NY30 P NA NA NA S S NT G G E S M El RSA5 P W 7.4 NA M D NT G G Sr S M El US154 P W 5.5 NA M M NT G G E S M El US155 P W 9.2 NA S M NT G G E S S El US45 P W 7 NA S S NT G G Sr S S Ol XAL13 P W 4.6 C S S NT G G E S M El XAL22 P W 3.3 NA S S NT G G E S S El ASU10 P LW 11.8 NA G G NT G G Sr S G O-El B14 P NA NA NA G G NT G G E G G O-La C19 P NA NA NA S S NT G S E S S Rh ILL3 P NA NA NA G S NT G G Sr G S O-La MICH1 P W 2.5 NA M M T G G E S S Rh/Tl MICH17 P NA NA NA G G NT G G Sr S G La MICH21 P W 6.75 NA G G NT G G E G G La MICH48 P LW 4 Dc S M NT G G E G S Oo/Rh MO29 P NA NA NA S M NT G G E G S Oo MO33 P NA NA Pr S S NT G G E G G Tl NY127 P NA NA NA M D NT G G E S D O NY162 P W 3.3 NA G G NT G G E G G Rh/El/La

129 Appendix 4 continued.

MarPub OTU Habit Root RootDiam Stem InNodPub NodPub StipShp StipAdPub StipAbPub StipMar Stip PetPub LfltShp US144 P LW 4.65 NA G G NT G G E G G El US145 P NA NA NA S S NT G G E G G Tl US147 P NA NA NA S M NT G G E G S Rh A2 P NA NA Pr M D NT G M E M D Oo A6 P W 2.8 NA M M NT G G E S M Oc POM2 P NA NA Pr D D T G M Sr M D El/Rh VT21 P LW 5.1 NA M D T S M Sr M D El XAL54 P W 1.2 NA D D NT G M E M M Oo ASU5 P W 3.5 R M D NT G G E S D Rh/Ot DAV5 P W 9.2 As S D NT G G E G D Oo/Rh GH49 P NA NA NA M D T G G E NA D Tl GH69 P W 6.4 NA S D NT G S E S M El MICH50 P LW 5 Pr S D T G G E M S Oo/Rh MO24 P NA NA Pr M D NT G G E S M Rh/Ot MO28 P NA NA NA M D NT G S Sr NA S Rh MU2 P LW 7.5 NA M D T G S Sr NA D Tl NY158 P W 4.6 R M D T G G Sr S D Oo/Rh NY172 P W 4.2 NA M M NT G S E S M Rh/Tl RSA4 P NA NA NA M D T G G E S D Rh TEX3 P NA NA NA M D T G S Sr NA D Rh US139 P NA NA NA M D NT G G Sr S D El XAL14 P NA NA NA M D NT G S Sr S D Oo XAL44 P NA NA NA M D NT G M Sr M M Rh-Oc

130 Appendix 4 continued.

fl OTU LfltAdPub LfltAbMdrbPub LfltAbPub LfltMar LfltMarPub LfltApxAng LfltMucr PetllePub In Bract Whorl PeduncPub PedicPub SepTubPub SepLobShp ARIZ34 G M S Sr S Oc Y S RU Lt 2 S S S NT DAV2 G S S Sr NA 180 Y M U Lt 1 D D D NT F28 G M S Ssr S Oc Y D CU Lt 1 D D S NT F75 G D M Ssr NA 90 Y D RU Lt 2 D M M NT GH17 G M M Sr S 120 Y D RU Lt 2 M S M NT GH37 G D M Sr G Oc N M U Lt 1 M M D NT GH40 G M G Sr NA 120 Y D U Lt 1 M D D NT GH48 G D S Ssr NA Oc Y S U Lt 2 D D M NT MO34 G S S Sr NA Oc N S U Lt 1 M M M NT NY28 G D S Ssr NA Oc Y M U Lt 1 D D D NT P76 G S S Ssr S Oc Y S CU Lt 1 D D S NT US109 G S S Sr NA Oc Y M U Lt 1 M D D NT US114 G M M Sr NA Oc Y D RU Lt 3 M S M NT US67 G M S Ssr S Oc Y D RU Lt 2 D M M NT US74 G S S Ssr S Oc Y M CU Lt 1 M D S NT US76 G M S Ssr S Oc Y S CU Lt 1 M D S NT US78 G M S Ssr S Oc Y M CU Lt 1 D D D NT US79 G S S Sr NA Oc Y D RU Lt 2 M D NA NT US87 G S G Sr NA Oc N S U Lt 2 D M M NT US89 G D G Sr NA Oc Y M U Lt 1 D D D NT US99 G S G Sr S 120 Y S RU Lt 2 S S G NT VT22 G M S Sr S 60 Y M RU Lt 2 M S S NT XAL48 G S S Sr S 105 Y S RU Lt 2 M S S NT ARIZ1 G M S Ssr S 75 Y D RU Lt 2 M M M NT ARIZ2 G S G Ssr S Oc Y D CU Lt 1 M S M NT ASU1 G S S Sr NA 105 Y S U Lt 1 M M M NT GH75 G S S Ssr NA 125 Y M U Lt 2 D M S NT NY8 G M G Sr NA 120 Y S U Lt 1 M M D NT NY9 G M G Sr NA 120 Y D RU Lt 2 M S M NT US191 G M G Sr NA Oc Y D U Lt 1 M D D NT GH105 G M M Ssr M Oc Y M RU Lt 2 M M M NT MEXU29 G M M Sr M Oc Y S RU Lt 2 M S D NT MEXU31 G M M Sr S 75 Y D CU Lt 1 M S D NT MEXU43 G D D Sr M 90 Y D RU Lt 2 D S S NT

131 Appendix 4 continued.

OTU LfltAdPub LfltAbMdrbPub LfltAbPub LfltMar LfltMarPub LfltApxAng LfltMucr PetllePub Infl Bract Whorl PeduncPub PedicPub SepTubPub SepLobShp NY248 G M M Ssr M 105 Y M RU Lt 2 D M D NT ARIZ16 G M S Ssr M 60 Y M RU Lt 3 M S S NT NY1 G M M Sr M 45 Y D CU Lt 1 M M D NT NY140 G M S Ssr S 60 Y D RU Bf 2 D S M NT P91 G S S Ssr S 60 Y S RU Lt 2 M S M NT POM8 G M S Ssr S 55 Y S CU Lt 1 M M M NT ARIZ39 G S S Ssr S 75 Y S RU Lt 4 M S G NT GH8 G S S Ssr S 75 Y S RU Lt 3 M S G NT MICH92 G S G Ssr S 90 Y S RU Lt 2 M S G NT MICH93 G S S Ssr S 75 Y S RU Lt 2 M S G NT MICH94 G S S Ssr S 90 Y S RU Lt 2 M S S NT GH1 G S S Sr NA 45 Y S U Lt 3 S D D NT GH74 G D S Ssr NA 90 N M U Lt 3 D S NA NT GH94 G S S Ssr S 45 Y M RU Bf 2 M S S NT NY142 G S G Sr G 75 Y M RU Lt 2 S S S NT NY30 G M S Ssr S 60 Y M RU Lt 3 M S S Lt RSA5 G S S Ssr S 75 Y M RU Lt 2 M M S NT US154 G M S Sr S Ro Y S RU Lt 3 M S S NT US155 G M S Ssr S 35 Y M RU Lt 3 M S S NT US45 G S S Ssr S 60 Y M RU Lt 2 M S G NT XAL13 G M S Ssr S 35 Y M CU Lt 1 M S S NT XAL22 G S S Ssr S Ro Y S CU Lt 1 M M S NT ASU10 G G G Ssr G 40 Y G RU Lt 4 G G G NT B14 G S G Ssr G 55 Y G RU Lt 4 S S S NT C19 G S S Ssr S 75 Y M RU Lt 3 M D S NT ILL3 G S S Ssr G 45 Y S RU Lt 4 S S S NT MICH1 G S S Ssr S 55 Y M RU Lt 3 M S S NT MICH17 G G G Ssr G 40 Y M RU Lt 5 S S S NT MICH21 G G G Ssr G 30 Y S RU Lt 3 M S G NT MICH48 G S S Ssr S 80 Y S RU Lt 3 M S G NT MO29 G S S Ssr G 60 Y M RU Lt 2 M S S NT MO33 G S G Ssr G 45 Y S RU Lt 3 M S S NT NY127 G M S Ssr S 75 Y D RU Lt 5 M S S NT NY162 G G G Ssr G 120 Y G RU Lt 3 S G G NT

132 Appendix 4 continued.

OTU LfltAdPub LfltAbMdrbPub LfltAbPub LfltMar LfltMarPub LfltApxAng LfltMucr PetllePub Infl Bract Whorl PeduncPub PedicPub SepTubPub SepLobShp US144 G S S Ssr S 50 Y G RU Lt 4 S S S NT US145 G S S Sr G 105 Y G RU Lt 3 M G G NT US147 G S S Ssr S 60 Y M RU Lt 2 M S G NT A2 G M M Sr M 60 Y D RU Lt NA M D D NT A6 G S S Sr S Oc Y M RU Lt 3 M S M NT POM2 G D M Ssr M 75 Y D RU Lt 3 D S D NT VT21 G M M Sr M 75 Y D RU Lt 3 M M S NT XAL54 G M M Sr M 75 Y D CU Lt 1 M D D NT ASU5 G M S Ssr S 80 Y S RU Lt 2 D M M NT DAV5 G M S Ssr S 75 Y D RU Lt 2 D S S NT GH49 G S G Ssr NA 74 Y G U NA NA D S NA Lt GH69 G M S Ssr S 45 Y D RU Lt 5 M S M NT MICH50 G M S Ssr S 100 Y M RU Lt 3 M S S NT MO24 G S S Ssr S 90 Y M RU Lt 4 M S S NT MO28 S M S Ssr NA 105 Y D RU Lt 2 D M M NT MU2 G M S Ssr NA 105 Y D U Lt 3 D M NA NT NY158 G M S Ssr S 90 Y D RU Lt 3 D S G NT NY172 G S M Ssr S 75 Y S RU Lt 4 M S S NT RSA4 G M S Ssr S 90 Y D RU Lt 4 D S S NT TEX3 G M S Ssr NA 75 Y D U Lt 3 D M NA Lt US139 G M S Ssr S 55 Y D RU Lt 4 D M S NT XAL14 G M S Ssr S Ro Y D RU Lt 4 D M M NT XAL44 G S S Ssr S Oc Y M RU Lt 3 M S S NT

133 Appendix 4 continued.

etLamMar OTU SepLobPub SepLobMar SepLobMarPub PetCol BPetShp WPetLam WPetLamBas WP WPetLamApx KPetLam KPetLamBas KPetLamMar KPetLamApx OvPub ARIZ34 S E S NA Oo Sp SA E Ro Sp Tr E Ro S DAV2 D E M Pk NA Sp SA E Ro Sp SA E Ro D F28 D E M NA Oo Sp SA E Ro Sp Tr E Ro S F75 M E M NA NA Sp Tr E Ro Sp Tr E Ro M GH17 M E M Pk Oo Sp SA E Ro Sp Tr E Ro S GH37 D E M NA NA Sp SA E Ro Sp SA E Ro G GH40 NA E D NA NA Sp SA E Ro Sp SA E Ro D GH48 M E M NA NA Sp SA E Ro Sp Tr E Ro M MO34 NA E M NA NA Sp SA E Ro Sp Tr E Ro S NY28 D E D NA NA Sp SA E Ro Sp Tr E Ro NA P76 S E D NA Oo Sp SA E Ro Sp Tr E Ro M US109 D E D NA NA Sp SA E Ro Sp Tr E Ro S US114 NA E S Pk NA Sp SA E Ro Sp Tr E Ro M US67 M E D NA Oo Sp SA E Ro Sp Tr E Ro S US74 S E D NA Oo Sp SA E Ro Sp Tr E Ro M US76 M E D NA Oo Sp SA E Ro Sp Tr E Ro M US78 S E D NA Oo Sp SA E Ro Sp Tr E Ro M US79 D E D NA NA Sp SA E Ro Sp Tr E Ro S US87 NA E D Pu NA Sp SA E Ro Sp Tr E Ro G US89 D E D Pu NA Sp SA E Ro Sp Tr E Ro D US99 G E S NA Oo Sp SA E Ro Sp Tr E Ro S VT22 M E M NA Oo Sp SA E Ro Sp Tr E Ro M XAL48 S E S Pu El/Oc Sp SA E Ro Sp Tr E Ro S ARIZ1 M Sr M NA Oo Sp SA E Ro Sp Tr E Ro D ARIZ2 M E M NA Oo Sp SA E Ro Sp Tr E Ro D ASU1 M E M Pk NA Sp SA E Ro Sp Tr E Ro M GH75 S E S Wh NA Sp SA E Ro Sp Tr E Ro D NY8 D E D NA NA Sp SA E Ro Sp Tr E Ro D NY9 NA E S Wh NA Sp SA E Ro Sp Tr E Ro M US191 NA E M NA NA Sp SA E Ro Sp Tr E Ro D GH105 S E M Pk El/Oc Sp Tr E Ro Sp Tr E Ro D MEXU29 D E D Pu El/Oc Sp Tr E Ro Sp Tr E Ro D MEXU31 M E D Wh El Sp SA E Ro Sp Tr E Ro D MEXU43 M E M Pk El/Oc Sp SA E Ro Sp Tr E Ro D

134 Appendix 4 continued.

OTU SepLobPub SepLobMar SepLobMarPub PetCol BPetShp WPetLam WPetLamBas WPetLamMar WPetLamApx KPetLam KPetLamBas KPetLamMar KPetLamApx OvPub NY248 M E D Wh El Sp SA E Ro Sp Tr E Ro D ARIZ16 M Sr M NA Oo Sp SA E Ro Sp Tr E Ro S NY1 D Sr D Yw Oo Sp SA E Ro Sp Tr E Ro M NY140 M Sr M NA Oo Sp Tr E Ro Sp Tr E Ro D P91 S Sr S NA Oo Sp SA E Ro Sp Tr E Ro D POM8 M Sr M NA Oo Sp Tr E Ro Sp Tr E Ro M ARIZ39 S E S Pu Oo Sp SA E Ro Sp Tr E Ro G GH8 S E S NA Oo Sp SA E Ro Sp Tr E Ro S MICH92 G E S Pu Oo Sp SA E Ro Sp Tr E Ro M MICH93 S E S Pu Oo Sp SA E Ro Sp Tr E Ro S MICH94 S E M Pu Oo Sp SA E Ro Sp Tr E Ro S GH1 NA E M NA NA Sp SA E Ro Sp Tr E Ro S GH74 M E S Wh NA Sp Tr E Ro Sp Tr E Ro S GH94 S E S NA Oo Sp SA E Ro Sp Tr E Ro S NY142 S E S NA Oo Sp SA E Ro Sp Tr E Ro S NY30 M E S NA Oo Sp SA E Ro Sp Tr E Ro S RSA5 S E S NA Oo Sp Tr E Ro Sp Tr E Ro D US154 M E S NA Oo Sp Tr E Ro Sp Tr E Ro M US155 M E M NA Oo Sp Tr E Ro Sp Tr E Ro S US45 M E S NA Oo Sp SA E Ro Sp Tr E Ro S XAL13 M E S Wh Oo Sp SA E Ro Sp Tr E Ro S XAL22 M E S NA Oo Sp SA E Ro Sp Tr E Ro M ASU10 G E S Wh Oo Sp SA E Ro Sp Tr E Ro G B14 S E S NA Oo Sp SA E Ro Sp Tr E Ro G C19 S E S NA Oo Sp Tr E Ro Sp Tr E Ro G ILL3 S E S NA Oo Sp SA E Ro Sp Tr E Ro G MICH1 S E S Wh Oo Sp Tr E Ro Sp Tr E Ro G MICH17 S E S Cr Oo Sp SA E Ro Sp Tr E Ro G MICH21 S E S NA Oo Sp SA E Ro Sp Tr E Ro G MICH48 S E S Yw Oo Sp SA E Ro Sp Tr E Ro G MO29 S E S Wh Oo Sp Tr E Ro Sp Tr E Ro G MO33 NA E S NA NA Sp SA E Ro Sp Tr E Ro G NY127 S E S Pk Oo Sp Tr E Ro Sp Tr E Ro G NY162 G E M NA Oo Sp SA E Ro Sp Tr E Ro G

135 Appendix 4 continued.

OTU SepLobPub SepLobMar SepLobMarPub PetCol BPetShp WPetLam WPetLamBas WPetLamMar WPetLamApx KPetLam KPetLamBas KPetLamMar KPetLamApx OvPub US144 S E S Wh Oo Sp Tr E Ro Sp Tr E Ro G US145 G E G Wh NA Sp SA E Ro Sp Tr E Ro G US147 G E S NA Oo Sp Tr E Ro Sp Tr E Ro G A2 M E D Pk Oo Sp Tr E Ro Sp Tr E Ro S A6 M E D NA Oo Sp Tr E Ro Sp Tr E Ro M POM2 D E D Pk Oo Sp Tr E Ro Sp Tr E Ro G VT21 M E M NA Oo Sp SA E Ro Sp Tr E Ro D XAL54 D E D Pk Oo Sp SA E Ro Sp Tr E Ro S ASU5 S E M Wh Oo Sp SA E Ro Sp Tr E Ro S DAV5 S E S NA Oo Sp SA E Ro Sp Tr E Ro M GH49 M E S NA NA Sp SA E Ro Sp Tr E Ro M GH69 M E S NA Oo Sp SA E Ro Sp Tr E Ro S MICH50 S E S Wh Oo Sp SA E Ro Sp Tr E Ro G MO24 S E S Wh Oo Sp SA E Ro Sp Tr E Ro S MO28 NA E M Wh NA Sp SA E Ro Sp Tr E Ro S MU2 S E M NA NA Sp SA E Ro Sp Tr E Ro S NY158 M E M Wh Oo Sp SA E Ro Sp Tr E Ro D NY172 S E S Wh Oo Sp SA E Ro Sp Tr E Ro S RSA4 D E M Wh Oo Sp SA E Ro Sp Tr E Ro D TEX3 M E M Cr NA Sp SA E Ro Sp Tr E Ro M US139 M E M NA Oo Sp SA E Ro Sp Tr E Ro M XAL14 S E S Wh Oo Sp SA E Ro Sp Tr E Ro D XAL44 M E S NA Oo Sp SA E Ro Sp Tr E Ro S

136 Appendix 4 continued.

OTU StylPub SeedShp SeedCol ARIZ34 G NA NA DAV2 G NA NA F28 G Gb RB F75 G NA Cu GH17 G NA NA GH37 G NA NA GH40 G NA Cu GH48 G NA NA MO34 G NA NA NY28 G NA NA P76 G Mt BO* US109 G NA NA US114 G NA NA US67 G Gb BO* US74 G Gb RB US76 G Mt RB US78 G NA NA US79 G NA NA US87 G NA RT* US89 G NA NA US99 G NA NA VT22 G NA NA XAL48 G NA NA ARIZ1 G NA NA ARIZ2 G Gb OB ASU1 G NA NA GH75 G NA NA NY8 G NA BO NY9 G NA NA US191 G NA RT* GH105 G Mt OB MEXU29 G Gb BO MEXU31 G Gb LB MEXU43 G Mt BY

137 Appendix 4 continued.

OTU StylPub SeedShp SeedCol NY248 G Mt LB* ARIZ16 G Gb BO* NY1 G NA NA NY140 G Mt DB P91 G NA NA POM8 G Mt DB ARIZ39 G NA NA GH8 G NA NA MICH92 S NA NA MICH93 G Mt RB* MICH94 G Mt BO* GH1 G NA RB GH74 G NA RB GH94 G Mt DB NY142 G NA NA NY30 G Mt OB* RSA5 G Mt Br US154 G Gb Br* US155 G Gb BO* US45 G NA NA XAL13 G Mt Bl XAL22 G Mt RB ASU10 G NA NA B14 G NA NA C19 G NA NA ILL3 G Mt LB MICH1 G Mt GY* MICH17 G NA NA MICH21 G NA NA MICH48 G NA NA MO29 G NA NA MO33 G NA Cu NY127 G NA NA NY162 G Mt DB

138 Appendix 4 continued.

OTU StylPub SeedShp SeedCol US144 G Mt OB US145 G NA NA US147 G NA NA A2 G NA NA A6 G NA NA POM2 G NA NA VT21 G Mt Br XAL54 G NA NA ASU5 G NA NA DAV5 G Mt GY GH49 G NA NA GH69 G Gb BO* MICH50 G Mt DB MO24 G NA NA MO28 G NA NA MU2 G NA NA NY158 G NA NA NY172 G NA NA RSA4 G NA NA TEX3 G NA NA US139 G NA NA XAL14 G NA NA XAL44 G NA NA Habit: P--perennial, A--annual; Root: L--large, W--woody; Stem: R--repent, C-- creeping, As--ascending, Pr--prostrate, Dc--decumbent; Pubescence: G-- glabrous, S--sparse, M--moderate, D--dense; Margin: E--entire, Sr--serrate, Ssr--serrate, often with alternating short, straight and larger, antrorsely curved teeth; Presence/Absence characters: Y--present, N--absent; Inflorescence: U-- umbel, RU--racemose umbel, CU--capitate umbel; Shapes: Oo--obovate, Oc-- obcordate, Ot--obtrullate, El--elliptic, Ro--rounded, Lt--linear triangular, Rh-- rhombic, Tl--trullate, Bf--bifid, Sp--spatulate, Gb--globular, Mt--mitten, Tr-- truncate, SA-- shortly auriculate, O--Ovate, La--lanceolate, Ol--oblanceolate, T- -triangular, NT--narrowly triangular; Colors: Pk--pink, Wh--white, Pu--purple, Yw--yellow, Cr--cream, Bl--blond, Br--brown, BO--brownish orange, BY--

139 brownish yellow, Cu--copper, DB--dark brown, GY--grayish yellow, LB--light brown, OB--olive brown, RB--reddish brown, RT-- reddish tan, *--dark speckled

140 Appendix 5. Results from PCA and SAHN clustering analyses from preliminary small group analyses in NTSYSpc. Groups are represented by different colors and symbols.

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152 Appendix 6. Final PCA results showing the final species, both 2D and 3D plots are presented.

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