Abstract a Morphological Analysis Of

ABSTRACT

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

by Justin Hendy

The Trifolium amabile Kunth species complex (Fabaceae) is a group of New World clovers found in mountainous regions ranging from southern Arizona in the United Sates, through Central America, where it occurs in Mexico, Guatemala, and Costa Rica, and south along the Andes Mountains, from Colombia, Ecuador, Peru, and Bolivia, to Argentina. Zohary and Heller (1984) circumscribe this group as a single species, T. amabile, with five varieties. Ahlquist (2012) examined material from North and Central America, and recognized nine species in that geographical range. This study focuses on the South American range of the species complex. Species boundaries were determined using morphometric analyses of herbarium specimens, including PCA and cluster analysis, using NTSYS-pc, Student’s t-tests, and box and whisker plots. Six taxa, including four new species, are recognized as distinct and described. These species include: Trifolium amabile, T. andesicola sp. nov., T. australe sp. nov., T. cochabambense sp. nov., T. peruvianum, and T. tafiense sp. nov.

A MORPHOLOGICAL ANALYSIS OF THE TRIFOLIUM AMABILE KUNTH SPECIES COMPLEX IN SOUTH 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

Justin Hendy

Miami University

Oxford, Ohio

2013

Advisor ______

Michael A. Vincent

Reader______

Richard C. Moore

Reader______

M. Henry H. Stevens

Table of Contents Introduction……………………………………………………………………….………………………………………………………1 Materials and Methods………………………………………………………………………………………………………………6 Results……………………….………………………………………………………………………………………………………………8 Discussion…………………………………………………………………………………..…………..……………………………….26 Taxonomic Treatment…………………………………………………………………………………..…………………………27 Conclusion……………………………………………………………………………………………………………………………….59 Literature Cited………………………………………………………………………………..………………………………………61 Appendix 1……………………………………………………………………………………………………………..……………….63 Appendix 2………………………………………………………………………………………………………..…………………….66 Appendix 3…………………………………………………………………………………………………………………..………….71 Appendix 4………………………………………………………………………………………………………………..…………….83 Appendix 5…………………………………………………………………………………………………………………………….107

iii

List of Tables 1. List of published names………………………………………………………………………………………………………….5 2. P-values resulting from student’s t-tests of each species against each other species…………..13

List of Figures 1. PCA of all 109 specimens…………………………………………………………………………………………………..…10 2. Dendrogram of all 109 specimens………………………………………………………………………..…….………..11 3. Box and whisker plot of characters used for the first couplet……………………………………….………14 4. Box and whisker plot of characters used for the second couplet………………………………….………17 5. Box and whisker plot of the characters used for the third couplet……………………………………....19 6. Box and whisker plot of the characters used for the fourth couplet…………………………………….21 7. Box and whisker plot of the characters used for the fifth couplet…………………………………….….23 8. Trifolium amabile……………………………………………………………………………………………………….………..33 9. Geographical range of T. amabile………………………………………………………………………………….……..34 10. Trifolium peruvianum……………………………….…………………………………………………………….………….39 11. Geographical range of T. peruvianum………………………………………………………………………..……….40 12. Trifolium tafiense………………………………………………………………………………………………………..……..43 13. Geographical range of T. tafiense……………………………………………………………………………….……..44 14. Trifolium andesicola…………………………………………………………………………………………..………………48 15. Geographical range of T. andesicola…………………………………………………………………………..………49 16. Trifolium australe……………………………………………………………………………………………………………...53 17. Geographical range of T. australe…………………………………………………………………………..………….54 18. Trifolium cochabambense…………………………………………………………………………………………….…….57 19. Geographical range of T. cochabambense………………………………………………………………………….58

Acknowledgements

First and foremost, I would like to thank my wife for all of her encouragement and support. Without her and my family none of this would have been possible. I would also like to thank my advisor, Dr. Mike Vincent, without whom I would have not accomplished something so great. I would also like to thank my thesis committee members, Drs. Hickey, Moore, and Stevens for all of their advice and help. Finally I would like to thank all of the other graduate students and my lab-mates for offering support, advice and keeping things fun.

Introduction

The genus Trifolium L. (clovers; Fabaceae, Faboideae, Trifolieae) consists of approximately 250 species (Ellison et al. 2006, Zohary & Heller 1984). The name Trifolium derives from tri, three, and folium, leaf, referring to the trifoliate leaf found commonly in the genus. Trifolium species, which are annual, biennial, or perennial herbs, occur in temperate regions, with a few species in subtropical areas. The range of the genus is North America, Central America, and South America, Europe, Africa, and temperate Asia; no native clovers are found in tropical Asia, Australia, or the Pacific Ocean islands. Three major centers for speciation in the genus are the Mediterranean region, the California region, and the eastern African highlands (Zohary & Heller 1984).

Within Fabaceae, Trifolium is placed in the tribe Trifolieae. Lewis et al. (2005) defined this tribe as consisting of six closely related genera: Ononis, Parochetus, Trigonella, Medicago, Melilotus, and Trifolium. Trigonella, Medicago, Parochetus, and Melilotus are Old World genera, but some species have been cultivated and have spread from cultivation outside their native ranges. Ononis (another Old World genus) can be distinguished from Trifolium by its alternately dorsifixed and basifixed dimorphic anthers and its usually glandular and spiny nature (Wei & Vincent 2010). Trifolium can be separated from Parochetus, Melilotus, Medicago, and Trigonella by its persistent corolla after anthesis, a legume that is included in or slightly exerted from the persistent corolla, and filaments that are dilated distally (Gleason & Cronquist 1991, Wei & Vincent 2010).

Clovers are important for livestock feed, and at least 16 species are actively cultivated (Ellison et al. 2006). Trifolium repens L., white clover, and T. pratense L., red clover, are the most widely cultivated. Clovers have been cultivated for several reasons. They produce a plentiful crop that will continue to grow after heavy grazing; they are nutritious for livestock; and they can grow in many climates due to their cold hardiness; they grow in a wide range of soil types (Taylor 1985). Due to these favorable properties T. pratense (red clover) has been cultivated since the 4th century A.D. in Europe (Zohary & Heller 1984).

Pre-Linnean botanists made preliminary attempts to classify Trifolium into subgeneric groups, but were only aware of a very small number of species. Linnaeus (1753) divided his 41 species of Trifolium into five units; Meliloti, Lotoidea, Lagopoda, Vesicaria, and Lupulina (Zohary & Heller 1984). Not all of his species are now placed in Trifolium, with some instead assigned to Trigonella and Melilotus. After Linnaeus, many scientists studied clovers in their own regions, but no world-wide studies were conducted until Seringe (1825) completed the first revisions of the entire genus, including 150 species from both the Old and New Worlds. Presl (1831) split Trifolium into nine genera: Paramesus, Amarenus, Lupinaster, Amoria, Micrantheum, Trifolium, Mistyllus, Galearia, and Calycomorphum. His work was not followed by others; however, most of his genera were later accepted as sections within Trifolium. Celakovsky (1874) examined the sections of Trifolium and attempted to describe their phyletic relationships. The last major work on the entirety of Trifolium before Zohary and Heller was that of Bobrov (1967). He organized eleven sections of Trifolium into two tribes, Trifolieae and Lupineae.

In the most recent monograph, Zohary and Heller (1984) recognized 237 species, and arranged them into eight sections: Lotiodea, Paramesus, Mistyllus, Vesicaria, Chronosemium, Trifolium, Trichocephalum, and Involucrarium. The New World species were included within Sections Lotoidea (containing both New and Old World species) and Involucrarium (containing only New World species).

Ellison et al. (2006) conducted a molecular phylogenetic analysis of Trifolium, including 255 species covering the geographical and morphological range of the genus. As a result of these analyses, they recognized two subgenera, Chronosemium and Trifolium, with the latter encompassing eight sections. Ellison et al. (2006) place all of New World Trifolum in a newly defined Section Involucrarium of subgenus Trifolium. Trifolium breweri S. Wats. (a Californian species) is basal within the Involucrarium clade, with T. amabile Kunth + T. peruvianum Vogel as sister to the remaining 65 species (Ellison et al. 2006).

Trifolium amabile was described by Kunth (in Humboldt et al. 1824) from material from Mexico collected by Humboldt and Bonpland. As circumscribed by Zohary and Heller (1984), T. amabile is a highly variable taxon, in which five varieties were recognized; T. amabile Kunth var.

2 amabile, T. amabile Kunth var. hemsleyi (Lojac.),Heller & Zoh., T. amabile Kunth var. pentlandii Ball, T. amabile var. mexicanum (Hemsl.) Heller & Zoh., T. amabile Kunth var. longifoliolum Hemsl., and which ranges from Arizona in the north to the Andes in the south including the countries of Ecuador, Argentina, Peru, and Bolivia. Zohary and Heller state that var. amabile is the most common and wide-ranging variety in the species, distinguishable from the rest of the varieties by its narrowly ovate leaflets and an inflorescence up to 1 cm. The large amount of variability within T. amabile, when delimited in this broad fashion, caused McVaugh (1987) to remark that, “In this species there is extensive variation, especially in the size of the leaves and the size of the flower heads”.

This study focuses on the South American range of the T. amabile species complex, since material from the North American and Central American part of the range was reexamined by Ahlquist (2012). The range studied by Ahlquist (2012) is geographically isolated from the South American range that I am to study, since there is a natural elevational split between populations found in Costa Rica and those in Columbia. The choice of splitting the species complex into two portions is recognized as potentially arbitrary, and the fact that there is a geographical break does not mean that none of the North American taxa may be found in South America because the current range could be a result of vicariance or long distance dispersal. In her study of the northern part of the range, Ahlquist (2012) split the T. amabile species complex into nine separate species.

Twelve names have been published for entities in the South American range T. amabile s.l (Table 1). The names that pertain to this study because they have a South American range include: T. amabile Kunth, T. humboldtii Spreng, T. mathewsii A. Gray, T. bolivianum Kennedy, T. hemsleyi (Lojac.) Heller & Zoh., T. amabile Kunth var. hemsleyi Lojac., T. macrorrhizum Ulbrich, T. amabile Kunth var. pentlandii Ball, T. chiclense Ball, T. peruvianum Vog., T. peruvianum var. chiclense (Ball) Macbr., and T. weberbaueri Ulbrich. The South American range follows the Andean mountain range and includes Columbia, Ecuador, Peru, Bolivia, and Argentina. Zohary and Heller (1984) include a South American range for T. hemslyi even though its type is from Mexico and due to this it is included in this study. I will integrate my results with those of

Ahlquist (2012) in order to make certain that a complete understanding of the taxonomy and nomenclature of the entire species complex is achieved.

Hypothesis

The hypothesis to be tested in this study is that Trifolium amabile within its South American range represents a single morphological species. If this hypothesis is rejected, I will delimit subset taxa. Subset taxa will be delimited based on the morphological cluster species concept in which taxa are distinguished as clusters separated by gaps (Tepe 2011, Mallet 1995).

The objectives of this study are:

1. To use statistical analyses of morphological data to test the hypothesis that the South American concept of T. amabile as delimited by Zohary and Heller (1984) represent a single morphological species;

2. If the hypothesis is rejected, to determine the morphological boundaries of subset taxa identified in the analyses and to determine the geographical ranges of these taxa;

3. To revise the nomenclature of the T. amabile species complex in South America; to integrate my results with those of Ahlquist’s study of the North and Central American specimens; to correlate existing names with newly delimited groups, if any; and to describe and formally name any group for which a current name is unavailable.

Table 1. Names published in Trifolium amabile species complex. Names in bold pertain to this study.

Material and Methods

Morphological characters of T. amabile were measured from herbarium specimens, following the methods described and illustrated by Ahlquist (2012). Herbarium specimens (operational taxonomic units [OTU’s]) measured were obtained from the following herbaria (acronyms from Thiers [continuously updated]): AAU, B, C, COLO, DAO, E, F, K, LPB, MICH, MO, MU, NY, P, POM, SI, US. All herbarium specimens were examined before any measurements were made, in order to discern the geographical range and the breadth of morphological variation within the OTU’s. One hundred and nine specimens were chosen from the full set of specimens to represent the full range of morphological variation and the entire geographical range. A full suite of vegetative and floral characters were measured; of these, 27 characters were used in analyses because they were informative and the rest were eliminated because they were hyper-variable or invariant, following Ahlquist (2012) (Appendix 2). For each OTU, three to five measurements were made for each character when possible, and the averages were used for analyses. If a particular OTU had limited material of a certain character, such as flowers, fewer measurements were used to derive the average. A complete list of specimens measured is included in Appendix 1.

The various morphological characters were measured using a micro-millimeter caliper, with a microscope with an ocular micrometer, and a millimeter ruler. Angle measurements were made using a clear protractor. Flowers and fruits were measured by rehydrating them in a 1% dilution of Aerosol OT in distilled water. A stock solution of 10% Aerosol OT, 20% ethanol, and 70% distilled water was used to make the dilution. Quantitative data used in the analyses are listed in Appendix 3. Qualitative and quantitative data not used in the analyses are listed in Appendix 4.

Statistical analyses of the averages were conducted using Numerical Taxonomic System (NTSYS-pc) software (Rohlf 2000). The data were standardized and a similarity matrix of interval data was generated using Correlation Coefficient (CORR). Principal Components Analysis (PCA) was carried out, and the results were visualized in three dimensional plots formed by utilizing the first three principal components. Further similarity matrices were

6 derived for OTUs using Correlation Coefficient, followed by clustering analyses using a sequential agglomerative, hierarchal, non-overlapping (SAHN) clustering analysis, and results were visualized in the form of a dendrogram using the TREE plot function. Multiple iterative runs of these analyses were done using NTSYS-pc with different characters to refine the groups. Once groups were developed, two-tailed Student’s t-tests were conducted for pairings of each of the putative species groups, and box and whisker plots were generated using Microsoft Excel 2010. T-tests were run as homoscedastic, assuming normal distributions of morphological characters.

The results of these analyses were used to further refine the groups and to analyze how consistently each held together when compared to each other group, using pairwise comparisons of the characters of the delimited groups. Some groups that had no or few differences were merged to form larger groups. With every move of specimens to new groups, all analyses were run again. This was repeated until each resultant group had significant differences from each other group and key characters could be discerned that separated the groups. Species will be delimited based similarities within clusters of OTUs and gaps between the clusters (Duley & Vincent 2003, Tepe 2011, Mallet 1995).

PCA figures and dendrogram were generated using NTSYS-pc (Rohlf, 2000). Maps of geographical ranges were created using ArcMap 10.2 (ESRI 2013).

Results Twenty-seven key characters were revealed as important in distinguishing differences within the T. amabile species complex, and using these characters, a dendrogram and PCA plot were generated using NTSYS-pc. The three dimensional plot resulting from the PCA of all 109 specimens is very cluttered, and made it difficult to discern any patterns that could be useful (Fig. 1). Focus was put on the dendrogram which showed three clear branches (Fig. 2). The top branch, A, was characterized by generally wider sepal lobes with shorter, wider banner petals. The bottom two branches, B and C, were both characterized by generally narrower sepal lobes with longer, narrower banner petals. The bottom two branches showed no significant differences in those three characters from each other and were analyzed together as one larger group. PCAs and SAHN clustering lead to the designation of four smaller subset groups within the top branch and two smaller subset groups within the bottom branches; specimens designated with black have banner petals >9mm wide, keel petal laminas ≥1.5mm wide at narrowest point, and wing petal laminas ≥1.6mm wide at narrowest point; the specimen designated with orange has pedicels <.6mm long, mature flowers not reflexing, inflorescences <7mm long X <8 mm wide, keel petal claws ≤1.3mm long, and wing petal claws ≤1.15mm long; specimens designated with green have stipules ≤6.8mm long, and terminal leaflets ≤5.7mm long X ≤3.75mm wide; specimens designated with brown have inflorescences ≥16 flowered, sepal circumferences ≤3.7mm, sepal lobes ≤.5mm wide, and wing petal claws ≤1.5mm long; specimens designated with purple have banner petal length/width ratios ≤1.2mm, keel petals ≤5.3mm long, peduncle length/petiole length ratios ≤1.3, sepal lobes ≥.7mm wide, and wing petals ≤6mm long; specimens designated with blue have banner petal length/width ratios >1.2mm, keel petals >5.3mm long; peduncle length/petiole length ratios >1.3, sepal lobes <.7mm wide, and wing petals <6mm long. With these six groups in mind, the same SAHN clustering can be used to show six subset groups (Fig. 2). Each of these six groups was run against each other group in both PCA and SAHN clustering. The PCA results from pair-wise comparisons show clear distinctions amongst all groups with limited overlap (Appendix 5). The SAHN clustering results show more

8 overlapping of specimens when compared with the PCA but still confirms six relatively clear groups. For each of the groups tentatively identified using PCA and SAHN clustering, t-tests confirmed statistically significant differences amongst the groups using the 27 characters. Table 2 gives the results for pair-wise comparisons of the six groups using the 27 characters. Box and whisker plots also showed pair-wise differences among the six groups using the 27 characters. These box and whisker plots were used to help construct the key. The t-tests and box and whisker plots support the PCA and SAHN clustering results, leading to the conclusion that these six groups are in fact distinct.

Figure 1a. Three dimensional PCA plot derived from analysis of 27 characters for all 109 specimens. Colors correspond with morphological groups as described in the text.

Figure 1b. Three dimensional PCA plot derived from analysis of 27 characters for all 109 specimens. Four individuals marked in black plotted higher in the third dimension than the rest of the specimens.

Figure 2a. Results (upper third of dendrogram – branch A) from SAHN clustering of 27 characters for all specimens used in the analysis. Colors to the right of the numbers correspond with morphological groups as described in the text.

Figure 2b. Results (lower two-thirds of dendrogram – branches B & C) from SAHN clustering of 27 characters for all specimens used in the analysis. Colors to the right of the numbers correspond with morphological groups as described in the text.

Table 2. P values from student’s t-tests of each species against each other species showing the 27 characters used in PCA and SAHN clustering analysis. Statistically significant results are highlighted. α=0.05.

T. peruvianum-T. T. tafiense peruvianum-T. T. andesicola peruvianum-T. T. australe peruvianum-T. T. amabile tafiense-T.andesicola T. tafiense-T.australe T. tafiense-T.amabile T. australeandesicola-T. T. amabile andesicola-T. T. amabile australe-T. T. StipL* 0.463159 1.7E-05 0.072909 0.139964 2.43E-05 0.720833 0.997264 2.36E-06 3.39E-07 0.469755 TermLfltL* 0.375778 3.39E-05 0.039857 0.15346 5.58E-05 0.67944 0.972864 1.93E-05 2.9E-05 0.395113 InflL* 0.692789 0.007505 0.017592 0.71454 0.017156 0.069771 0.590027 0.442334 0.018127 0.029422 InflW* 0.632255 0.002628 0.093611 0.609595 0.006123 0.125914 0.815519 0.058529 0.000584 0.022989 SepLobL* 0.990191 0.00296 0.714523 0.052418 0.020442 0.819906 0.366962 0.001674 0.000196 0.164511 SepLobW* 0.742093 0.002533 1.13E-08 7.7E-10 0.008729 2.15E-05 0.001105 0.000198 0.3686 0.00069 BPetL* 0.360782 0.008533 0.007615 1.81E-07 0.003583 0.061121 0.061393 0.834346 2.34E-08 6.55E-11 BPetW* 8.65E-06 1.52E-05 1.21E-07 1.21E-06 5.57E-10 2.01E-11 2.31E-15 0.949182 0.004647 0.000308 WPetL* 0.012205 0.009781 0.000669 1.01E-07 0.395726 0.872539 1.73E-06 0.538168 4.93E-09 1.29E-13 WPetLamWw* 0.147163 0.000153 5.12E-05 0.186498 5.45E-05 0.001192 0.410134 0.938086 2.05E-06 1.53E-08 KPetL* 0.01088 0.02941 0.00013 3.6E-07 0.278442 0.979689 1.18E-06 0.198593 3.79E-08 1.15E-14 KPetLamWw* 0.022998 0.008466 0.002298 0.084316 0.004272 0.001726 0.036719 0.817351 4.44E-05 4.58E-07 PedPetR* 0.060818 0.029587 5.16E-07 0.001006 0.770821 0.085861 0.729231 0.00909 0.451626 0.005466 StipW* 0.217632 0.001705 0.22442 0.609058 0.000294 0.45033 0.232958 3.65E-06 6.95E-05 0.331288 PetL* 0.985687 3.04E-05 0.059282 0.223067 2.53E-05 0.270117 0.60046 0.011691 0.001054 0.018849 TermLfltW* 0.549934 1.39E-07 0.249399 0.029442 0.000226 0.913085 0.60697 4.63E-06 6.89E-08 0.443911 FlNo 0.240254 0.023195 3.49E-05 0.042192 0.012023 0.179725 0.019339 1.88E-05 0.25228 3.57E-11 PeduncL* 0.016737 0.32896 2.42E-06 2.9E-06 5.35E-06 0.367571 0.672219 5.15E-06 9.29E-05 0.546048 PedicL* 0.051993 0.002281 0.128396 0.560768 0.583787 0.20975 0.052131 0.033411 0.001517 0.182409 SepL* 0.654381 0.001164 0.276386 0.062268 0.020297 0.838877 0.151541 0.003158 8.24E-07 0.002646 SepCir* 0.4142 0.000473 2.1E-08 5.23E-07 9.68E-05 1.59E-07 7.33E-05 0.031244 0.211867 3.88E-05 WPetClwL* 0.005009 0.244029 6.01E-06 0.011554 0.125173 0.962103 0.00036 0.013748 0.006798 2.57E-10 WPetLamWn* 4.88E-06 0.000769 6.51E-09 0.000687 4.79E-06 2.07E-09 3.25E-12 0.022386 0.052575 5.54E-08 KPetClwL* 2.3E-05 0.833715 2.53E-05 0.004647 0.002797 0.044544 7.03E-08 0.003983 0.036254 7.09E-11 KPetLamWn* 9.66E-06 0.625841 4.2E-05 0.569525 6.57E-06 4.91E-08 3.31E-08 0.00191 0.882891 2.28E-06 ConFilL* 0.317708 0.228329 0.022611 5.44E-05 0.863243 0.622159 0.006159 0.396776 0.000167 5.26E-08 ConFilW* 0.317546 0.000163 0.001899 0.016571 0.034886 0.340132 0.023732 0.236434 1.21E-08 2.95E-08

A. PedicL 4.5 4 3.5 3 2.5 2 1.5 1 0.5

B. InflL 20 18 16 14 12 10 8 6

Figure 3a. Box and whisker plots of the characters used for the first couplet in the key. All measurements are in mm. A: PedicL=pedicle length, B: InflL=inflorescence length.

C. InflW 20 18 16 14 12 10 8 6

D. KPetClwL

2.6 2.4 2.2 2 1.8 1.6 1.4 1.2

Figure 3b. Box and whisker plots of the characters used for the first couplet in the key. All measurements are in mm. C: InflW=inflorescence width, D: KPetClwL=keel petal claw length.

E. WPetClwL 2.5 2.25 2 1.75 1.5 1.25 1

Figure 3c. Box and whisker plots of the characters used for the first couplet in the key. All measurements are in mm. E: PetClwL=wing petal claw length.

A. BPetW 10

3 T. peruvianum T. tafiense T. andesicola T. australe T. amabile

B. KPetLamWn 1.7

1.5

1.3

1.1

0.9

0.7 T. peruvianum T. tafiense T. andesicola T. australe T. amabile

Figure 4a: Box and whisker plots of the characters used for the second couplet in the key. All measurements are in mm. A: BPetW=banner petal width, B: KPetLamWn=keel petal lamina width at its narrowest point.

C. WPetLamWn

2 1.8 1.6 1.4 1.2 1 0.8 0.6 T. peruvianum T. tafiense T. andesicola T. australe T. amabile

Figure 4b: Box and whisker plots of the characters used for the second couplet in the key. All measurements are in mm. C: WPetLamWn=wing petal lamina width at its narrowest point.

A. StipL 19

5 T. peruvianum T. andesicola T. australe T. amabile

B. TermLfltL

22 20 18 16 14 12 10 8 6 4 T. peruvianum T. andesicola T. australe T. amabile

Figure 5a. Box and whisker plots for of the characters used for the third couplet in the key. All measurements are in mm. A: StipL=stipule length, B: TermLfltL=terminal leaflet length.

C. TermLfltW 16

2 T. peruvianum T. andesicola T. australe T. amabile

Figure 5b. Box and whisker plots for of the characters used for the third couplet in the key. All measurements are in mm. C: TermLfltW=terminal leaflet width.

A. FlNo

38 33 28 23 18 13 8 3 T. peruvianum T. australe T. amabile

B. SepCir

5.5

4.5

3.5

2.5 T. peruvianum T. australe T. amabile

Figure 6a: Box and whisker plots of characters used for the fourth couplet in the key. All measurements are in mm. A: FlNo=flower number per inflorescence, B: SepCir=sepal circumference.

C. SepLobW 1.5

1.3

1.1

0.9

0.7

0.5

0.3 T. peruvianum T. australe T. amabile

D. WPetClwL 2.5

2.25

1.75

1.5

1.25

1 T. peruvianum T. australe T. amabile

Figure 6b: Box and whisker plots of characters used for the fourth couplet in the key. All measurements are in mm. C: SepLobW=sepal lobe width, D: WPetClwL=wing petal claw length.

A. BPetLWR 1.8

1.6

1.4

1.2

0.8

0.6 T. peruvianum T. amabile

B. KPetL

3 T. peruvianum T. amabile

Figure 7a: Box and whisker plots of characters used for the fifth couplet in the key. All measurements are in mm. A: BPetLWR=banner petal length/width ratio, B: KPetL=keel petal length.

C. PedPetR 9 8 7 6 5 4 3 2 1 0 T. peruvianum T. amabile

D. SepLobW 1.5

1.3

1.1

0.9

0.7

0.5

0.3 T. peruvianum T. amabile

Figure 7b: Box and whisker plots of characters used for the fifth couplet in the key. All measurements are in mm. C: PedPetR=peduncle length/petiole length ratio, D: SepLobW=sepal lobe width.

E. WPetL 9

3 T. peruvianum T. amabile

Figure 7c: Box and whisker plots of characters used for the fifth couplet in the key. All measurements are in mm. E: WPetL=wing petal length.

Discussion

This study was the first study to look in depth at the T. amabile species complex in South America. In doing a study of this type, I was able to use a large number of characters in order to find species boundaries within the complex. This investigation of specimens indicates that the Trifolium amabile species complex in South America is not one morphological entity and instead is composed of six morphologically distinct species. Based on reexamination of all the type specimens and descriptions, disposition of the names given in table 1 is as follows: T. humboldtii Spreng, T. mathewsii A. Gray, T. macrorrhizum Ullbr., and T. pauciflorum Willd. ex Steud., are synonyms of T. amabile s. str.; T. weberbaueri Ulbrich, T. bolivianum Kennedy, T. chiclense Ball, T. peruvianum Vogel var. chiclense, and T. amabile Kunth var pentlandii Ball, are synonyms of T. peruvianum Vogel. I also investigated the possibility that any of the South American species matched the North American species described by Ahlquist (2012). The group of specimens that I delimited as T. amabile fits within the circumscription of that species as given by Ahlquist (2012), and the range of T. amabile s.str. was found to be both North and South America. Since no molecular work was done in this study or by Ahlquist (2012), future studies on this group should focus on molecular data to further test that T. amabile s.str. as circumscribed in these two studies is in fact a single species.

Disposition of the varieties of T. amabile that were recognized by Zohary and Heller (1984) as occurring in South American is as follows: T. amabile Kunth var. amabile = T. amabile s. str., T. amabile Kunth var. hemsleyi Lojac. = T. amabile s.str., and T. amabile Kunth var. pentlandii Ball = T. peruvianum Vogel.

Assigning names to each of the groups recognized in this study required examination of published descriptions and type specimens. Of these groups, two have previously been described in the literature: T. amabile Kunth and T. peruvianum Vogel. The remaining four did not correspond to any known previously described species, and will be described as new: T. tafiense Hendy et Vincent sp. nov., T. andesicola Hendy et Vincent sp. nov., T. australe Hendy et Vincent sp. nov., and T. cochabambense Hendy et Vincent sp. nov.

Taxonomic Treatment

Key to the South American Trifolium amabile species complex

1. Pedicels <.6mm long; mature flowers not reflexing; inflorescences <7mm long X <8 mm wide; keel petal claws ≤1.3mm long; wing petal claws ≤1.15mm long………….….….6.T. cochabambense 1. Pedicels >.6mm long; mature flowers reflexing or not; inflorescence >7mm long X >8mm wide; keel petal claws >1.3mm long; wing petal claws >1.15mm long…………………...... …………2 2. Banner petals >9mm wide; keel petal laminas ≥1.5mm wide at narrowest point; wing petal laminas ≥1.6mm wide at narrowest point ……………………………………………3.T. tafiense 2. Banner petals <8mm wide; keel petal laminas <1.5mm wide at narrowest point; wing petal laminas ≤1.6mm wide at narrowest point………………….…………………………………….……3 3. Stipules ≤6.8mm long; terminal leaflets ≤5.7mm long X ≤3.75mm wide …………………………………………………………………………………..……………….4.T. andesicola 3. Stipules >6.8mm long; terminal leaflets >5.7mm X >3.75mm wide…...... 4 4. Inflorescences ≥16 flowered; sepal circumferences ≤3.7mm; sepal lobes ≤.5mm wide; wing petal claws ≤1.5mm long………………..5.T. australe 4. Inflorescences <16 flowered; sepal circumference >3.7mm; sepal lobes >.5mm wide; wing petal claws >1.5mm long……………….…………………….....5 5. Banner petal length/width ratios ≤1.2mm; keel petals ≤5.3mm long; peduncle length/petiole length ratios ≤1.3; sepal lobes ≥.7mm wide; wing petals ≤6mm long…………………..2.T. peruvianum 5. Banner petal length/width ratios >1.2mm; keel petals >5.3mm long; peduncle length/petiole length ratios >1.3; sepal lobes <.7mm wide; wing petals <6mm long…………………………1.T. amabile

1. Trifolium amabile Kunth, Fig. 8. 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. Crescit in pratis prope Toluccam, s.d., Humboldt & Bonpland s.n. (holotype: P!; isotype: B (destroyed), photos GH!, NY!). Trifolium humboldtii Spreng, Syst. Veg. 3: 213. 1826. – TYPE: ECUADOR. Napo: Antisana, s.d., Humboldt s.n. (holotype: B-W, photo MU!). Trifolium mathewsii A. Gray, U.S. Expl. Exped. , Phan. 1: 398. 1854. – TYPE: PERU. Obrajillo, “1836”, Matthews s. n. (holotype: K!; possible isotypes: GH! (photos: F!, NY!)US!). Trifolium macrorrhizum Ullbr., Fedde Repert. 2: 2. 1906. – TYPE: Peru. Pampas Romas, 31 May 1903, A. Weberbauer 3213 (holotype: B (destroyed, photos NY! (2)), photo and specimen fragment F!, [LECTOTYPE to be designated]. Trifolium pauciflorum Willd. ex Steud., Nomencl. Bot. (ed. 2) 2(12-13) : 707. 1841. (non d’Urv. 1822) – TYPE: ECUADOR. Napo: Antisana, s.d., Humboldt, s.n. (holotype: B-W, photo MU!).

Plants perennial; roots woody, 1.9-37.5 mm in diameter; stems decumbent-prostrate 6- 66+ cm long, 0.8-2 mm in diameter, internodes sparsely-moderately pubescent, nodes sparsely- moderately pubescent; leaves trifoliate, ± uniform in size throughout; stipules triangular- narrowly triangular-sub-equal, 6.5-18.4x2-6.3 mm, glabrous adaxially, glabrous-sparsely pubescent abaxially, margins subentire-entire, glabrous-moderately pubescent, apices acute; petioles 3.8-60.2x≤0.7 mm, sparsely-densely pubescent; petiolules 0.5-1.1 mmx≤0.7 mm, sparsely-densely pubescent; leaflets elongate-ovoid-obcordate, glabrous adaxially, sparsely- moderately pubescent along midrib abaxially, glabrous-½ laterally sparse abaxially, bases acute, margins weakly serrate-serrate, glabrous-moderately pubescent, apices rounded-acute- emarginate, weakly mucronulate, terminal leaflets 5-22.2x3.8-15.5 mm; peduncles 6-86.8x≤1 mm, sparsely-densely pubescent; inflorescences axillary, racemose-umbellate, 7.3-18.8x6.7- 19.5 mm; flowers 3.5-26, each subtended by a linear triangular bract, in 1-3 whorls, terminal sterile appendage sometimes present; pedicels 1-4x≤0.6 mm, sparsely-moderately pubescent;

28 calyx 3.7-7 mm long, tube 2.9-5.7 mm in circumference, glabrous-densely pubescent, lobes narrowly triangular, 1.8-5.4x0.3-1.05 mm, glabrous-densely pubescent abaxially, margins entire-subentire, glabrous-densely pubescent, apices attenuate; petals color; banner petals obovate shallow notch, 5.7-10.3x3.3-6.8 mm; wing petals 5.3-8.9 mm long, asymmetrically clawed, claws 1.3-2.4x0.4-0.7 mm, lamina spatulate, 3.5-6.5 mm long, 1.2-2.8 mm wide at widest point, 0.9-1.6 mm wide at narrowest point, bases shortly-broadly auriculate, margins entire, apices rounded-slightly acute; keel petals 4.6-7.3 mm long, asymmetrically clawed, claws 1.5-2.6x0.4-0.7 mm, lamina spatulate, 2.7-4.7 mm long, 1.2-2.2 mm wide at widest point, 0.9-1.7 mm wide at narrowest point, bases obtusely auriculate, auricles rounded, margins entire, apices rounded-acute; stamens diadelphous, connate filaments 3.1-5.5x1-2 mm, fused for 26-61% of their length, free filaments 1.9-3.9x≤0.2mm; pistils sessile, ovaries 1.5-3.5x0.4-1.1 mm, glabrous-moderately pubescent, styles 1-2.3x≤0.2 mm, glabrous-sparsely pubescent, ovules 2-4; legumes laterally compressed, 3.3-6.1x1.3-3.2 mm; seeds 1-3, ovoid-mitten shaped, yellowish brown-brown occasionally dark speckeled,1.5-2.4x0.9-2.3 mm.

Representative specimens ARGENTINA. Salta: Santa Victoria, Arroyo Peña Negra, -22.250833°, -65.0775°, 3360m, 15 Feb 2009, F.O. Zuloaga, et al. 10745 (MU); Tucumán: Tafí, Cerro, San José, -27.7°, -65.5°, 2700m, 11 Feb 1925, S. Venturi, 3569 (US); Catamarca: Andalgalá, Estancia Yunka Suma, Cerro Chapina, -27.2°, -66.1°, 2650m, 7 Mar 1950, E. Pedersen & J.P. Hjerting 1661 (C); Jujuy: Valle Grande, Cerro Hermosos, -23.591111°, -64.873056°, 3302m, 20 Feb 2009, F.O. Zuloagaet al. 11086 (MU); Córdoba: Sierra Achala, Las Ramalas, Pampa de San Miguel, -31.9°, -65°, 1400m, 11 Feb 1890, F. Kurtz, 6876 (NY). BOLIVIA. La Paz: Murillo, La Paz-Cota Cota, 16.56°, -68.06°, 3600m, 20 Dec 1981, St. G. Beck 8007 (LPB); Chuquisaca: Belisario Boeto, Comunidad Nuevo Mundo, -19.004167°, - 64.289444°, 2298m, 28 Nov 2005, J. Villalobos et al. 379 (MO); Tarija: José María Aviléz, -21.8°, - 64.95°, 2800m, 23 Apr 2000, S. G. Beck & N. Paniagua 27426 (LPB); Cochabamba: Cumbre de Bombeo, 3km SE Pongo, -17.6785°, -66.425528°, 3518m, 3 Oct 2003, L. Rico et al. 1509 (MO); Oruro: Sebastian Pagador, Cantón Huari, Municipio de Huari, Ayllu Yucasa, en el borde del

29 camino, lado de la acequia, -19.021101°, -66.767984°, 3825m, 27 Jan 2008, S. G. Beck et al. 29979 (MU). COLOMBIA. Distrito Capital: In montibus bogotensibus Andinum oriental, Novae granatae, [Andes near Bogotá], 4.6°, -74.1°, 2800m, 29 Dec 1875, E. André 917 (NY); Cundinamarca: Guasca—Andes, 4.9°, -73.9°, 3000m, Aug 1931, P.E. Arbelaez 1130 (US); Valle del Cauca : Cartago, 4.75°, -75.9°, 1600m, 24 Aug 1924, R.T. Rojas 72 (US); Santander: Vicinity of Tona, 7.2°, -72.9°, 2000m, 17 Feb 1927, E.P. Killip & A.C. Smith 19514 (F); Nariño: Pasto, Andes de Tuquerres, 1.19°, -77.27°, 3000m, J. Triana s. n. (P). ECUADOR. Cotopaxi: Cordillera Occidental; Cordillera de Angamarca y Zumbagua; Páramo de Zumbagua, rocky slope above large rock outcrop cut by stream; surrounded by cultivated fields and stream; surrounded by cultivated and open páramo. Punto Quindeseye, between first crest above Pujilí and Indian village, Zumbagua, -0.95°, -78.9°, 3700m, 17 Jul 1959, H.G. Barclay & P. Juajibioy 8053 (US); Carchi: Ca. 2km along the road El Angel-Tulcán, 0.633333°, -77.916667°, 3300m, 14 Mar 1973, L. Holm-Nielsen, et al. 5222 (NY); Imbabura: Cotacachi Canton, Reserva Ecológica Cotacachi-Cayapas, Laguna de Cuicocha, Sendero alrededor do la Laguna, bosque Húmedo Montano, Suelo Volcánico, 0.3°, -8.366667°, 3100m, 23 Dec 1991, M. Peñafiel et al. 685 (MO); Loja: Jera, 10km north of Saraguro, -3.532°, -79.238°, 2400m, 15 Feb 1989, L. Elleman 66986 (AAU); Cañar : near the village of San Marcos, (5-8km northeast of Azogues), -2.7°, -78.8°, 3000m, 13 Apr 1945, F. Prieto E-2509 (F); Bolívar: Simiatug: Hacienda Talahua, -1.35°, -79°, 3300m, 29 Apr 1939, C.W. Penland & R. H. Summers 569 (US); Pichincha: Quito, -0.2°, -78.5°, 2900m, C. Hartweg 940 (P); Chimborazo: Alausi, -2.2°, -78.8°, 2300m, 09 Jun 1939, E. Asplund 6799 (F); Azuay: Carretera Gulaquiza Sigsig Gualaquiza colecciones en Altarurco a 50km de Gualaceo, -3.011126°, -78.647296°, 2950m, 31 Aug 1984, J. Jaramillo 7147 (MO). PERU. Cajamarca: Lucmacucho, 1 km al NO de Cajamarca, sobre la ladera que converge al Río Ronquillo, -7.15°, -78.5°, 2780m, 20 Feb 1992, I. M. Sánchez Vega 6099 (F); Puno: Sandia, 2-6 km S. of Limbani, -14.18°, -69.7°, 3650m, 11 May 1942, R. D. Metcalf 30441 (US); Huancavelica: Ayán, abajo de Conaica, -12.5°, -75°, 3300m, 13 Mar 1951, O. Tovar 157 (US); Piura: Rio Blanco, old mine road, -4.866°, -79.35°, 3600m, 8 May 1922, MacBride &

Featherstone 821 (F); Pasco: Hacienda 9 miles upriver from Yanahuanca, Chinche, -10.494°, - 76.561°, 3500m, 21 Jun 1922, Macbride & Featherstone 1268 (F); Ancash: Huaylas road from Pamparomas to Caraz, steep, weedy slope on deep, black clay soil, forest remnants of Cantua cf. buxifolia, -9.091194°, -77.972472°, 3153m, 27 Apr 2006, M. Weigend et al. 8475 (F); Junín: Mito, shrubby southwestern slope, -11.937°, -75.339°, 3400m, 08 Jul 1922, Macbride & Featherstone 1440 (F); Cusco: Valle de Apurímac, -13.4°, -72.8°, 2200m, Feb 1929, F. L. Herrera, s. n. (F); Ayacucho: Lucanas, -14.7°, -74.5°, 3300m, 20 Mar 1949, R. Ferreyra 5527 (US); La Libertad: mountain slopes above Yamobamba, -7.8°, -78.1°, 3100m, 23 Feb 1974, J. Conrad 2733 (US); Apurímac: Abancay, Laderas del Ampay, -13.568°, -72.922°, 3400m, Jan 1938, C. Vargas 767 (F); Lima: along Río Chillón, above Obrajillo, -11.45°, -76.6°, 2700m, 13 Jun 1925, F. W. Pennell 14375 (NY).

Distribution and habitat Trifolium amabile spans the entire South American range of the Andes. It is found in Colombia, Ecuador, Peru, Bolivia, and Argentina (Fig. 9). This species occupies a wide range of habitats, including open hillsides, along roadsides, grasslands, open moist places, among rocks, very firm sandy soil, steep slopes, and alpine conditions. Collections have been made from an elevation range of 1400m to 3800m. This species also occurs in México, Guatemala, and Costa Rica (Ahlquist 2012). It should be noted that Zohary and Heller (1984) list a range for T. amabile that includes Chile. No known specimens of T. amabile from Chile were found in this study.

Comments Trifolium amabile most closely resembles T. australe but can be distinguished by its looser inflorescences with fewer flowers 3.5-26 compared to 13-40 respectively. Trifolium amabile also has larger flowers than T. australe, with correspondingly larger sepals, 3.7-6.95mm compared to 3.15-5.4mm, wing petals 5.3-8.9mm compared to 3.45-6.2mm, and keel petals 4.6-7.3mm compared to 3.25-5.05mm, respectively. Trifolium amabile can be distinguished from T. peruvianum by having longer narrower banner petals, 5.75-10.3x3.35-6.8mm,

31 compared to 5.1-7.7x3.85-7.9mm, respectively, and longer narrower sepal lobes, 1.8-5.4x.35- 1.05mm, compared to 1.95-4.05x0.6-1.45mm, respectively. Additional synonymy for material from North America and Central America is provided by Ahlquist (2012). In the course of this study, the holotype of T. amabile (which was unknown to Ahlquist 2012)) was found to be intact in the Paris herbarium. As a result, the neotype designated by Ahlquist (2012) is superfluous. The type specimen of T. matthewsii A. Gray (K), is labeled “Mathews, 1836, Prov. of Chachapoyas, Peru”. A specimen purporting to be an isotype (US) is labeled “Herbarium of the South Pacific Exploring Expedition under the command of Capt. Wilkes, U.S.N., 1838-42”. If the US specimen is a duplicate of the K material, it appears that the K material is mislabeled as to the year in which it was collected.

Figure 8. Representative specimen of Trifolium amabile (Zuloaga et al. 10745 (MU)).

Figure 9. Geographical range of Trifolium amabile in South America, based on representative specimens (map generated using ESRI 2013).

2. Trifolium peruvianum Vogel, Fig. 10. Trifolium peruvianum Vogel, Nov. Act. Nat. Car. 19: Suppl. 1: 12. 1843. TYPE: Peru. Puno, Laguna de Titicaca. Jul 1931, Meyen s. n. (holotype: B (destroyed photos NY! (2)) photo and specimen fragment F! EPITYPE to be designated: Bolivia. Depto. La Paz, Prov. Murrillo, 27 Feb 1982, E. Valenzuela 83 (MU), isoepitype: LPB). Trifolium weberbaueri Ulbrich, Feddes Repert. 2: 2. 1906. TYPE: Peru: Cusco, Pucará, bahnstation Puno–Cuzco, 24 Feb 1902, A. Weberbauer 404. (holotype: B (destroyed photos NY!, F!); isotype: F!, [LECTOTYPE to be designated]. Trifolium bolivianum Kennedy, Muhlenbergia 7: 97. 1911. TYPE: Bolivia. s.d., Bang 2819. (holotype: MICH!, isotype: K!). Trifolium chiclense Ball, Jour. Linn. Soc. Bot. 22: 35. 1885. T. peruvianum Vogel var. chiclense (Ball) Macbr., Field Mus. Publ. Bot. 13: 451. 1943. TYPE: Peru. Chicla, 21-23 Apr 1882, J. Ball s.n. (holotype: K!) Trifolium amabile Kunth var pentlandii Ball, Jour. Linn. Soc. Bot. 22: 35. 1885. TYPE: Peru. Chicla. 21-23 Apr 1882, J. Ball s.n. (holotype: K!).

Plants perennial; roots woody-large 4.1-5.3 mm in diameter; stems prostrate 10-41+ cm long, 1.3-1.9 mm in diameter, internodes sparsely-densely pubescent, nodes sparsely- densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules sub-equal, 6.6- 16.5x1.9-5.8 mm, glabrous adaxially, glabrous-sparsely pubescent abaxially, margins entire, sparsely-moderately pubescent, apices acute; petioles 9.3-40.5-60.2x≤0.7 mm, sparsely-densely pubescent; petiolules 0.5-1.0 mmx≤0.5 mm, sparsely-moderately pubescent; leaflets elongate- ovoid-obcordate, glabrous adaxially, sparsely pubescent along midrib abaxially, glabrous-½ laterally sparse abaxially, bases acute, margins subentire-serrate, glabrous-sparsely pubescent, apices rounded-acute-emarginate, weakly mucronulate, terminal leaflets 5.7-15.2x3.4-9.3 mm; peduncles 3.7-51x≤1 mm, glabrous-densely pubescent; inflorescences axillary, racemose- umbellate, 9.9-19.2x9.1-17.6 mm; flowers 7-24, each subtended by a linear triangular bract, in 1-4 whorls; pedicels 1.3-4.2x≤0.3 mm, sparsely-densely pubescent; calyx 3.6-6.6 mm long, tube 2.8-6.1 mm in circumference, sparsely-densely pubescent, lobes broadly triangular, 1.9-4.1x0.6-

1.45 mm, sparsely-densely pubescent abaxially, margins entire-subentire, sparsely-densely pubescent, apices acute; petals white-pink; banner petals obcordate-rounded shallow notch, 5.1-7.7x3.8-7.9 mm; wing petals 4.3-6.9 mm long, asymmetrically clawed, claws 1.4-2.0x0.4-0.5 mm, lamina spatulate, 3.1-5.1 mm long, 1.2-2.2 mm wide at widest point, 0.9-1.6 mm wide at narrowest point, bases shortly-longly-broadly auriculate, margins entire, apices rounded; keel petals 3.8-5.7 mm long, asymmetrically clawed, claws 1.6-2.4x0.4-0.6 mm, lamina spatulate, 2.7-3.7 mm long, 1.2-2 mm wide at widest point, 0.8-1.45 mm wide at narrowest point, bases smally-auriculate to no auricle, auricles rounded, margins entire, apices rounded-acute- acuminate; stamens diadelphous, connate filaments 3.2-4.6x1-1.8 mm, fused for 29-45% of their length, free filaments 2-2.8x≤0.2mm; pistils sessile, ovaries 2.2-4x0.5-1.6 mm, glabrous- moderately pubescent, styles 0.8-1.4x≤0.2 mm, glabrous pubescent, ovules 2-4; legumes laterally compressed, 4.7x2.7-2.9 mm; seeds 2, mitten shaped, yellowish brown-brown occasionally dark speckeled,1.9-2.1x1.7 mm.

Representative specimens ARGENTINA. Jujuy: Cochinoca, just E of Abra Rechaite, along Rte. 74, ca. 70km W of Abra Pampa, dry hillsides with corrals and low scrub, -22.856389°, -66.189722°, 3820m, 17 Mar 1994, C.M. Taylor et al. 11320 (MO); Córdoba: Tierra Schala, -31.42°, -64.22°, Feb 1990, F. Kurtz s. n. (NY); Tucuman: Dpto Tafi, Bajada del Infiernillo, -26.85°, -65.70°, 2000m, 19 Mar 1979, A. L. Cabrera, et al. 30832 (MU); Salta: Dto. Chicoana, Cuesta del Obispo, -25.10°, -65.53°, 3100, 26 Feb 1987, E. Nicora, et al. 9212 (NY). BOLIVIA. Cochabamba: Mizque, Localidad de Sacha Loma, -17.44445°, -65.3431°, 3700m, 15 Mar 2000, M. Mercado 2280 (MO); La Paz: -16.5°, -68.14°, 3700m, 19 Mar 1920, E. W. D. Holway & M. M. Holway 426 (US); Tarija: Aviles, Lagunas de Tajzara, -21.7°, -65.1°, 3800m, 1 Mar 1960, T. Meyer et al. 21532 (SI). ECUADOR. Cotopaxi: Laguna Quilotoa, Paramo, -0.883333°, -78.916667°, 3550m, 30 Nov 1998, D. Benyamini 591 (MO); Bolívar: ca. 10 km W of Pogyos, Paramo, -1.443611°, - 78.944444°, 4050m, 1 Feb 2009, E. J. Tepe & S. Stern 2682 (MU); Chimborazo: Urbina, Paramo

36 on east flank of Mt. Chimborazo, -1.5°, -78.7°, 3600m, 5 Oct 1923, A. S. Hitchcock 22027 (NY); Pichincha: Quito, -0.2°, -78.5°, 2800m, Nov 1902, M. Rivet 277 (P). PERU. Junín: Between Tarma and Jauja, Puna, -11.6°, -75.5°, 4500m, 24 Apr 1929, E.P. Killip & A. C. Smith 21972 (NY); Puno: Chuquibambilla, thin soil over limestone on puna, - 14.73°, -70.68°, 4000m, 19 Apr 1925, F. W. Pennell 13393 (F); Cusco: Sacsahuaman, above Cusco, open rocky slope, -13.505°, -71.980°, 3580m, 24 Apr 1925, F. W. Pennell 13596 (F); Lima: Huarochiri, Chicla, hard gravelly ground, -11.7°, -76.2°, 4500m, 6 Jun 1940, E. Asplund 11461 (US); Piura: Rio Blanco, prostrate on steep W. bunch-grass slope, -4.866°, -79.35°, 3600m, 08 May 1922, Macbride & Featherstone 749 (F); Ancash: Huaylas Province, Huascarán National Park, ruins at Auquispuquio, grassland/shrubland, intermittantly grazed or farmed; steep slopes, southern aspects, -8.833333°, -77.966667°, 3900m, 08 Apr 1986, D.N. Smith e tal. 12041 (F).

Distribution and habitat Trifolium peruvianum is a wide-ranging species found in Ecuador, Peru, Bolivia, and Argentina (Fig. 11). It has been collected mainly on steep rocky slopes as a small, often growing as cushion-type plant. It is sometimes found on grassy areas, shrublands, or grazed areas. It has been collected at high elevations from a range of 2800m to 4500m.

Comments Trifolium peruvianum is similar to T. tafiense and T. andesicola. It can be distinguished from T. tafiense by its banner petals 3.85-7.9mm that are narrower than T. tafiense 9.05- 9.95mm. It can be distinguished from T. andesicola by its banner petals 3.85-7.9mm that are wider than T. andesicola 3.8-5mm. Trifolium peruvianum can also look like T. amabile at times especially when it has longer peduncles. It can be distinguished from T. amabile by its shorter and wider banner petals, 5.1-7.7x3.85-7.9mm compared to 5.75-10.3mmx3.35-6.8, and wider sepal lobes 0.6-1.45mm compared to 0.35-1.05mm, respectively. An epitype is being designated to represent Trifolium peruvianum since the only known original material, a small fragment at F in a packet on sheet 649106 (with a photograph of the

37 type specimen from B) consists only of a single flower and a single leaf, and is inadequate to serve as the type.

Figure 10. Representative specimen of Trifolium peruvianum (Valenzuela 83 (MU)).

Figure 11. Geographical range of Trifolium peruvianum based on representative specimens (map generated using ESRI 2013).

3. Trifolium tafiense Hendy et Vincent, sp. nov. (ined.), Fig. 12. TYPE: Argentina. Tucuman: Tafi, Ruta 307, camino de Tafí del Valle a Amaicha del Valle, El Infernillo, 10 Feb 2002, A. M. Cialdella, N. B. Deginani, & L. M. Giussani 148 (holotype: MU!; isotype: SI!).

Plants perennial; roots woody, 2.9-6.4 mm in diameter; stems decumbent 28-29+ cm long, 0.9-1.5 mm in diameter, internodes sparsely pubescent, nodes sparsely-moderately pubescent; leaves trifoliate, ± uniform in size throughout; stipules triangular-narrowly triangular-sub-equal, 8.9-14.7x3.4-6 mm, glabrous adaxially, glabrous-sparsely pubescent abaxially, margins entire, moderately-densely pubescent, apices acute; petioles 17-23.8x≤0.6 mm, sparsely-moderately pubescent; petiolules ≤1.0x ≤0.4 mm, sparsely-moderately pubescent; leaflets elongate-ovoid-obcordate, glabrous adaxially, sparsely-moderately pubescent along midrib abaxially, ½ laterally sparse abaxially, bases acute, margins weakly serrate-serrate, glabrous-sparsely pubescent, apices rounded-acute-emarginate, weakly mucronulate, terminal leaflets 9-15x4.9-10 mm; peduncles 29.9-43.3x≤0.7 mm, sparsely- densely pubescent; inflorescences axillary, racemose-umbellate, 10.8-15.5x11.3-16 mm; flowers 8-24, each subtended by a linear triangular bract, in 1-3 whorls, terminal sterile appendage sometimes present; pedicels 1.6-1.8x≤0.3 mm, moderately pubescent; calyx 4.2-5.2 mm long, tube 4.7-5.4 mm in circumference, moderately-densely pubescent, lobes triangular, 2.4-3.6x0.6-1.05 mm, moderately-densely pubescent abaxially, margins entire-subentire, moderately-densely pubescent, apices acute; petals pink-red; banner petals broadly elliptical shallow notch, 6.1-6.8x9.05-9.95 mm; wing petals 3.9-4.9 mm long, asymmetrically clawed, claws 1.3-1.7x0.4 mm, lamina spatulate, 3.1 mm long, 1.8-2.3 mm wide at widest point, 1.6-2 mm wide at narrowest point, bases truncate, margins entire, apices rounded-obtuse; keel petals 3.7-4 mm long, asymmetrically clawed, claws 1.3-1.6x0.4 mm, lamina spatulate, 3.1 mm long, 1.5-2.6 mm wide at widest point, 1.5-1.6 mm wide at narrowest point, bases truncate- shortly auriculate, auricles rounded, margins entire, apices rounded; stamens diadelphous, connate filaments 3.5x1.2-1.6 mm, fused for 33-44% of their length, free filaments 2.2-

2.3x≤0.1mm; pistils sessile, ovaries 1.9-2x0.6 mm, glabrous-sparsely pubescent, styles 0.9- 1x≤0.2 mm, glabrous, ovules 2-3; legumes not seen; seeds not seen.

Paratypes Argentina. Tucuman: Tafi, Sierras Calchaquíes: La Queñua. No Frecente en quebradas, -26.3833°, -65.6667°, 3000m, 28 Jan 1933, A. Burkart 5473 (F); Tucuman: Tafi, Valle de Tafí, - 26.845°, -65.7°, 2050m, 1908, C. Bruch s.n. (NY); Tucuman: Tafi, Tafí del Valle. En quebradas, - 26.845°, -65.7°, 2250m, 3 Feb 1933, A. Burkart 5178 (F).

Distribution and habitat Trifolium tafiense has a very limited range. All specimens of T. tafiense examined were collected in ravines from 2050m to 3000m in elevation, in Tucumán Province, Argentina, in the Tafi Valley (Fig. 13).

Comments Trifolium tafiense is unique in that it has very wide banner petals. It is easy to recognize by its blocky-looking flowers. Trifolium tafiense is most similar to T. peruvianum, but T. tafiense differs in having much wider banner petals, 9.05-9.95mm compared to 3.85-7.9mm respectively, and wider wing petal laminas at their narrowest point 1.6-2mm compared to 0.9- 1.6mm and keel petal laminas at their narrowest point 1.5-1.55mm compared to 0.8-1.35mm respectively. Trifolium tafiense was named for the Tafi Valley in Argentina. This is the only known location for the species.

Figure 12. Trifolium tafiense (Cialdella et al. 148 (MU)).

Figure 13. Geographical range of Trifolium tafiense, based on all known specimens (map generated using ESRI 2013).

4. Trifolium andesicola Hendy et Vincent, sp. nov. (ined.), Fig. 14 TYPE: Argentina. Salta: Dpto. Chicoana, Ruta 33, de Cachi a Ciudad de Salta, pasando desvío a Ruta 42, 15 Feb 2002, A. M. Cialdella, N. B. Deginani, & L. M. Giussani 310 (holotype: MU!, isotypes: MO!, SI!).

Plants perennial; roots woody 2.4-4.7 mm in diameter; stems prostrate 5-15+ cm long, 1-1.2 mm in diameter, internodes sparsely-moderately pubescent, nodes glabrous-moderately pubescent; leaves trifoliate, ± uniform in size throughout; stipules sub-equal, 5.5-6.8x1.7-3 mm, glabrous adaxially, glabrous-sparsely pubescent abaxially, margins entire-slightly serrate, glabrous-sparsely pubescent, apices acute; petioles 7-12.3x≤0.4 mm, sparsely-moderately pubescent; petiolules ≤0.7 mmx≤0.3 mm, glabrous-sparsely pubescent; leaflets elongate- ovoid-obcordate, glabrous adaxially, sparsely-moderately pubescent along midrib abaxially, glabrous-½ laterally sparse abaxially, bases acute, margins serrate, glabrous-sparsely pubescent, apices rounded-acute-emarginate, weakly mucronulate, terminal leaflets 4.7- 6.4x2.5-3.8 mm; peduncles 9-20.5x≤0.4 mm, sparsely pubescent; inflorescences axillary, racemose-umbellate, 7.7-12.8x8.2-13.2 mm; flowers 6-13, each subtended by a linear triangular bract, in 1-2 whorls; pedicels 1.1-2.1x≤0.2 mm, sparsely-moderately pubescent; calyx 2.7-4.4 mm long, tube 2.8-4.1 mm in circumference, sparsely-densely pubescent, lobes triangular, 1.9-2.9x0.6-0.8 mm, sparsely-moderately pubescent abaxially, margins entire- subentire, sparsely-moderately pubescent, apices acute; petals white-pink-red; banner petals obcordate shallow notch-notch absent, 4.8-6.4x3.8-5 mm; wing petals 4.2-5.9 mm long, asymmetrically clawed, claws 1.4-2.0x0.3-0.4 mm, lamina spatulate, 2.7-3.2 mm long, 1-1.6 mm wide at widest point, 0.8-1.2 mm wide at narrowest point, bases broadly auriculate-no auricle, margins entire, apices rounded-narrowly rounded; keel petals 3.9-5.4 mm long, asymmetrically clawed, claws 1.6-2.3x0.4-0.5 mm, lamina spatulate, 2.7-3.2 mm long, 1.2-1.6 mm wide at widest point, 1-1.3 mm wide at narrowest point, bases smally-auriculate to no auricle, auricles rounded, margins entire, apices rounded-acute; stamens diadelphous, connate filaments 3.3- 4.2x1-1.4 mm, fused for 30-33% of their length, free filaments 1.9-2.4x≤0.4mm; pistils sessile, ovaries 1.9-3x0.7-0.9 mm, sparsely-moderately pubescent, styles 0.9-1x≤0.2 mm, glabrous

45 pubescent, ovules 2-3; legumes laterally compressed, 5.4x2.3 mm; seeds 2, sub-reniform-ovoid shaped, dark yellow green-brown 2x1.9 mm.

Paratypes ARGENTINA. Tucumán: Tafí, Ruta 307, camino de Tafí del Valle a Amaicha del Valle, El Infernillo, -26.75°, -65.75°, 2970m, 10 Feb 2002, A.M. Cialdella et al. 148 (MU); Catamarca: Andalgalá, El Candado, -27.316°, -66.2°, 5200m, 1916, P. Jörgensen 1339 (US); Jujuy: Cochinoca, Abra pueblo de Cochinoca, -22.733333°, -65.883333°, 3690m, 8 Feb 1995, N.B. Deginani et al. 496 (MO); Córdoba: Pampa de Achala, -31.4166°, -64.5166°, 689m, 01 Feb 1936, A. P. Rodrigo 475 (NY). BOLIVIA. Tarija: Aviles, Tajzara, -21.9°, -65.03°, 3650m, 10 Mar 1986, E. Bastión 1011 (NY); La Paz: Pacajes Caquiaviri, Kalla Centro, -17.023°, -68.594°, 3950m, 29 Jan 1983, T. Johns 83-17 (LPB); Potosí: Tomás Frias, “Las Lecherías", -19.58°, -67.75°, 3800m, 06 Feb 1988, Schulte, 135 (LPB). PERU. Pasco: La Quinua, path along stream, -10.6°, -76.767°, 3700m, 14 May 1922, Macbride & Featherstone 2024 (F); Cajamarca: surroundings of Cajamarca, capital of the Departmento Cajamarca, - northern Peru, 7.162°, -78.51°, 2700m, 03 Apr 1986, B. Becker & H. Terrones 927 (NY); Cusco: Urubamba, Chincheros, above community of Taucca, -13.41667°, - 72°, 4000m, 09 Feb 1982, S. King, et al. 228 (F).

Distribution and habitat Trifolium andesicola is found in Peru, Bolivia, and Argentina (Fig. 15). It grows on exposed ground, along streams, and in ravines, at an elevation range of 680m to 5200m.

Comments Trifolium andesicola is most similar to T. cochabambense. It can be distinguished from T. cochabambense by having longer pedicels, 1.18-2.3mm compared to 0.52mm, respectively. Trifolium andesicola can be distinguished from T. peruvianum by having much smaller inflorescences, 7.75-12.83x8.25-13.27mm compared to 9.9-19.2x9.1-17.6mm, and fewer

46 flowers per inflorescence, 6.8-13 compared to 7.4-24, respectively. Trifolium andesicola and T. australe both have smaller flowers then T. amabile, T. peruvianum, and T. tafiense, but T. andesicola can be distinguished by its sparser inflorescences 6.8-13 flowers per inflorescence as compared to 13-40, and its smaller leaves and stipules. Trifolium andesicola was named for the Andes Mountains in which it is found.

Figure 14. Trifolium andesicola (Cialdella et al. 310 (MU)).

Figure 15. Geographical range of Trifolium andesicola based on representative specimens (map generated using ESRI 2013).

5. Trifolium australe Hendy et Vincent sp. nov. (ined.), Fig. 16. TYPE: Bolivia: Tarija. Cerca Tucumille, 22 Jan 1986, E. Bastión, 489 (holotype: LPB!; isotype: NY!)

Plants perennial; roots woody-large 2-10.4 mm in diameter; stems decumbent- prostrate 7-52+ cm long, 1.1-1.7 mm in diameter, internodes sparsely-densely pubescent, nodes sparsely-densely pubescent; leaves trifoliate, ± uniform in size throughout; stipules sub- equal, 8.2-16.7x3-6 mm, glabrous adaxially, glabrous-sparsely pubescent abaxially, margins entire-slightly serrate, glabrous-sparsely pubescent, apices acute; petioles 7.3-33.6x≤0.7 mm, sparsely-densely pubescent; petiolules 0.6-1.1 mmx≤0.5 mm, sparsely-densely pubescent; leaflets elongate-ovoid-obcordate, glabrous adaxially, sparsely-moderately pubescent along midrib abaxially, glabrous-½ laterally sparse abaxially, bases acute, margins serrate, glabrous- sparsely pubescent, apices rounded-acute-emarginate, weakly mucronulate, terminal leaflets 6.9-18.7x4.3-1 mm; peduncles 16-78x≤0.75 mm, sparsely-densely pubescent; inflorescences axillary, racemose-umbellate, 8.2-15x8.3-15.2 mm; flowers 13-40, each subtended by a linear triangular bract, in 1-2 whorls; pedicels 1.3-3.5x≤0.3 mm, sparsely-densely pubescent; calyx 3.1-5.4 mm long, tube 2.8-4.3 mm in circumference, sparsely-densely pubescent, lobes triangular, 1.6-3.7x0.3-0.8 mm, sparsely-moderately pubescent abaxially, margins entire, sparsely-densely pubescent, apices acute; petals white-pink-red-purple; banner petals obcordate shallow notch, 3.6-7.7x3.1-5.5 mm; wing petals 3.4-6.2 mm long, asymmetrically clawed, claws 1.1-1.9x0.3-0.5 mm, lamina spatulate, 2.3-4.6 mm long, 0.8-2.1 mm wide at widest point, 0.6-1.1 mm wide at narrowest point, bases deeply auriculate-no auricle, margins entire, apices rounded; keel petals 3.2-5.1 mm long, asymmetrically clawed, claws 1.2-1.9x0.3- 0.5 mm, lamina spatulate, 1.9-3.3 mm long, 0.9-1.8 mm wide at widest point, 0.7-1.2 mm wide at narrowest point, bases no auricle, margins entire, apices rounded-acute-acuminate; stamens diadelphous, connate filaments 2.6-4.3x0.7-1.6 mm, fused for 23-47% of their length, free filaments 1.8-2.8x≤0.4mm; pistils sessile, ovaries 1.7-2.7x0.5-1 mm, glabrous-moderately pubescent, styles 0.8-1.9x≤0.2 mm, glabrous-sparsely pubescent, ovules 2-4; legumes laterally compressed, 3.9x5.8 mm; seeds 2-3, mitten shaped, light brown-brown 1.5-1.6x1.2-1.3 mm.

Paratypes ARGENTINA. La Rioja: La Ciénega; Sierra de Tucuman, -30.733°, -66.233°, 560m, 10 Jan 1874, P. G. Lorentz & Hieronymus s.n. (F); Salta: Orán, San Andres, -23.06°, -64.6°, 1800m, 08 Feb 1945, S. A. Pierotti 264 (NY); Catamarca: Andalgalá, El Candado, -27.316°, -66.2°, 5200m, 1916, P. Jörgensen 1339 (F); Córdoba: Pampa de Achala, -31.416668°, -64.516670°, 689m, 15 Jan 1940, A. Burkart 10290 (MU); Jujuy: Cuesta de Garabatal, -24.2°, -64.4°, 1400m, Jan 1874, Lorentz & Hieronymus 1057 (NY); Tucumán: Tafí, Cumbre de Siambon, -26.845°, -65.7°, 2250m, 25 Nov 1944, D. Olea 1 (NY). BOLIVIA. Potosi: Cantón Tuero Saavedra, Comunidad de Despensa. -19.333333°, - 65.333333°, 3000m, 17 Mar 1989, C. J. de Dios Romero 48 (LPB); La Paz: Inquisivi, "Carabuco" - Along the road between Choquetanga and the Carabuco Power Station. 0.5-1.5 km N. of Choquetanga, -16.833333°, -67.316667°, 3200m, 27 Jan 1990, M. Lewis 37006 (MO); Cochabamba: Ayopaya, Cuenca Rio Tambillo, Barbecho en direccion NE a fuera y arriba de Independencia, -17.08°, -66.81°, 2820m, 15 Mar 1989, R. Baar 39 (MU); Santa Cruz: Carretera entre Vallegrande y Pucara, -18.636333°, -64.122333°, 2793m, 05 Mar 2005, J.R.I. Wood et al. 21781 (LPB); Oruro, Poopó, Pazña, a 1km al este del balneario de Urmiri pasando la ex central eléctrica, -18.576639°, -66.877278°, 3729m, 12 Mar 2006, L. Torrico & G. Castillo 571 (MO); Chuquisaca: Oropeza, 1km from Sucre towards Tarabuco, -19.0823°, -65.2321°, 2880m, 06 Feb 1994, J. R. I. Wood 7924 (MU).

Distribution and habitat Trifolium australe has a southern range within South America being found in Bolivia and Argentina (Fig. 17). This species was collected along roadsides and on open sandy areas by streams at an elevation of 680m to 5200m.

Comments Trifolium australe most closely resembles T. amabile. It can be distinguished by its much denser inflorescences, 13-40 flowers per inflorescence compared to 3.5-26, generally smaller inflorescences, 8.2-15x8.3-15.7mm compared to 7.3-18.8x6.75-19.5mm. Trifolium

51 australe can be distinguished from T. andesicola by its larger vegetative characters in general, often purple flowers, and much denser inflorescences, with 13-40 flowers per inflorescence compared to 6.8-13, respectively.

Figure 16. Trifolium australe (Bastión 489 (LPB)).

Figure 17. Geographical range of Trifolium australe (map generated using ESRI 2013).

6. Trifolium cochabambense Hendy et Vincent sp. nov. (ined.), Fig. 18. TYPE: Bolivia. Cochamamba: Quillacollo. Bosque de Polylepis, camino Sipe-Sipe – Lipichi, 02 Mar 1990, I. Hensen 569 (holotype: MU!; isotype: LPB!).

Plants perennial; roots woody, 2.3 mm in diameter; stems 10+ cm long, 1.2 mm in diameter, internodes sparsely pubescent, nodes glabrous-sparsely pubescent; leaves trifoliate, ± uniform in size throughout; stipules narrowly triangular, 8.5x2.8 mm, glabrous adaxially, glabrous pubescent abaxially, margins entire, sparsely pubescent, apices acute; petioles 20.6x≤0.3 mm, sparsely pubescent; petiolules ≤1.0x≤0.15 mm, glabrous-sparsely pubescent; leaflets ovoid-obcordate, glabrous adaxially, glabrous-sparsely pubescent along midrib abaxially, glabrous abaxially, bases acute, margins weakly serrate, glabrous-sparsely pubescent, apices rounded-emarginate, weakly mucronulate, terminal leaflets 7.6x5.3 mm; peduncles 30.8x≤0.3 mm, sparsely pubescent; inflorescences axillary, racemose-umbellate, 6.6x8 mm; flowers 4-8, each subtended by a linear triangular bract, in 1-2 whorls; pedicels ≤0.6x≤0.3 mm, sparsely pubescent; calyx 2.15 mm long, tube 3.2 mm in circumference, sparsely pubescent, lobes triangular, 2.15x0.5 mm, sparsely pubescent abaxially, margins entire, sparsely- moderately pubescent, apices acute; petals white; banner petals obcordate shallow notch, 4.85x5.5 mm; wing petals 4.45 mm long, asymmetrically clawed, claws 1.15x0.3 mm, lamina spatulate, 3.2 mm long, 1.5 mm wide at widest point, 1 mm wide at narrowest point, bases auriculate, auricles rounded, margins entire, apices rounded; keel petals 4 mm long, asymmetrically clawed, claws 1.3x0.5 mm, lamina spatulate, 2.7 mm long, 1.3 mm wide at widest point, 0.9 mm wide at narrowest point, bases truncate-shortly auriculate, auricles rounded, margins entire, apices acute; stamens diadelphous, connate filaments 3.4x1.3 mm, fused for 37% of their length, free filaments 2.2x≤0.1mm; pistils sessile, ovaries 2.2x0.7 mm, glabrous pubescent, styles 1.3x≤0.2 mm, glabrous, ovules 2; legumes not seen; seeds not seen.

Distribution and habitat Trifolium cochabambense is found in the province of Quillacollo in Cochabamba, Bolivia (Fig. 19). No information was given about its habitat on the label of the single specimen known.

Comments Trifolium cochabambense is easy to recognize with its near sessile flowers. It most closely resembles T. andesicola, because of their smaller inflorescences with a small number of flowers per inflorescence, but T. cochabambense differs in having nearly sessile flowers with a pedicel length of 0.52mm compared to 1.18-2.13mm in T. andesicola. Trifolium cochabambense was named for the area in which it was collected, Cochabamba, Bolivia.

Figure 18. Trifolium cochabambense (Hensen 569 (MU)).

Figure 19. Known geographical range of T. cochabambense, based on only known specimen (map generate using ESRI 2013).

Conclusion

Zohary and Heller (1984) recognized five varieties of Trifolium amabile: T. amabile Kunth var. amabile, found in Arizona in the United States, Mexico, Peru, Colombia, Bolivia, Guatemala, Chile, Argentina, and Ecuador; T. amabile Kunth var. hemsleyi (Lojac.) Heller & Zoh., found in Mexico, Peru, Ecuador, Argentina, Bolivia, Coast Rica, Colombia, and Guatemala; T. amabile Kunth var. pentlandii Ball, found in Peru and Bolivia; T. amabile var. mexicanum (Hemsl.) Heller & Zoh., found in Mexico, Guatemala, and Bolivia; and T. amabile Kunth var. longifoliolum Hemsl, found in Mexico. In-depth reexamination of specimens from the North and Central American range of the T. amabile species complex carried out by Ahlquist (2012) provided evidence that nine distinct species exist in this portion of the range: T. amabile Kunth, from Mexico, Guatemala and Costa Rica; T. blanquitum Ahlquist & Vincent (ined.), from Mexico; T. cognatum House, from Mexico; T. goniocarpum Lojac., from Mexico; T. hickeyi Ahlquist & Vincent (ined.), from Guatemala and Mexico; T. laciae Ahlquist & Vincent (ined.), from Mexico; T. lozani House, from Mexico; T. mexicanum Hemsl., from Mexico; and T. sonoranensis Ahlquist & Vincent, from Mexico and the United States (Arizona).

Research into the South American ranges of the T. amabile species complex carried out in this study provided evidence of six distinct species, and extended the range of T. amabile s. str. through South America in Argentina, Bolivia, Colombia, Ecuador, and Peru. Other species recognized in this study include: T. peruvianum, from Argentina, Bolivia, Ecuador, and Peru; T. tafiense, from Argentina; T. andesicola, from Argentina, Bolivia, and Peru; T. australe, from Argentina and Bolivia; and T. cochabambense, from Bolivia.

Additional field work in Colombia would be beneficial, since the fewest specimens in this species complex were collected in that country. Additional material would be helpful in clarifying the northern limits of T. peruvianum, for which specimens were seen from Ecuador but not from Colombia. More specimens from Colombia could also be used to better compare the northernmost range of the South American T. amabile s.str and the southernmost range of the North American populations. This could be used to further support the notion that both populations of T. amabile s.str are in fact the same species.

Another area that would benefit from more collections would be Cochabamba, Bolivia. Only one known specimen of T. cochabambense exists and more are needed to better understand the species boundaries and to confirm its existence as a species.

Ellison et al. (2006) used herbarium specimens for the molecular work that they carried out. The voucher specimen that they used for T. peruvianum was found in this study to actually be a part of T. amabile s. str and not what I have circumscribed as T. peruvianum. The voucher specimen cited by Ellison et al. 2006 as T. amabile could not be located, and therefore a determination of its true identity could not be made. Further molecular work needs to be done on the newly defined species as described in this paper and in Ahlquist (2012) in order to fully understand how they are related to each other.

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

Appendix 1. Table of specimens measured.

Species Tag # Barcode/Accession Collector T. peruvianum US14 23632 Rusby, H. H. 1015 T. peruvianum NY51 Pennell, F. W. 13596 T. peruvianum F121 557984 Pennell, F. W. 13596 T. peruvianum NY66 Buchtien, O. 8742 T. peruvianum MU27 000094187/268731 Loza de la Cruz, F. 104 T. peruvianum US171 603585 Cook, O. F. 356 & Gilbert, G. B. T. peruvianum US169 1059583 Watkins, C. s. n. T. peruvianum F118 557819 Pennell, F. W. 13393 T. peruvianum MO44 MO-1737315/5565670 Mercado, M. 2280 T. peruvianum P97 P00735620 Gay, M. C. 2038 T. peruvianum NY60 Beck, St. G. 2297 T. peruvianum MU26 000094185/268732 Valenzuela, E. 83 T. peruvianum US32 32892 Bang, M. 2819 T. peruvianum MO48 MO-1737319/04939491 Benyamini, D. 591 T. peruvianum NY119 Feuerer, T. 22733b T. peruvianum MO49 MO-1737318/04976101 Taylor, C. M. 11320 & Múlgura, M. & Careno, P. T. peruvianum MUX 000015344/221729 Cabrera, A. 30832 et al. T. peruvianum NYX Nicora, E. 9212 T. peruvianum US29 1057975 Holway, E. W. D. 485 & Holway, M. M. T. peruvianum NY50 Killip, E. P. 21972 Smith, A. C. T. peruvianum NY89 Kurtz, F. s. n. T. peruvianum US30 603552 Cook, O. F. 323 & Gilbert, G. B. T. peruvianum US28 1057962 Holway, E. W. D. 426 & Holway, M. M. T. tafiense F48 838890 Burkart, A. 5473 T. tafiense MU9 000035740/232469 Cialdella, A. M. 148 & Deginani, N. B. & Giussani, L. M. T. tafiense F131 1007423 Burkart, A. 5178 T. tafiense NY85 Bruch, C. s. n. T. andesicola LPB8 Schulte 135 T. andesicola MU10 000035739/232468 Cialdella, A. M. 310 & Deginani, N. B. & Giussani, L. M. T. andesicola NY104 Bastión, E. 1011 T. andesicola LPB11 Johns, T. 83-17 T. andesicola LPB19 Ehrich, R. 409 T. andesicola NY25 Solomon, J. C. 17782 T. andesicola B4 B 10 0358267 Villavicencio L., X. 285 T. andesicola LPB18 Fisel, U. 192 T. andesicola MU17 000091411/266515 Cialdella, A. M. 148 & Deginani, N. B. & Giussani, L. M. T. australe NY90 Olea, D. 1 T. australe US165 2220038 Spiaggi, L. 63 T. australe NY79 Feuerer, T. 8214b T. australe LPB4 Mostacedo, A. M. s. n. T. australe NY91 Venturi, S. 4764 T. australe F52 838902 Burkart, A. 5479 T. australe NY21 Lorentz 1057 & Hieronymus T. australe MU23 000094192/268726 Baar, R. 39 T. australe AHUC2 2464 Burkart, A. 11583 & Troncoso, N. S. T. australe LPB14 Bastión, E. 489 T. australe F57 955949 Hammarlund, C. 384

Appendix 1 continued.

Species Tag # Barcode/Accession Collector T. australe LPB15 Beck, St. G. 11956 T. australe LPB17 de Dios Romero C., J. 48 T. australe MU6 000089674/166816 Lewis, M. 37006 T. australe MO43 MO-1737317/5153453 Lewis, M. 37006 T. australe LPB6 Wood, J. R. I. 21781 et al. T. australe NY98 Beck, St. G. 11143 T. amabile US4 2100443 Tovar, O. 157 T. amabile P62 P00735624 Harlweg, C. 940 T. amabile P86 P00735601 Benoist, M. R. 2012 T. amabile P6 P00735597 Rivet, M. 86 T. amabile NY69 Holm-Nielsen, L. 5222 et al. T. amabile US8 2799987 Hudson, J. 1148 T. amabile US100 1208808 Rojas, R. T. 72 T. amabile US12 1776256 Penland, C. W. 569 & Summers, R. H. T. amabile F45 1015678 Penland, C. W. 569 & Summers, R. H. T. amabile AAU6 Elleman, L. 66986 T. amabile US2 1472155 Jamesen s. n. T. amabile F44 638092 Killip, E. P. 19514 & Smith, A. C. T. amabile F47 1426397 Prieto, F. E-2509 T. amabile US35 1364079 Lehman, F. C. 134a T. amabile MO10 MO-1737313/5335351 Peñafiel, M. 185 & Bersosa, F., Gaibor, P. & Toasa, G. T. amabile NY101 André, E. 917 T. amabile F135 2285464 Weigend, M. 8475 (MW 6_216) et al. T. amabile P83 P00735554 Reichler 225 T. amabile F91 2211271 Binder, M. 1990/80 et al. T. amabile F104 2150075 Sánchez V., I. 380 T. amabile MO37 MO-2050563/5878253 Rico, L. 1509 & Windsor-Shaw, T., Malavisi, A., Viscarra, J. & Remy, F. T. amabile F114 589505 Herrera, F. L. s. n. T. amabile F126 2055057 Asplund, E. 6799 T. amabile F109 517942 Macbride 1440 & Featherstone T. amabile US131 1592882 MacBride 821 & Featherstone T. amabile F122 534066 MacBride, J. F. 2996 T. amabile F111 518511 Macbride 2029 & Featherstone T. amabile F110 517778 MacBride 1268 & Featherstone T. amabile B11 B 10 0358260 Troll, C. 724 T. amabile NY122 [illegible] P377 T. amabile US13 1196611 Hitchcock, A. S. 22027 T. amabile US26 2482677 Barclay, H. G. 8053 & Juajibioy, P. T. amabile F124 1544064 Acosta Solís, M. 7580 T. amabile US20 1517748 P. Arbelaez, E. 1130 T. amabile US21 1185040 Ariste-Jospeh 966 T. amabile US129 1549668 Venturi, S. 3569 T. amabile F133 1900336 Davis, E. W. 1499 T. amabile MO32 MO-2040800/5952268 Villalobos, J. 379 & Paredes, M. & Villalobos, D. T. amabile MO19 MO-2136370/6111991 Villalobos, J. 934 & Jimenez, M. & Apaza, O. T. amabile US34 2103984 Cardenas 4924 T. amabile F98 517349 MacBride 821 & Featherstone

Appendix 1 continued.

Species Tag # Barcode/Accession Collector T. amabile US132 1549669 Venturi, S. 3642 T. amabile LPB3 Thomas, E. 97 T. amabile NY97 Sánchez V., J. 55 & Cabanillas, J. T. amabile P8 P00735626 Gay, M. C. 365 T. amabile MU19 000091413/266518 Zuloaga, F. O. 10745 & Aagesen, L., Biganzoli, F., Panizza, A. T. amabile NY63 Rusby, H. H. 1015 T. amabile SI6 Krapovickas, A 7073 & Fuchs de K., A. M. T. amabile LPB22 Beck, St. G. 8007 T. amabile US19 1875884 Metcalf, R. D. 30441 T. amabile MU21 000094189/268730 Vargas, E. 60 T. amabile NY24 Solomon, J. C. 15237 T. amabile MO3 MO-1737309/5290233 Quipuscoa, V. 574 & Angulo, O. & Yahuana R. T. amabile F2 2149952 Sánchez Vega, I. M. 6099 T. amabile F90 1903161 Dillon, M. 1567 & Turner, B. L. T. cochabambense MU34 000094174/268739 Hensen, I. 569

Appendix 2. Table of characters measured for selected OTU. Table courtesy of Ahlquist (2012).

Abbreviation Character Description

Habit Habit Qualitative description of the overall habit of the plant.

Roots Root Qualitative description of the taproot (woody, large/small, etc.).

Root Diameter RootDiam* Measurement of the taproot at 1cm below the crown. Qualitative description of the stems (erect, ascending, prostrate, Stems Stem§ etc.) based on the lable data. Measurement of the length of the stem from the crown to the apex Stem Length StemL* of the terminal leaf or inflorescence of the longest stem.

Stem Diameter StemD* Measurement of the diameter of the stem at 1cm above the crown.

Internode Pubescence InNodPub† Qualitative description of the pubescence of the internodes. Measurement of the internode between the first and second nodes Internode Length InNodL* from the crown, or the basal most nodes if crown was absent.

Node Pubescence NodPub† Qualitative description of the pubescence of the nodes. Measurement of the length of the stipules from the base to the Stipule Length StipL* apex.

Stipule Width StipW* Measurement of the width of the stipules at the widest point.

Stipule Shape StipShp‡ Qualitative description of the shape of the stipules. Qualitative description of the pubescence of the adaxial surface of Stipule Adaxial Pubescence StipAdPub† the stipules. Qualitative description of the pubescence of the abaxial surface of Stipule Abaxial Pubescence StipAbPub† the stipules. Qualitative description of the margin of the stipules (entire, Stipule Margins StipMar§ serrate). Qualitative description of the pubescence of the margin of the Stipule Margin Pubescence StipMarPub† stipules.

Stipule Apex Angle StipApxAng Measurement of the apex angle of the stipules. Measurement of the length of the petioles from the base to the Petiole Length PetL* apex.

Petiole Diameter PetD* Measurement of the diameter of the petioles.

Petiole Pubescence PetPub† Qualitative description of the pubescence of the petioles.

Leaflet Shape LfltShp‡ Qualitative description of the shape of the leaflets. Qualitative description of the pubescence of the adaxial surface of Leaflet Adaxial Pubescence LfltAdPub† the leaflets. Leaflet Abaxial Midrib Qualitative description of the pubescence of the abaxial surface of LfltAbMdrbPub† Pubescence the midrib. Qualitative description of the pubescence of the abaxial surface of Leaflet Abaxial Pubescence LfltAbPub† the leaflets.

Appendix 2 continued.

Abbreviation Character Description

Leaflet Base Angle LfltBasAng Measurement of the base angle of the leaflets.

Leaflet Margins LfltMar§ Qualitative description of the margin of the leaflets. Qualitative description of the pubescence of the margin of the Leaflet Margin Pubescence LfltMarPub† leaflets.

Leaflet Apex Angle LfltApxAng Measurement of the apex angle of the leaflets.

Leaflet Mucronulate LfltMucr Qualitative description of the mucron of the leaflets. Measurement of the length of the terminal leaflets from the base to Terminal Leaflet Length TermLfltL* the most distal point of the lamina. Measurement of the width of the terminal leaflets at the widest Terminal Leaflet Width TermLfltW* point.

Petiolule Length PetlleL* Measurement of the length of petiolule from the base to the apex.

Petiolule Diameter PetlleD* Measurement of the diameter of the petiolules.

Petiolule Pubescence PetllePub† Qualitative description of the pubescence of the petiolules.

Inflorescence Infl Qualitative description of the type of inflorescence. Measurement of the length of the inflorescence parallel to the Inflorescence Length InflL* peduncle from the tip of the basalmost to the tip of the distalmost Measurement of the width of the inflorescence perpendicular to Inflorescence Width InflW* the peduncle from tip to tip of the lateral flowers.

Flower # FlNo Count of the number of flowers in an inflorescence. Qualitative description of the bracts subtending the flowers of the Bracts Bract inflorescence. Count of the number of whorls of flowers comprising the Whorls Whorl inflorescences.

Peduncle Length PeduncL* Measurement of the length of peduncles from the base to the apex.

Peduncle Diameter PeduncD* Measurement of the diameter of the peduncles.

Peduncle Pubescence PeduncPub† Qualitative description of the pubescence of the peduncles.

Pedicel Length PedicL* Measurement of the length of pedicels from the base to the apex.

Pedicel Diameter PedicD* Measurement of the diameter of the pedicels.

Pedicel Pubescence PedicPub† Qualitative description of the pubescence of the pedicels. Measurement of the length of the sepals from the base of the tube Sepal Length SepL* to the apex of the lobes. Measurement of the circumference of the opened sepal tube at the Sepal Circumference SepCir* base of the lobes.

Appendix 2 continued.

Abbreviation Character Description

Sepal Tube Pubescence SepTubPub† Qualitative description of the pubescence of the fused sepal tubes.

Sepal Lobes Shape SepLobShp‡ Qualitative description of the shape of the sepal lobes. Measurement of the length of the free sepal lobes from the base to Sepal Lobe Length SepLobL* the apex.

Sepal Lobe Width SepLobW* Measurement of the width of the free sepal lobes at the base.

Sepal Lobe Pubescence SepLobPub† Qualitative description of the pubescence of the free sepal lobes.

Sepal Lobe Margins SepLobMar§ Qualitative description of the margin of the free sepal lobes. Sepal Lobe Margin Qualitative description of the pubescence of the margins of the free SepLobMarPub† Pubescence sepal lobes.

Sepal Lobe Apex Angle SepLobApxAng Measurement of the apex angle of the sepal lobes Qualitative description of the color of the flower petals based on Petals Color PetCol the label data.

Banner Petal Shape BPetShp‡ Qualitative description of the shape of the banner petals. Measurement of the length of the banner petal from the base to the Banner Petal Length BPetL* apex.

Banner Petal Width BPetW* Measurement of the width of the banner petal at the widest point. Measurement of the wing petals from the base of the claw to the Wing Petal Length WPetL* apex of the lamina. Measurement of the length of the claw of the wing petals from the Wing Petal Claw Length WPetClwL* base of the claw to the base of the lamina.

Wing Petal Claw Width WPetClwW* Measurement of the width of the claw of the wing petals. Qualitative description of the shape of the lamina of the wing Wing Petal Lamina WPetLam§ petals. Measurement of the length of the lamina of the wing petals from Wing Petal Lamina Length WPetLamL* the base to the apex of the lamina. Wing Petal Lamina Width Measurement of the width of the lamina of the wing petals at the WPetLamWw* W widest point. Wing Petal Lamina Width Measurement of the width of the lamina of the wing petals at the WPetLamWn* N narrowest point. Qualitative description of the shape of the base of the lamina of the Wing Petal Lamina Base WPetLamBas§ wing petal. WPetLamArcAp Wing Petal Lamina Auricle xAng Apex Angle Measurement of the apex angle of the auricle of the lamina. Qualitative description of the margin of the lamina of the wing Wing Petal Lamina Margin WPetLamMar§ petals.

Wing Petal Lamina Apex WPetLamApx§ Qualitative description of the wing petal lamina apices. Measurement of the keel petals from the base of the claw to the Keel Petal Length KPetL* apex of the lamina.

Appendix 2 continued.

Abbreviation Character Description Measurement of the length of the claw of the keel petals from the Keel Petal Claw Length KPetClwL* base of the claw to the base of the lamina.

Keel Petal Claw Width KPetClwW* Measurement of the width of the claw of the keel petals. Qualitative description of the shape of the lamina of the keel Keel Petal Lamina KPetLam§ petals. Measurement of the length of the lamina of the keel petals from Keel Petal Lamina Length KPetLamL* the base to the apex of the lamina. Measurement of the width of the lamina of the keel petals at the Keel Petal Lamina Width W KPetLamWw* widest point. Measurement of the width of the lamina of the keel petals at the Keel Petal Lamina Width N KPetLamWn* narrowest point. Qualitative description of the shape of the base of the lamina of the Keel Petal Lamina Base KPetLamBas§ keel petal. Qualitative description of the margin of the lamina of the keel Keel Petal Lamina Margin KPetLamMar§ petals.

Keel Petal Lamina Apex KPetLamApx§ Qualitative description of the keel petal lamina apices. Measurement of the length of the connate filaments from the base Connate Filaments Length ConFilL* to the apex of the longest filament. Measurement of the width of the connate filaments at the widest Connate Filaments Width ConFilW* point. Connate Filament Percent ConFilPcFus Fusion The proportion of the connate filaments that is fused.

Free Filament Length FreFilL* Measurement of the length of the free filament.

Free Filament Diameter FreFilD* Measurement of the width of the free filament at the widest point. Measurement of the length of the ovary from the base of the ovary Ovary Length OvL* to the base of the style.

Ovary Width OvW* Measurement of the width of the ovary at the widest point.

Ovary Pubescence OvPub† Qualitative description of the pubescence of the ovaries. Measurement of the length of the style from the apex of the ovary Style Length StylL* to the base of the stigma.

Style Diameter StylD* Measurement of the diameter of the style.

Style Pubescence StylPub† Qualitative description of the pubescence of the styles.

Ovule Number OvlNo Count of the number of ovules in the ovary. Measurement of the length of the fruit from the base of the fruit to Fruit Length FruitL* the base of the style remnant.

Fruit Width FruitW* Measurement of the width of the fruit at the widest point.

Seeds Per Fruit SeedpFruit Count of the number of seeds in the fruit.

Appendix 2 continued.

Abbreviation Character Description Qualitative description of the shape of the seeds based on Gillett et Seed Shape SeedShp al., 2001. Qualitative description of the color of the seeds based on Kornerup Seed Color SeedCol and Wanscher, 1978.

Seed Length SeedL* Measurement of the length of the seeds.

Seed Width SeedW* Measurement of the width of the seeds. * Measurements taken in millimeters. † sparse= 1mm≤ between hairs, very sporadic; moderate= <1mm between haris, uniformly spaced, able to see the underlying structure beneath the hairs; dense= so many hairs that the underlying structure is not visible ‡ based on Systematics Association Committee for Descriptive Biological Terminology, 1962. § based on Harris and Harris, 2001.

Appendix 3. Table of qualitative data used for analysis.

OTUs StipL* StipW* TermLfltL* TermLfltW* PedicL* PeduncL* PetL* PedPetR* InflL* InflW* US14 15.10 5.20 13.70 8.60 3.64 35.40 36.40 0.97 15.50 16.50 NY51 10.00 3.67 8.40 5.35 2.22 12.60 17.00 0.74 15.10 15.50 F121 10.60 3.70 9.00 6.20 3.22 9.60 15.88 0.60 14.70 15.90 NY66 12.10 4.90 12.40 7.90 4.18 16.00 28.60 0.56 15.70 16.60 MU27 11.90 4.70 11.10 7.20 2.38 5.30 19.80 0.27 14.40 14.60 US171 9.33 3.50 9.83 7.33 2.07 7.33 35.00 0.21 12.33 12.33 US169 9.40 3.10 8.60 5.85 2.18 3.70 23.80 0.16 11.20 12.10 F118 9.10 2.70 5.70 3.40 1.76 4.67 17.70 0.26 10.50 10.00 MO44 10.67 2.10 6.33 4.83 2.10 8.83 21.67 0.41 11.50 11.50 P97 9.90 3.40 8.00 5.05 2.08 17.00 15.20 0.95 10.33 11.50 NY60 11.90 4.10 11.20 7.30 2.12 13.75 22.60 0.61 12.25 13.63 MU26 15.10 5.30 13.40 7.80 1.84 4.50 40.50 0.11 13.20 12.30 US32 16.50 4.40 15.20 9.30 2.72 18.60 26.20 0.71 14.70 14.40 MO48 7.40 2.90 7.00 5.10 1.38 7.90 9.30 0.85 9.90 9.10 NY119 9.70 3.70 8.60 6.20 2.64 17.00 16.80 1.01 11.10 13.70 MO49 6.67 1.90 7.00 4.83 2.00 8.00 17.67 0.45 10.83 11.33 MUX 8.33 2.60 8.83 6.00 1.53 34.33 26.33 1.30 11.75 11.75 NYX 10.17 4.50 10.00 8.75 2.73 44.33 18.00 2.46 15.50 14.50 US29 12.83 5.40 14.17 8.00 2.20 51.00 28.33 1.80 15.50 16.33 NY50 7.00 1.90 11.50 5.00 1.90 10.00 17.00 0.59 13.00 12.00 NY89 8.70 3.10 8.90 5.80 2.24 16.20 18.90 0.86 10.80 10.30 US30 9.30 3.20 9.60 6.20 3.22 22.33 22.33 1.00 15.00 14.00 US28 11.40 5.80 12.90 7.30 3.28 38.80 11.20 3.46 19.20 17.60 F48 10.80 3.40 10.05 7.70 1.60 29.90 25.80 1.16 10.80 11.30 MU9 8.90 3.90 9.00 4.90 1.70 38.40 18.20 2.11 15.10 15.10 F131 14.67 6.00 15.00 10.00 1.80 43.33 27.33 1.59 15.50 16.00 NY85 12.00 4.75 11.33 5.50 1.70 31.50 17.00 1.85 13.50 13.50 LPB8 6.80 2.75 4.85 2.80 1.80 9.00 9.20 0.98 8.60 8.80 MU10 6.17 2.50 5.17 3.00 2.13 14.00 7.83 1.79 12.00 11.50 NY104 5.50 1.70 4.70 3.15 1.18 9.10 11.50 0.79 11.00 11.00 LPB11 6.30 2.60 5.40 2.70 1.66 10.40 7.80 1.33 11.80 10.60 LPB19 6.00 2.10 6.33 3.75 1.70 9.67 12.33 0.78 11.00 10.67 NY25 6.33 3.00 6.33 2.80 1.63 15.67 8.67 1.81 12.83 13.17 B4 6.25 2.40 6.40 3.20 1.30 14.60 9.00 1.62 10.70 9.90 LPB18 6.33 2.30 5.67 2.50 1.60 14.67 7.17 2.05 11.00 11.50

Appendix 3 continued.

OTUs StipL* StipW* TermLfltL* TermLfltW* PedicL* PeduncL* PetL* PedPetR* InflL* InflW* MU17 5.83 2.00 6.33 3.00 1.57 20.5 7.00 2.93 7.75 8.25 NY90 9.20 3.00 8.20 5.80 1.68 25.67 7.30 3.52 10.63 11.13 US165 9.30 3.70 9.70 6.20 2.00 41.00 10.60 3.87 11.00 13.33 NY79 8.33 3.80 10.50 5.75 2.20 30.00 8.17 3.67 9.75 11.00 LPB4 9.40 3.70 10.60 5.60 2.96 50.50 16.70 3.02 9.63 11.38 NY91 12.10 3.30 8.90 6.10 1.56 47.20 9.80 4.82 9.30 10.50 F52 8.20 3.40 6.90 4.30 1.48 16.00 9.50 1.68 8.20 8.30 NY21 13.60 4.67 17.60 11.00 2.10 48.40 25.80 1.88 11.13 11.50 MU23 16.67 6.00 18.67 10.75 3.50 53.00 15.67 3.38 14.00 14.17 AHUC2 14.90 4.40 13.30 6.90 2.44 48.75 28.20 1.73 9.38 10.25 LPB14 12.50 3.60 10.00 5.40 1.36 35.80 20.60 1.74 10.20 10.70 F57 10.40 4.70 14.00 7.40 2.74 44.40 11.40 3.89 13.30 13.40 LPB15 14.60 4.60 13.40 7.30 1.44 41.75 27.00 1.55 12.00 12.67 LPB17 13.00 3.70 12.70 6.70 1.84 27.80 28.80 0.97 8.60 9.40 MU6 15.60 3.45 15.00 11.00 1.52 78.00 33.60 2.32 14.38 14.75 MO43 12.90 4.10 10.50 7.35 2.18 58.20 11.80 4.93 15.00 14.80 LPB6 16.50 5.75 18.00 9.00 2.10 56.00 9.33 6.00 13.50 15.17 NY98 9.50 4.40 8.60 5.10 2.50 27.60 13.60 2.03 13.70 14.00 US4 10.20 3.45 11.20 7.30 2.36 49.20 12.00 4.10 14.10 15.20 P62 8.90 3.30 9.40 7.40 2.50 32.80 9.90 3.31 13.80 13.90 P86 11.00 3.70 12.00 9.20 3.50 36.80 10.30 3.57 14.90 13.90 P6 10.40 3.75 8.80 7.70 3.00 23.40 17.60 1.33 13.20 13.10 NY69 11.20 3.30 10.90 8.90 3.36 25.40 49.00 0.52 12.00 11.50 US8 10.10 2.40 8.70 6.60 2.92 38.40 29.00 1.32 9.80 13.10 US100 9.60 2.90 10.60 7.70 2.04 28.00 27.40 1.02 12.80 12.20 US12 9.80 3.20 9.20 9.70 2.40 33.80 17.20 1.97 16.10 13.30 F45 10.40 2.90 11.75 9.80 2.56 56.60 25.20 2.25 10.33 11.00 AAU6 9.10 3.00 8.70 7.50 2.30 26.70 12.70 2.10 11.10 9.20 US2 10.20 3.20 9.30 8.80 2.72 30.20 20.60 1.47 10.60 11.70 F44 7.70 2.00 5.00 4.70 1.50 15.60 14.10 1.11 7.38 6.75 F47 8.50 2.40 6.70 5.30 1.62 19.30 11.80 1.64 8.40 10.50 US35 11.10 3.67 8.10 7.50 1.82 38.40 45.60 0.84 7.30 9.90 MO10 10.70 3.20 7.40 6.70 1.76 27.80 19.80 1.40 7.80 10.00 NY101 9.10 2.10 5.40 4.50 1.04 20.67 19.00 1.09 8.83 11.17 F135 11.50 3.00 10.80 5.60 2.18 50.60 23.40 2.16 11.80 15.00

Appendix 3 continued.

OTUs StipL* StipW* TermLfltL* TermLfltW* PedicL* PeduncL* PetL* PedPetR* InflL* InflW* P83 10.10 3.30 13.20 6.20 2.86 86.80 35.20 2.47 13.00 13.33 F91 11.90 4.70 13.40 6.80 1.68 55.00 14.50 3.79 10.83 15.50 F104 10.70 4.00 11.80 4.90 2.08 54.40 17.80 3.06 10.50 13.30 MO37 7.80 3.00 9.40 5.25 2.37 30.00 17.20 1.94 10.40 11.70 F114 6.50 2.80 7.00 4.60 2.48 22.20 8.90 2.49 11.83 12.67 F126 10.30 3.10 11.50 9.70 2.30 44.40 24.60 1.80 9.00 13.40 F109 11.50 3.70 13.20 9.20 1.92 55.60 16.70 3.33 12.40 13.50 US131 11.67 5.00 13.67 7.50 2.53 40.67 19.00 2.14 15.17 16.17 F122 8.17 3.00 6.83 6.17 1.30 11.67 22.17 0.53 12.00 12.33 F111 9.10 4.10 12.20 9.80 3.60 37.20 16.50 2.25 15.60 19.50 F110 11.00 4.50 11.50 7.80 2.26 28.80 33.20 0.87 15.80 17.10 B11 9.90 3.60 7.25 4.63 2.00 22.60 9.00 2.51 14.40 14.50 NY122 10.40 4.50 11.00 6.00 2.60 34.80 13.40 2.60 16.00 16.00 US13 10.30 3.60 5.70 4.20 1.14 8.60 13.40 0.64 11.90 11.90 US26 9.25 2.75 5.35 3.80 1.50 6.00 12.40 0.48 10.83 11.67 F124 9.80 4.20 10.40 8.20 1.54 15.10 9.90 1.53 15.70 14.90 US20 10.80 3.95 9.90 8.20 2.02 33.50 30.80 1.09 15.00 12.83 US21 10.30 3.20 9.80 6.40 1.70 18.00 16.00 1.13 11.70 12.10 US129 13.50 3.50 9.20 5.70 1.86 41.20 18.90 2.18 12.80 11.90 F133 14.40 5.00 14.90 10.60 2.82 64.40 19.60 3.29 16.33 17.00 MO32 16.30 6.10 20.20 10.20 2.94 82.80 32.80 2.52 14.40 17.40 MO19 17.85 6.30 17.00 9.00 1.98 58.80 25.40 2.31 16.40 15.70 US34 18.10 6.00 11.90 8.50 2.64 70.20 41.20 1.70 18.80 18.40 F98 11.60 5.90 14.80 6.90 2.60 48.20 31.67 1.52 17.90 17.00 US132 12.30 4.60 11.00 8.20 1.72 39.80 27.00 1.47 11.50 12.40 LPB3 17.40 4.50 13.10 6.80 2.28 49.50 49.80 0.99 11.25 13.13 NY97 18.40 5.50 16.00 10.70 2.82 68.60 53.40 1.28 14.80 14.60 P8 16.30 3.90 13.00 8.60 2.24 60.00 57.60 1.04 14.00 15.75 MU19 15.60 4.30 12.80 7.40 1.96 43.80 30.00 1.46 14.20 13.60 NY63 16.40 4.90 14.80 9.60 2.34 63.00 60.20 1.05 11.90 13.60 SI6 13.40 3.20 16.00 6.90 2.50 24.40 32.20 0.76 13.40 12.80 LPB22 13.80 4.50 10.80 8.70 3.28 22.00 33.75 0.65 15.00 15.70 US19 12.40 3.70 10.80 8.40 2.26 24.40 41.20 0.59 13.30 13.10 MU21 14.33 5.00 17.83 10.50 3.50 45.00 50.00 0.90 17.00 18.00 NY24 13.20 5.00 18.50 12.90 3.96 58.20 60.00 0.98 14.90 13.70

Appendix 3 continued.

OTUs StipL* StipW* TermLfltL* TermLfltW* PedicL* PeduncL* PetL* PedPetR* InflL* InflW* MO3 13.67 5.50 22.17 15.50 2.13 59.33 52.33 1.13 15.00 17.00 F2 10.00 3.50 10.30 5.80 2.52 78.80 15.10 5.22 13.60 12.20 F90 9.33 3.20 8.33 4.50 2.07 32.00 3.83 8.35 15.17 15.50 MU34 8.50 2.80 7.60 5.30 0.52 30.80 20.60 1.50 6.60 8.00

Appendix 3 continued.

OTUs SepLobL* SepLobW* SepL* SepCir* BPetL* BPetW* FlNo ConFilL* ConFilW* US14 3.55 1.00 5.95 4.70 7.55 7.80 21.60 4.55 1.35 NY51 3.20 0.90 5.15 4.60 7.15 6.80 14.40 3.60 1.45 F121 3.25 0.90 5.35 4.65 6.90 7.40 17.60 3.90 1.55 NY66 3.05 0.95 5.70 5.35 6.45 6.10 20.00 4.45 1.40 MU27 2.95 1.00 5.45 5.30 5.70 6.00 18.60 4.15 1.45 US171 2.35 0.70 4.20 4.00 5.85 7.90 13.00 3.50 1.40 US169 2.65 0.85 4.60 4.80 5.35 6.35 9.60 3.80 1.65 F118 2.35 1.00 3.80 5.05 5.60 5.85 8.60 3.60 1.60 MO44 2.90 1.25 4.50 4.70 5.75 5.75 8.50 3.40 1.60 P97 3.00 0.95 4.80 4.55 6.40 5.70 10.00 4.15 1.40 NY60 3.85 1.45 6.55 5.65 5.65 7.75 9.50 3.55 1.50 MU26 3.15 1.25 6.05 6.05 5.95 6.45 12.80 4.00 1.65 US32 4.05 1.00 6.00 5.65 6.35 7.80 21.00 3.75 1.40 MO48 2.10 0.70 4.25 3.95 5.10 3.85 7.40 3.70 1.70 NY119 3.50 0.70 5.30 4.45 6.70 5.35 14.20 3.95 1.55 MO49 1.95 0.85 3.60 3.80 5.65 5.80 17.00 3.30 1.15 MUX 2.10 0.70 3.60 4.10 5.30 6.60 14.00 3.30 1.20 NYX 2.65 1.00 4.50 4.60 6.20 6.50 14.00 3.40 1.30 US29 3.00 0.80 4.50 4.50 6.10 7.05 24.00 3.80 1.50 NY50 2.35 0.80 4.20 2.80 5.35 4.00 9.70 3.20 1.05 NY89 2.75 0.60 4.40 3.75 5.50 4.60 10.20 3.60 1.30 US30 2.80 0.60 4.25 4.05 6.90 6.70 17.50 4.45 1.75 US28 2.90 0.75 4.45 4.50 7.70 6.95 21.20 4.05 1.80 F48 3.55 0.80 4.90 4.75 6.40 9.95 8.25 3.55 1.55 MU9 2.70 0.65 4.25 4.75 6.70 9.25 23.60 3.60 1.20 F131 2.85 1.05 5.20 5.40 6.75 9.95 17.00 3.80 1.40 NY85 2.43 0.95 4.20 4.70 6.10 9.05 23.00 3.40 1.30 LPB8 2.05 0.65 4.00 3.65 4.80 3.80 6.80 3.50 1.15 MU10 2.40 0.60 3.90 3.50 6.35 5.00 12.50 4.20 1.30 NY104 2.03 0.60 3.67 4.00 5.00 4.27 11.00 3.63 1.20 LPB11 2.15 0.70 4.30 4.10 5.85 4.00 12.60 3.80 1.15 LPB19 1.95 0.65 3.30 3.40 4.85 3.90 8.00 3.30 1.20 NY25 2.00 0.68 3.30 3.40 5.95 4.50 13.00 3.70 1.10 B4 2.80 0.65 4.40 3.70 4.90 3.90 11.40 3.30 1.00 LPB18 2.85 0.80 4.20 3.70 5.20 4.45 8.40 3.70 1.00

Appendix 3 continued.

OTUs SepLobL* SepLobW* SepL* SepCir* BPetL* BPetW* FlNo ConFilL* ConFilW* MU17 2.00 0.63 2.70 2.80 5.45 4.80 10.00 3.40 1.40 NY90 2.80 0.50 4.90 2.90 5.10 4.55 19.25 3.60 1.30 US165 3.25 0.50 5.15 3.70 7.55 5.45 16.67 4.05 1.55 NY79 2.65 0.48 4.30 3.30 5.40 3.90 13.00 3.80 1.40 LPB4 2.40 0.50 3.90 3.30 5.20 4.20 16.00 3.30 1.30 NY91 2.45 0.55 3.60 2.95 4.50 3.40 19.80 3.10 1.30 F52 1.60 0.45 3.15 2.80 3.60 3.80 16.00 2.80 1.00 NY21 2.48 0.52 4.06 3.15 4.54 3.74 27.00 3.28 1.22 MU23 3.25 0.80 4.70 4.30 5.05 5.25 40.00 3.30 1.40 AHUC2 3.05 0.35 4.40 2.90 3.85 3.10 25.25 3.05 1.30 LPB14 3.00 0.40 4.55 2.95 4.10 3.60 25.20 2.65 1.25 F57 3.50 0.40 5.25 3.10 4.90 3.70 24.40 3.45 1.15 LPB15 3.70 0.40 5.30 3.40 4.90 4.20 23.33 3.70 1.40 LPB17 2.75 0.45 4.40 3.05 5.05 4.40 19.40 2.95 0.75 MU6 3.40 0.45 4.85 2.90 6.55 4.95 31.75 4.30 1.00 MO43 3.40 0.65 4.95 3.05 6.60 5.05 35.20 4.25 1.15 LPB6 3.55 0.50 5.40 3.40 7.70 5.30 24.00 3.90 1.40 NY98 2.93 0.50 4.80 3.63 5.23 4.07 23.00 3.37 1.50 US4 2.65 0.50 4.45 3.60 7.25 5.45 16.00 4.70 1.45 P62 2.05 0.40 3.90 3.15 5.75 3.35 12.20 4.10 1.50 P86 3.75 0.50 5.65 3.65 7.40 4.65 13.20 4.55 1.85 P6 2.90 0.50 5.25 3.05 7.10 4.65 8.80 4.10 1.40 NY69 2.34 0.60 4.60 3.90 6.24 3.90 7.50 4.10 1.78 US8 2.85 0.50 4.60 3.50 6.95 4.20 7.80 4.00 1.65 US100 2.70 0.35 4.40 2.90 7.25 4.20 9.00 3.90 1.60 US12 3.55 0.75 6.40 4.30 9.55 6.05 7.00 5.50 1.55 F45 3.45 0.70 5.45 3.65 7.15 5.35 7.00 5.10 1.50 AAU6 2.90 0.55 5.10 3.55 7.05 4.85 8.40 4.20 1.65 US2 2.80 0.60 5.80 3.45 6.90 4.70 8.00 3.85 1.65 F44 2.10 0.60 4.05 3.60 6.85 4.40 3.50 3.70 1.30 F47 2.50 0.60 4.35 3.95 6.65 4.90 6.00 3.75 1.65 US35 4.00 0.63 5.80 4.17 7.17 4.20 4.60 4.10 1.80 MO10 3.13 0.53 5.20 3.53 6.93 4.20 5.40 3.95 1.55 NY101 3.10 0.50 5.10 3.40 6.90 4.65 6.33 3.65 1.30 F135 3.55 0.50 5.45 3.50 7.55 5.10 13.60 3.90 1.60

Appendix 3 continued.

OTUs SepLobL* SepLobW* SepL* SepCir* BPetL* BPetW* FlNo ConFilL* ConFilW* P83 3.40 0.45 5.05 3.40 8.20 5.70 11.33 4.40 1.60 F91 3.55 0.65 5.85 3.70 8.15 5.25 12.33 4.70 1.60 F104 4.20 0.65 5.75 3.65 7.70 5.35 8.60 4.75 1.60 MO37 2.95 0.60 4.65 4.00 6.90 5.45 10.00 4.35 1.55 F114 2.75 0.50 4.35 3.45 7.00 5.00 13.00 4.40 1.55 F126 3.57 0.53 5.57 3.97 6.83 4.27 9.40 3.97 1.77 F109 4.80 0.55 6.70 3.90 7.50 5.15 8.20 4.20 1.85 US131 3.45 0.90 4.80 5.10 7.85 6.05 14.00 4.30 1.65 F122 3.00 0.83 5.10 3.70 6.75 4.65 10.00 4.55 1.40 F111 3.95 1.00 6.35 5.30 10.30 6.60 11.60 5.20 2.00 F110 3.37 0.63 5.97 3.83 8.63 6.23 16.40 5.03 1.73 B11 2.10 0.65 4.25 4.25 8.10 6.30 17.40 4.70 1.70 NY122 2.90 0.80 5.00 4.25 9.10 6.35 15.00 5.15 1.65 US13 1.80 0.65 3.70 3.80 7.00 4.35 12.40 4.20 1.65 US26 2.10 0.60 4.00 3.70 5.80 3.40 10.00 4.00 1.60 F124 2.60 0.50 4.85 3.60 7.20 4.40 13.20 4.25 1.80 US20 2.72 0.76 4.62 4.16 7.22 5.06 10.00 3.60 1.64 US21 2.85 0.60 5.15 3.75 6.70 4.50 9.20 3.15 1.95 US129 3.05 0.55 4.85 3.45 6.15 4.40 14.00 3.65 1.45 F133 3.72 0.72 5.68 4.02 8.12 4.86 14.40 4.02 1.62 MO32 3.65 0.55 5.65 3.85 8.00 6.40 23.40 4.95 1.60 MO19 3.30 0.60 5.20 3.50 8.15 4.70 16.40 4.85 1.30 US34 3.42 0.66 5.26 3.50 8.34 5.80 20.20 4.62 1.60 F98 3.36 0.72 5.38 3.68 7.86 5.64 13.20 5.04 1.74 US132 2.95 0.65 4.80 3.45 6.30 5.25 10.80 4.05 1.60 LPB3 2.95 0.70 5.25 4.30 7.15 5.40 12.75 4.35 1.65 NY97 3.55 0.65 5.40 3.90 8.20 5.00 17.00 4.30 1.65 P8 3.00 0.60 4.90 3.60 6.90 4.85 15.25 4.20 1.50 MU19 3.10 0.67 4.76 4.00 6.76 4.94 15.00 3.88 1.46 NY63 4.70 1.05 6.95 5.70 7.45 6.80 11.80 4.00 1.30 SI6 2.85 0.70 6.15 4.15 6.80 4.60 11.20 3.95 1.05 LPB22 4.05 0.65 6.25 3.85 7.15 6.20 19.00 4.45 1.50 US19 3.55 0.60 5.15 3.60 6.85 4.90 14.00 4.10 1.40 MU21 5.40 0.65 6.90 3.50 6.55 5.85 26.00 3.90 1.20 NY24 4.10 0.60 6.05 3.80 6.35 5.85 14.80 3.75 1.40

Appendix 3 continued.

OTUs SepLobL* SepLobW* SepL* SepCir* BPetL* BPetW* FlNo ConFilL* ConFilW* MO3 4.35 0.45 5.40 3.20 8.95 5.35 13.00 4.10 1.65 F2 2.70 0.40 4.45 3.35 6.45 4.45 17.00 4.00 1.55 F90 2.70 0.75 4.20 4.50 8.35 6.75 14.00 4.30 1.40 MU34 2.15 0.50 3.55 3.20 4.85 5.50 8.40 3.35 1.25

Appendix 3 continued.

OTUs WPetL* WPetLamWw* WPetClwL* WPetLamWn* KPetL* KPetLamWw* KPetClwL* KPetLamWn* US14 6.55 2.20 1.95 1.25 5.70 2.00 2.10 1.15 NY51 5.85 2.05 1.75 1.25 5.05 1.70 1.95 1.20 F121 6.00 2.15 1.85 1.15 5.25 1.80 2.00 1.00 NY66 6.20 1.90 2.00 1.45 5.45 1.85 2.35 1.20 MU27 5.40 1.40 2.00 1.30 5.05 1.40 2.20 1.15 US171 5.10 1.65 1.75 1.00 4.55 1.45 1.85 0.95 US169 4.70 1.70 1.60 1.35 4.40 1.60 1.90 1.35 F118 5.05 1.40 1.80 1.15 4.70 1.50 2.00 1.15 MO44 5.10 1.65 1.60 1.30 4.20 1.40 1.85 1.00 P97 6.05 1.75 2.00 1.15 5.20 1.40 2.10 1.15 NY60 5.60 2.05 1.65 1.50 5.10 1.80 1.95 1.30 MU26 5.40 1.75 1.85 1.30 5.25 1.60 2.05 1.20 US32 5.55 2.10 1.80 1.60 4.85 1.60 1.90 1.30 MO48 4.90 1.45 1.40 1.05 4.35 1.45 1.60 1.10 NY119 5.80 1.75 1.85 1.25 5.50 1.75 2.05 1.30 MO49 4.40 1.55 1.45 1.20 3.85 1.35 1.60 1.05 MUX 4.45 1.35 1.70 1.20 4.15 1.40 1.60 1.20 NYX 5.85 1.60 1.70 1.30 5.20 1.80 1.70 1.30 US29 5.35 2.05 1.60 1.30 4.70 1.60 1.80 1.20 NY50 4.30 1.20 1.50 0.90 4.05 1.20 1.80 0.80 NY89 4.80 1.40 1.60 1.10 4.65 1.30 1.90 1.00 US30 5.65 2.00 1.80 1.20 5.05 1.80 1.70 1.20 US28 6.90 2.15 1.85 1.35 5.55 1.95 1.85 1.20 F48 4.70 2.30 1.30 2.00 4.00 2.60 1.30 1.50 MU9 4.90 1.80 1.70 1.60 4.75 1.85 1.60 1.55 F131 4.35 1.90 1.50 1.80 3.90 1.55 1.30 1.50 NY85 3.95 1.95 1.30 1.60 3.75 1.75 1.30 1.55 LPB8 4.85 1.25 1.85 1.10 4.75 1.40 2.30 1.15 MU10 5.90 1.55 2.00 1.20 5.35 1.55 2.20 1.20 NY104 4.23 1.17 1.50 0.87 4.03 1.23 1.93 1.03 LPB11 4.95 1.15 1.80 1.05 4.70 1.35 2.05 1.05 LPB19 4.35 1.15 1.45 1.10 3.95 1.30 1.65 1.30 NY25 4.90 1.40 1.60 1.00 4.40 1.45 1.70 1.20 B4 4.20 1.05 1.40 0.80 4.05 1.30 1.70 1.00 LPB18 4.65 1.45 1.80 1.10 4.30 1.55 1.75 1.10

Appendix 3 continued.

OTUs WPetL* WPetLamWw* WPetClwL* WPetLamWn* KPetL* KPetLamWw* KPetClwL* KPetLamWn* MU17 4.60 1.40 1.50 1.00 4.15 1.25 1.70 1.10 NY90 4.45 1.20 1.50 0.95 4.45 1.55 1.60 1.20 US165 6.00 1.75 1.40 1.10 5.00 1.75 1.65 1.15 NY79 4.85 1.55 1.50 0.90 4.40 1.60 1.90 1.00 LPB4 4.30 1.10 1.50 0.90 4.10 1.30 1.80 1.00 NY91 3.68 0.83 1.28 0.73 3.23 0.98 1.25 0.78 F52 3.45 1.00 1.10 0.75 3.25 1.10 1.30 0.90 NY21 4.16 1.08 1.30 0.88 3.84 1.24 1.62 0.82 MU23 4.65 1.40 1.40 1.10 4.00 1.50 1.60 1.10 AHUC2 3.68 0.83 1.33 0.68 3.33 0.95 1.33 0.83 LPB14 3.70 0.85 1.30 0.70 3.40 1.05 1.50 0.85 F57 4.35 1.15 1.85 0.80 3.95 1.25 1.60 0.85 LPB15 4.10 1.20 1.40 1.10 4.00 1.50 1.50 1.10 LPB17 4.15 1.20 1.45 0.95 3.95 1.20 1.80 1.00 MU6 5.35 1.60 1.65 0.90 4.70 1.45 1.75 0.85 MO43 5.75 1.65 1.75 0.95 4.90 1.50 1.85 0.90 LPB6 6.20 2.10 1.50 0.80 5.05 1.60 1.80 0.90 NY98 4.47 1.20 1.37 0.97 4.30 1.53 1.43 1.03 US4 6.40 2.05 1.85 1.10 5.60 1.70 2.00 1.00 P62 5.50 1.40 1.80 1.00 5.25 1.35 2.10 1.10 P86 7.25 1.85 2.20 1.20 6.25 1.75 2.30 1.20 P6 6.70 1.60 2.30 1.10 6.20 1.45 2.45 1.00 NY69 6.26 1.48 1.84 1.04 5.44 1.46 1.98 1.04 US8 6.45 1.65 1.75 1.05 5.40 1.60 1.85 1.15 US100 5.85 1.40 1.75 0.90 4.70 1.25 2.00 1.00 US12 8.90 2.35 2.40 1.60 7.30 2.10 2.60 1.65 F45 7.40 2.25 1.90 1.25 5.95 1.85 2.05 1.40 AAU6 6.60 1.80 1.85 1.20 5.85 1.70 2.05 1.20 US2 6.75 1.75 2.00 1.15 5.75 1.55 2.30 1.20 F44 6.35 1.85 1.65 1.05 5.50 1.60 2.10 1.05 F47 6.55 1.85 1.80 1.15 5.35 1.70 1.80 1.25 US35 6.67 1.63 2.10 1.20 5.70 1.50 2.03 1.13 MO10 6.20 1.70 1.85 1.00 5.25 1.60 1.90 1.15 NY101 6.15 1.80 1.75 1.05 5.35 1.55 1.85 1.00 F135 7.05 2.15 1.90 1.00 5.50 1.80 2.00 1.05

Appendix 3 continued.

OTUs WPetL* WPetLamWw* WPetClwL* WPetLamWn* KPetL* KPetLamWw* KPetClwL* KPetLamWn* P83 6.80 2.05 1.70 1.15 5.40 1.80 1.85 1.15 F91 6.70 2.05 1.85 1.10 5.70 1.75 2.05 1.10 F104 7.00 1.70 2.15 1.05 5.70 1.70 2.15 1.20 MO37 5.75 1.85 1.75 1.05 5.10 1.65 1.90 1.05 F114 6.00 1.85 1.75 0.95 5.00 1.65 1.80 0.90 F126 6.33 1.73 1.33 1.13 5.57 1.50 1.97 1.23 F109 6.70 2.30 1.60 1.10 5.45 1.85 1.80 1.15 US131 7.00 1.90 2.20 1.30 6.00 1.80 2.30 1.10 F122 5.80 1.70 1.70 1.10 5.25 1.75 2.20 1.05 F111 8.65 2.70 2.10 1.40 6.75 2.20 2.40 1.25 F110 7.97 2.80 2.07 1.23 6.73 2.07 2.30 1.27 B11 6.95 2.10 1.95 1.30 6.15 2.00 2.20 1.10 NY122 7.60 2.00 2.15 1.40 6.60 2.05 2.45 1.25 US13 6.30 1.45 1.95 1.20 5.35 1.55 2.05 1.10 US26 5.40 1.50 1.60 1.10 5.10 1.30 1.80 1.10 F124 6.55 1.85 1.75 1.30 5.65 1.65 1.90 1.15 US20 6.64 2.10 1.78 1.10 5.60 1.72 1.98 1.18 US21 6.00 1.90 1.75 1.15 5.50 1.55 1.85 1.15 US129 5.40 1.25 1.70 0.90 4.80 1.60 1.85 1.00 F133 6.58 1.90 1.88 1.05 5.68 1.82 1.96 1.06 MO32 6.90 2.05 2.20 1.25 6.20 1.80 2.35 1.25 MO19 6.75 1.85 1.85 0.95 5.80 1.65 2.10 1.20 US34 7.30 2.50 2.14 1.12 6.30 1.96 2.38 1.12 F98 6.82 1.94 2.26 1.04 6.06 1.78 2.46 1.24 US132 5.30 1.45 1.65 1.00 4.60 1.35 1.55 1.00 LPB3 6.40 2.25 1.60 1.20 5.55 1.80 2.10 1.15 NY97 7.35 1.75 1.95 1.10 5.65 1.85 2.05 1.20 P8 5.65 1.65 1.45 1.00 4.95 1.45 1.95 1.10 MU19 5.80 1.70 1.66 1.08 5.40 1.52 1.76 1.00 NY63 6.15 2.05 1.80 1.30 5.55 1.95 2.05 1.15 SI6 5.45 1.45 1.95 1.05 5.00 1.50 2.00 1.20 LPB22 6.30 1.95 2.05 1.15 5.65 1.75 2.10 1.15 US19 6.25 1.75 2.00 0.95 5.45 1.65 2.20 1.00 MU21 5.70 1.60 1.80 1.10 5.15 1.75 2.10 1.10 NY24 5.60 1.75 1.85 1.00 4.75 1.60 1.75 0.95

Appendix 3 continued.

OTUs WPetL* WPetLamWw* WPetClwL* WPetLamWn* KPetL* KPetLamWw* KPetClwL* KPetLamWn* MO3 6.45 1.95 1.70 1.05 5.30 1.70 1.85 1.05 F2 5.90 1.50 1.70 0.90 5.00 1.35 2.00 1.00 F90 6.00 1.55 2.00 1.20 6.00 1.90 2.20 1.30 MU34 4.45 1.50 1.15 1.00 4.00 1.30 1.30 0.90

Appendix 4. Table of qualitative and quantitative data not used for analysis. Quantitative data

in mm.

OTUs Habit Root InNodPub† NodPub† StipAdPub† StipAbPub† StipMarPub† PetPub† LfltAdPub† LfltAbMdrbPub† LfltAbPub† US14 P NA S SM G G S S G S S NY51 P NA S S G S S SM G S 1/2S F121 P NA SM S G GS S SM G S 1/2S NY66 P LW S SM G S S M G S 1/2S MU27 P LW S GS G G S SD G S 1/2S US171 P NA NA NA NA NA NA NA NA NA NA US169 P NA GS GS G G SM SM G S G1/2S F118 P NA S SM G S S SM G S G1/2S MO44 P NA NA NA NA NA NA NA NA NA NA P97 P NA GM SM G G SM MD G S G1/2S NY60 P NA SM SM G GS S SM G S 1/2S MU26 P LW S SM G S S S G S 1/2S US32 P W S S G G S SM G S 1/2S MO48 P LW MD GD G G S SM G S 1/2S NY119 P LW SM SM G S S S G S G1/2S MO49 P NA NA NA NA NA NA NA NA NA NA MUX P NA NA NA NA NA NA NA NA NA NA NYX P NA NA NA NA NA NA NA NA NA NA US29 P NA NA NA NA NA NA NA NA NA NA NY50 P NA NA NA NA NA NA NA NA NA NA NY89 P W S GM G G S SM G GS G1/2S US30 P LW S SM G G S SM G S G1/2S US28 P NA S SM G S S S G S 1/2S F48 P W S S G G S SM G S 1/2S MU9 P W S SM G GS S M G M 1/2S F131 P NA NA NA NA NA NA NA NA NA NA NY85 P NA NA NA NA NA NA NA NA NA NA LPB8 P NA S SM G G S S G S 1/2S MU10 P NA NA NA NA NA NA NA NA NA NA NY104 P NA S G G G S S G S G1/2S LPB11 P W SM SM G GS S SM G S 1/2S LPB19 P NA NA NA NA NA NA NA NA NA NA NY25 P NA NA NA NA NA NA NA NA NA NA B4 P W S GM G G GS S G SM 1/2S LPB18 P NA NA NA NA NA NA NA NA NA NA

Appendix 4 continued.

OTUs Habit Root InNodPub† NodPub† StipAdPub† StipAbPub† StipMarPub† PetPub† LfltAdPub† LfltAbMdrbPub† LfltAbPub† MU17 P NA NA NA NA NA NA NA NA NA NA NY90 P LW MD D G S S MD G SM 1/2S US165 P W M MD G S S D G S 1/2S NY79 P NA NA NA NA NA NA NA NA NA NA LPB4 P NA S MD G S G S G S 1/2S NY91 P LW S M G G G MD G S 1/2S F52 P LW SM SM G G S M G SM G1/2S NY21 P LW S S G G S S G S 1/2S MU23 P NA NA NA NA NA NA NA NA NA NA AHUC2 P W S S G G S S G S 1/2S LPB14 P LW S SM G G S S G S 1/2S F57 P W S SM G G S S G S 1/2S LPB15 P W S M G S S MD G S 1/2S LPB17 P LW S S G G S S G S G1/2S MU6 P LW S SM G S S S G S 1/2S MO43 P W S SM G S S SM G S 1/2S LPB6 P NA NA NA NA NA NA NA NA NA NA NY98 P LW SM M G S S SM G S 1/2S US4 P W S M G S S M G S 1/2S P62 P W M MD G S S M G M 1/2M P86 P W SM SM G S S M G S 1/2S P6 P NA S M G S S SM G S 1/2S NY69 P LW S GM G S G M G M G1/2S US8 P NA SM M G S S S G S 1/2S US100 P W S SM G G S SM G S G1/2S US12 P W S SM G S GS S G S 1/2S F45 P NA S SM G G S S G S 1/2S AAU6 P NA S SM G S S S G S 1/2S US2 P NA S GS G G G SM G SM 1/2S F44 P NA SM SM G S S SM G S 1/2S F47 P W S GS G G S SM G M 1/2M US35 P NA S SM G S S S G S 1/2S MO10 P W S SM G S S S G M 1/2M NY101 P NA SM M G S S SM G M 1/2M F135 P W S GS G G GS S G S 1/2S

Appendix 4 continued.

OTUs Habit Root InNodPub† NodPub† StipAdPub† StipAbPub† StipMarPub† PetPub† LfltAdPub† LfltAbMdrbPub† LfltAbPub† P83 P NA S GM G G S S G S G1/2S F91 P NA S SM G S S SM G S G1/2S F104 P W S SM G G S S G S G MO37 P NA S SM G S S S G S 1/2S F114 P NA M SM G G SM MD NA SM G1/2S F126 P W S SM G G G GS G S 1/2S F109 P W S SM G G S S G S 1/2S US131 P NA NA NA NA NA NA NA NA NA NA F122 P NA NA NA NA NA NA NA NA NA NA F111 P LW SM SM G GS S S G SM G1/2S F110 P LW M MD G S S SM G SM 1/2SM B11 P LW MD D G G S MD G SM G1/2S NY122 P NA M SD G S S M G S 1/2M US13 P W S SM G G S SM G SM G US26 P W NA NA G G S S G S 1/2S F124 P NA SM SD G G S SM G SM 1/2S US20 P W S M G G G S G S 1/2S US21 P W M MD G S S SM G M 1/2M US129 P NA S S G S S SM G M 1/2S F133 P W S S G G S GS S M S MO32 P NA SM SD G S S SM G S 1/2S MO19 P LW S SM G G S S G S 1/2S US34 P NA SM MD G G S M G SM 1/2S F98 P LW SM M G G S M G M 1/2M US132 P W S SM G S SM S G S 1/2S LPB3 P W S SM G G S SM G S 1/2S NY97 P W S MD G S S S G S 1/2S P8 P W SM M G G S SM G S G1/2S MU19 P NA S M G S G S G S 1/2S NY63 P NA S SD S S S S G S G1/2S SI6 P NA S S G G S S G S 1/2S LPB22 P W SM SM G GS S S G S G1/2S US19 P W SM SM G GS S S G S 1/2S MU21 P NA NA NA NA NA NA NA NA NA NA NY24 P W S SM G S S S G S 1/2S

Appendix 4 continued.

OTUs Habit Root InNodPub† NodPub† StipAdPub† StipAbPub† StipMarPub† PetPub† LfltAdPub† LfltAbMdrbPub† LfltAbPub† MO3 P NA NA NA NA NA NA NA NA NA NA F2 P W S SM G S S S G S G1/2S F90 P NA NA NA NA NA NA NA NA NA NA MU34 P W S GS G G S S G GS G

Appendix 4 continued.

OTUs LfltMarPub† PetllePub† PeduncPub† PedicPub† SepTubPub† SepLobPub† SepLobMarPub† StipShp‡ StipMar§ LfltShp‡ LfltMar§ US14 G S SM S S S S SeA E OcO Sr NY51 S SM SM M M S M SeA E OcO Sr F121 S SM M M M S S SeA E OcO Sr NY66 S SM M SM MD MD MD SeA E OcO Sr MU27 S S S SM MD SM M SeA E OcO Sr US171 NA NA NA NA NA NA NA NA NA NA NA US169 S S GS S S S SM SeA Se OcO Sr F118 S S GS S GS GS S T E EOcO Sr MO44 NA NA NA NA NA NA NA NA NA NA NA P97 S SM M MD MD MD MD SeA E EOcO Sr NY60 S M SM S M S M SeA E Oc Sr MU26 S SM S SM M M M SeA E OcO Sr US32 GS S SM S S S S SeA E OcO Sr MO48 S S D MD MD MD MD SeA E OcO SeSr NY119 S S M S M S M SeA E Oc SeSr MO49 NA NA NA NA NA NA NA NA NA NA NA MUX NA NA NA NA NA NA NA NA NA NA NA NYX NA NA NA NA NA NA NA NA NA NA NA US29 NA NA NA NA NA NA NA NA NA NA NA NY50 NA NA NA NA NA NA NA NA NA NA NA NY89 GS S SM S D D D SeA E OcO SeSr US30 G S D M S S S SeA E EOcO SeSr US28 GS S SM S S S S SeA E EOcO Sr F48 S SM SM M M M M SeA E OcO Sr MU9 GS M MD M D D D SeA E EOcO Sr F131 NA NA NA NA NA NA NA NA NA NA NA NY85 NA NA NA NA NA NA NA NA NA NA NA LPB8 S GS S S SM SM SM SeA E OcO Sr MU10 NA NA NA NA NA NA NA NA NA NA NA NY104 GS S S M MD M S Sr NA Sr LPB11 S S S S M S M SeA E EO Sr LPB19 NA NA NA NA NA NA NA NA NA NA NA NY25 NA NA NA NA NA NA NA NA NA NA NA B4 S S S S MD SM SM SeA E O Sr LPB18 NA NA NA NA NA NA NA NA NA NA NA

Appendix 4 continued.

OTUs LfltMarPub† PetllePub† PeduncPub† PedicPub† SepTubPub† SepLobPub† SepLobMarPub† StipShp‡ StipMar§ LfltShp‡ LfltMar§ MU17 NA NA NA NA NA NA NA NA NA NA NA NY90 G M M S M M M SeA E OcO Sr US165 S S MD D D D D SeA E EO Sr NY79 NA NA NA NA NA NA NA NA NA NA NA LPB4 G S S S M M M T SrE OcO Sr NY91 G M M SM SM S GS NA E OcO Sr F52 G SM MD D D M D SeA E EOcO Sr NY21 G D S SM D MD MD NA E OcO Sr MU23 NA NA NA NA NA NA NA NA NA NA NA AHUC2 S M S S S S M T E O Sr LPB14 G S SM S GS GS SM SeA E EOcO Sr F57 S SM SM S M S M SeA E EO Sr LPB15 S SM SM S S S S T E EOcO Sr LPB17 GS S SM S S S MD NT E O Sr MU6 GS S S S S S S SeA E Oc SeSr MO43 S S SM S S G S SeA E OcO Sr LPB6 NA NA NA NA NA NA NA NA NA NA NA NY98 G S M M SM SM SM T E OcO Sr US4 G SM SM S M M M SeA Se OcO Sr P62 M M SM M M M M T E OcO Sr P86 S M SM MD MD MD MD SeA E OcO SeSr P6 M SM M M SM SM SM SeA E Oc Sr NY69 S M M MD S S S NA E NA SeSr US8 S SM M MD D D D SeA E Oc Sr US100 S SM MD MD MD SM MD SeA E Oc Sr US129 S SM SM SM M S M SeA Se OC Sr F45 S S SM S S S S SeA E Oc Sr AAU6 S S S S M S M SeA E Oc Sr US2 S S M MD SM S S T E Oc Sr F44 S S SM M M S M SeA E Oc SeSr F47 S S S SM S S SM SeA E OcO Sr US35 G S S M M M M T SrSeE Oc SeSr MO10 S S MD MD S M M T E Oc Sr NY101 S S MD M MD SM MD SeA E OcO SeSr F135 S SM SM S GS GS GS SeA E EOC Sr

Appendix 4 continued.

OTUs LfltMarPub† PetllePub† PeduncPub† PedicPub† SepTubPub† SepLobPub† SepLobMarPub† StipShp‡ StipMar§ LfltShp‡ LfltMar§ P83 S GS S SM S S S SeA E EO SeSr F91 G SM SM S S S S SeA E EO SeSr F104 GS SM SM SM SM S S SeA E El Sr MO37 S SM MD S MD MD MD SeA E O Sr F114 S M SM M S S S SeA E OcO Sr F126 GS GS S M M M M T E Oc Sr F109 S SM SD M S S S SeA E OcO SeSr US131 NA NA NA NA NA NA NA NA NA NA NA F122 NA NA NA NA NA NA NA NA NA NA NA F111 S M SM SM D D D SeA E OcO Sr F110 S D D MD MD MD MD T E OcO Sr B11 G SM D M M M M SeA E EO Sr NY122 S MD MD M SM S S SeA E El Sr US13 G S D MD MD MD MD SeA E Oc SeSr US26 S S SD MD D D D T E OcO Se F124 S SM MD M M M M SeA E Oc Sr US20 G S SM M MD M M T E Oc Sr US21 M M M M M M M SeA E OcO SeSr US129 S SM MD M D M D SeA E EOcO Sr F133 G M SM SM SM SM SM NA E OcO SeSr MO32 GS SM S S S S SM SeA E El Sr MO19 S MD M S S S M SeA E EO Sr US34 S D MD D D M M NA E OcO Sr F98 G M M M S M M NA E Sr US132 S M M MD MD S M SeA Se OcO SeSr LPB3 S SM SM GS S GS S SeA E El Sr NY97 S SM SM M S S S SeA E OcO SeSr P8 S D M S S G S T E OcO SeSr MU19 G M MD SM S M M NA E NA Sr NY63 GS SM MD S M GS S SeA E O Sr SI6 S M SM M MD MD MD SeA E EO Sr LPB22 S S SM S GS GS GS SeA E OcO Sr US19 S S M S SM S M SeA E Oc Sr MU21 NA NA NA NA NA NA NA NA NA NA NA NY24 G SM SM S SM S S SeA E OcO Sr

Appendix 4 continued.

OTUs LfltMarPub† PetllePub† PeduncPub† PedicPub† SepTubPub† SepLobPub† SepLobMarPub† StipShp‡ StipMar§ LfltShp‡ LfltMar§ MO3 NA NA NA NA NA NA NA NA NA NA NA F2 G S S S S S M SeA E EOcO SeSr F90 NA NA NA NA NA NA NA NA NA NA NA MU34 GS GS S S S S SM SeA E OcO SeSr

Appendix 4 continued.

OTUs LfltMucr Bract SepLobShp‡ SepLobMar§ RootDiam* StemD* StemL* InNodL* StipApxAng LfltApxAng LfltBasAng US14 Y Bf Nt E NA NA 408.0 6.0 10.0 176.0 80.0 NY51 Y Nt Nt E NA NA 141.0 11.0 10.0 172.0 72.0 F121 N Bf Nt Se NA NA 205.0 7.0 10.0 180.0 78.0 NY66 Y Bf Nt Se 4.8 1.5 200.0 10.7 10.0 180.0 78.0 MU27 N Bf Nt Se 4.1 1.7 206.0 4.3 10.0 160.0 82.0 US171 NA NA NA NA NA NA NA NA NA NA NA US169 Y Bf Nt Se NA NA 120.0 5.0 10.0 156.0 78.0 F118 N Bf Nt Se NA NA 62.0 5.5 10.0 144.0 76.0 MO44 NA NA NA NA NA NA NA NA NA NA NA P97 Y Bf Nt Se NA NA 164.0 6.4 10.0 152.0 72.0 NY60 Y Nt Nt Se NA NA 183.0 5.0 10.0 180.0 72.0 MU26 Y Nt A E 4.9 1.5 261.0 7.5 10.0 172.0 80.0 US32 Y Nt Nt E 5.4 1.7 359.0 NA 10.0 140.0 88.0 MO48 Y Bf Nt E 5.0 1.9 102.0 5.0 10.0 168.0 86.0 NY119 Y Nt Nt E 4.3 1.6 148.0 7.4 10.0 180.0 66.0 MO49 NA NA NA NA NA NA NA NA NA NA NA MUX NA NA NA NA NA NA NA NA NA NA NA NYX NA NA NA NA NA NA NA NA NA NA NA US29 NA NA NA NA NA NA NA NA NA NA NA NY50 NA NA NA NA NA NA NA NA NA NA NA NY89 Y Bf Nt E 2.2 1.4 108.0 6.0 10.0 164.0 82.0 US30 Y Bf Nt Se 5.2 1.3 400.0 4.0 10.0 144.0 64.0 US28 Y Bf Nt E NA 1.6 474.0 12.3 10.0 152.0 70.0 F48 Y Nt Nt E 2.9 0.9 289.0 5.0 10.0 172.0 76.0 MU9 Y Nt Nt E 6.4 1.5 275.0 8.2 10.0 140.0 92.0 F131 NA NA NA NA NA NA NA NA NA NA NA NY85 NA NA NA NA NA NA NA NA NA NA NA LPB8 N Bf Nt E NA NA 49.4 3.7 10.0 134.0 68.0 MU10 NA NA NA NA NA NA NA NA NA NA NA NY104 N Nt NA Se NA 1.2 85.0 5.4 15.0 130.0 71.0 LPB11 Y Bf Nt E 3.7 1.0 152.0 8.0 10.0 112.0 64.0 LPB19 NA NA NA NA NA NA NA NA NA NA NA NY25 NA NA NA NA NA NA NA NA NA NA NA B4 Y Bf Nt E 2.4 1.4 134.3 6.7 10.0 156.0 74.0 LPB18 NA NA NA NA NA NA NA NA NA NA NA

Appendix 4 continued.

OTUs LfltMucr Bract SepLobShp‡ SepLobMar§ RootDiam* StemD* StemL* InNodL* StipApxAng LfltApxAng LfltBasAng MU17 NA NA NA NA NA NA NA NA NA NA NA NY90 Y Nt Nt E 5.1 1.1 74.0 8.0 10.0 168.0 72.0 US165 Y Nt Nt E 3.9 1.3 215.0 3.3 10.0 120.0 78.0 NY79 NA NA NA NA NA NA NA NA NA NA NA LPB4 Y Nt Nt E NA NA 289.0 6.5 10.0 146.0 72.0 NY91 Y NA Nt E 4.0 1.1 281.0 12.5 10.0 106.0 60.0 F52 Y Bf Nt E 8.2 1.5 123.0 8.5 10.0 112.0 58.0 NY21 N NA Nt E 7.0 1.3 426.0 10.2 10.0 126.0 81.0 MU23 NA NA NA NA NA NA NA NA NA NA NA AHUC2 Y Nt Nt E 2.0 1.6 367.0 8.9 10.0 136.0 72.0 LPB14 Y NA Nt E 5.5 1.6 319.0 9.7 10.0 148.0 66.0 F57 Y Nt Nt E 4.9 1.5 489.0 7.5 10.0 132.0 76.0 LPB15 Y Bf Nt E 4.2 1.2 392.0 4.0 10.0 132.0 60.0 LPB17 Y Bf Nt E 6.6 1.7 228.0 13.0 10.0 112.0 72.0 MU6 Y Nt Nt Se 2.3 1.1 468.0 9.0 10.0 180.0 80.0 MO43 Y Nt Nt E 6.2 1.7 521.0 13.0 10.0 168.0 66.0 LPB6 NA NA NA NA NA NA NA NA NA NA NA NY98 Y Nt Nt E 10.4 1.4 234.0 9.5 10.0 148.0 68.0 US4 Y Nt Nt E 3.9 1.1 275.0 9.0 10.0 152.0 98.0 P62 Y Bf Nt E 3.9 1.1 289.0 5.0 10.0 164.0 68.0 P86 Y Bf Nt E 4.7 1.8 441.0 10.0 10.0 180.0 74.0 P6 Y Nt Nt E NA NA 433.0 9.0 10.0 180.0 74.0 NY69 N Nt A E 37.5 1.5 480.0 10.0 10.0 180.0 56.0 US8 Y Bf Nt E NA 1.4 162.0 9.0 10.0 180.0 58.0 US100 Y Bf Nt E 0.8 1.0 330.0 5.5 10.0 180.0 64.0 US129 Y Bf Nt Se 3.5 1.3 300.0 11.0 10.0 180.0 72.0 F45 N Bf Nt E NA NA 216.0 8.5 10.0 180.0 86.0 AAU6 Y Nt Nt E NA NA 304.0 11.0 10.0 180.0 64.0 US2 Y Bf Nt E NA NA 269.0 10.5 10.0 180.0 74.0 F44 Y Nt Nt E NA 1.3 74.0 6.0 10.0 180.0 76.0 F47 Y Bf Nt E 5.1 1.7 189.0 5.3 10.0 176.0 66.0 US35 Y Bf Nt E NA NA 324.0 8.5 10.0 180.0 68.0 MO10 Y Bf Nt E 3.7 1.0 223.0 7.0 10.0 180.0 74.0 NY101 Y Bf Nt E NA 1.0 113.0 10.3 10.0 168.0 70.0 F135 Y Bf Nt E 3.7 1.4 290.0 4.3 10.0 172.0 76.0

Appendix 4 continued.

OTUs LfltMucr Bract SepLobShp‡ SepLobMar§ RootDiam* StemD* StemL* InNodL* StipApxAng LfltApxAng LfltBasAng P83 Y Bf Nt E NA NA 324.7 6.3 10.0 118.0 66.0 F91 Y Bf Nt E NA NA 269.0 9.0 10.0 140.0 72.0 F104 Y Bf Nt E 4.5 1.5 559.0 10.6 10.0 100.0 56.0 MO37 Y Bf Nt E NA NA 102.8 5.5 10.0 152.0 66.0 F114 Y Nt Nt E NA NA 97.0 12.0 10.0 140.0 58.0 F126 N Bf Nt E 4.0 1.3 550.0 13.5 10.0 180.0 78.0 F109 Y Bf Nt E 4.8 1.1 621.0 8.8 10.0 172.0 74.0 US131 NA NA NA NA NA NA NA NA NA NA NA F122 NA NA NA NA NA NA NA NA NA NA NA F111 Y Bf Nt E 7.9 1.5 658.0 11.0 10.0 160.0 72.0 F110 Y Bf A E 7.8 1.8 301.0 15.9 10.0 164.0 80.0 B11 Y Bf Nt Se 8.0 1.3 118.0 8.0 10.0 118.0 78.0 NY122 Y Bf Nt E NA NA 217.5 9.5 10.0 136.0 96.0 US13 Y Bf Nt E 4.0 1.6 81.3 8.0 10.0 180.0 68.0 US26 Y Bf Nt E NA NA 59.0 6.5 10.0 176.0 86.0 F124 Y Bf Nt E NA NA 460.0 5.0 10.0 180.0 64.0 US20 Y Nt Nt E 1.5 1.0 343.0 9.0 10.0 180.0 72.0 US21 Y Bf Nt E 4.2 2.0 193.0 9.5 10.0 162.0 68.0 US129 Y Nt Nt E NA 1.7 358.0 9.0 10.0 156.0 72.0 F133 Y Nt Nt E 4.0 1.5 446.0 35.0 10.0 78.0 76.0 MO32 Y Nt Nt E NA NA 392.5 12.0 10.0 116.0 72.0 MO19 Y Nt Nt E 7.3 1.6 288.0 10.3 10.0 126.0 60.0 US34 Y Nt Nt E NA NA 549.0 26.5 10.0 180.0 67.0 F98 N Nt Nt E 6.0 0.8 506.0 21.0 10.0 70.0 60.0 US132 Y NA Nt E 5.6 1.8 406.0 8.0 10.0 176.0 72.0 LPB3 Y Bf Nt Se 5.1 1.2 218.0 8.5 10.0 100.0 88.0 NY97 Y Bf Nt E NA 1.5 471.0 18.8 10.0 180.0 72.0 P8 Y Nt Nt E 3.8 1.6 301.0 8.5 10.0 148.0 70.0 MU19 N Nt Nt E NA NA 285.0 25.8 10.0 150.0 56.0 NY63 N Bf Nt Se NA NA 408.0 27.0 10.0 180.0 78.0 SI6 Y Bf Nt E NA NA 179.8 13.0 10.0 112.0 72.0 LPB22 N Bf Nt Se 7.1 1.8 236.0 13.5 10.0 156.0 84.0 US19 Y Nt Nt E 3.2 1.5 420.0 9.5 10.0 180.0 68.0 MU21 NA NA NA NA NA NA NA NA NA NA NA NY24 N Bf Nt Se 2.8 1.3 540.0 11.0 10.0 176.0 92.0

Appendix 4 continued.

OTUs LfltMucr Bract SepLobShp‡ SepLobMar§ RootDiam* StemD* StemL* InNodL* StipApxAng LfltApxAng LfltBasAng MO3 NA NA NA NA NA NA NA NA NA NA NA F2 Y Bf Nt E 1.9 1.4 373.0 9.0 10.0 176.0 56.0 F90 NA NA NA NA NA NA NA NA NA NA NA MU34 Y Bf Nt E 2.3 1.2 105.5 7.0 10.0 132.0 78.0

Appendix 4 continued.

OTUs PetD* PetlleL* PetlleD* PeduncD* PedicD* BPetShp‡ WPetLamBas§ WPetLamL* KPetClwW* WPetLamMar§ WPetLamApx§ US14 0.64 1.0 0.42 1.00 0.3 Ro Sa 4.6 0.6 E Ro NY51 0.40 0.6 0.36 0.68 0.2 Oc Sa 4.2 0.6 E Ro F121 0.46 0.9 0.40 0.44 0.2 Ro Tr 4.1 0.6 E Ro NY66 0.60 1.0 0.40 0.92 0.3 Ro Sa 4.2 0.6 E Ro MU27 0.48 1.0 0.36 0.60 0.3 Oc Sa 3.4 0.5 E Ro US171 NA NA NA NA NA NA NA NA NA NA NA US169 0.56 0.7 0.36 0.30 0.2 Oc Sa 3.1 0.4 E Ro F118 0.46 0.5 0.26 0.60 0.3 Oc Sa 3.3 0.4 E Ro MO44 NA NA NA NA NA NA NA NA NA NA NA P97 0.58 1.0 0.32 0.47 0.2 Oc Tr 4.1 0.5 E Ro NY60 0.68 0.8 0.40 1.00 0.2 Ro Sa 3.9 0.5 E Ro MU26 0.60 0.8 0.40 0.60 0.3 Oc Sa 3.6 0.5 E Ro US32 0.68 1.0 0.44 0.85 0.2 Ro Sa 3.8 0.5 E Ro MO48 0.40 0.5 0.30 0.56 0.2 Ro Sa 3.5 0.5 E Ro NY119 0.36 0.8 0.32 0.52 0.3 Oc Tr 4.0 0.5 E Ro MO49 NA NA NA NA NA NA NA NA NA NA NA MUX NA NA NA NA NA NA NA NA NA NA NA NYX NA NA NA NA NA NA NA NA NA NA NA US29 NA NA NA NA NA NA NA NA NA NA NA NY50 NA NA NA NA NA NA NA NA NA NA NA NY89 0.44 0.8 0.36 0.40 0.2 Oc Sa 3.3 0.4 E Ro US30 0.58 0.7 0.40 0.53 0.2 Oc Tr 3.9 0.6 E Ro US28 0.48 1.0 0.40 0.68 0.2 Oc Sa 5.1 0.5 E Ro F48 0.46 1.0 0.38 0.54 0.3 Ro NA NA NA NA NA MU9 0.56 0.7 0.38 0.64 0.3 Ro Tr 3.1 0.4 E Ro F131 NA NA NA NA NA NA NA NA NA NA NA NY85 NA NA NA NA NA NA NA NA NA NA NA LPB8 0.34 0.7 0.20 0.40 0.2 Ro Sa 3.1 0.5 E Ro MU10 NA NA NA NA NA NA NA NA NA NA NA NY104 0.34 0.7 0.24 0.34 0.2 Oc NA 2.7 0.4 E NA LPB11 0.40 0.7 0.30 0.38 0.2 Oc Sa 3.2 0.5 E Ro LPB19 NA NA NA NA NA NA NA NA NA NA NA NY25 NA NA NA NA NA NA NA NA NA NA NA B4 0.34 0.7 0.28 0.38 0.2 Oc Sa 2.7 0.4 E Ro LPB18 NA NA NA NA NA NA NA NA NA NA NA

Appendix 4 continued.

OTUs PetD* PetlleL* PetlleD* PeduncD* PedicD* BPetShp‡ WPetLamBas§ WPetLamL* KPetClwW* WPetLamMar§ WPetLamApx§ MU17 NA NA NA NA NA NA NA NA NA NA NA NY90 0.52 0.6 0.40 0.47 0.2 Oc Sa 2.9 0.5 E Ro US165 0.48 0.8 0.40 0.60 0.2 Oc Sa 4.6 0.5 E Ro NY79 NA NA NA NA NA NA NA NA NA NA NA LPB4 0.32 0.9 0.40 0.60 0.2 Oc Sa 2.8 0.4 E Ro NY91 0.44 0.7 0.34 0.68 0.2 Oc Sa 2.4 0.3 E Ro F52 0.40 0.8 0.36 0.60 0.2 Oc Tr 2.4 0.3 E Ro NY21 0.68 0.6 0.44 0.72 0.2 Ro Sa 2.8 0.4 E Ro MU23 NA NA NA NA NA NA NA NA NA NA NA AHUC2 0.48 0.9 0.32 0.65 0.2 Oc Sa 2.3 0.5 E Ro LPB14 0.40 1.0 0.32 0.64 0.2 Oc Sa 2.4 0.3 E Ro F57 0.60 0.5 0.40 0.52 0.2 Oc Tr 2.8 0.4 E Ro LPB15 0.66 0.9 0.40 0.60 0.2 Oc Sa 2.6 0.5 E Ro LPB17 0.36 0.8 0.38 0.50 0.3 Oc Sa 2.7 0.5 E Ro MU6 0.40 1.1 0.38 0.60 0.2 Oc Tr 3.8 0.4 E Ro MO43 0.40 0.9 0.38 0.52 0.2 Oc Sa 4.0 0.4 E Ro LPB6 NA NA NA NA NA NA NA NA NA NA NA NY98 0.32 0.9 0.40 0.52 0.2 Oc Sa 3.0 0.5 E Ro US4 0.60 1.0 0.34 0.60 0.2 Ro Tr 4.5 0.6 E Ro P62 0.48 1.0 0.36 0.56 0.2 Ro Sa 3.7 0.5 E Ro P86 0.48 0.8 0.44 0.68 0.2 Oc Sa 5.1 0.6 E Ro P6 0.48 1.0 0.40 0.56 0.2 Oc Sa 4.4 0.4 E Ro NY69 0.56 1.2 0.56 0.54 0.2 Ro NA 4.4 0.5 E NA US8 0.40 1.1 0.64 0.44 0.2 Oc Sa 4.6 0.5 E Ro US100 0.42 0.9 0.32 0.40 0.2 Oc Sa 4.1 0.5 E Ro US129 0.56 0.9 0.40 0.56 0.2 Oc Sa 6.5 0.7 E Ro F45 0.56 1.0 0.42 0.56 0.3 Oc Sa 5.5 0.6 E Ro AAU6 0.58 1.0 0.40 0.40 0.2 Oc Sa 4.8 0.6 E Ro US2 0.56 1.0 0.40 0.56 0.6 Oc Sa 4.7 0.5 E Ro F44 0.32 0.6 0.36 0.44 0.3 Oc Tr 4.8 0.4 E Ro F47 0.38 0.5 0.34 0.44 0.2 Oc Sa 4.8 0.5 E Ro US35 0.36 0.8 0.26 0.40 0.2 Oc Sa 4.6 0.5 E Ro MO10 0.40 0.7 0.40 0.40 0.2 Oc Sa 4.3 0.5 E Ro NY101 0.48 0.7 0.38 0.67 0.2 Oc Sa 4.5 0.5 E Ro F135 0.44 0.9 0.32 0.64 0.2 Oc Tr 5.0 0.5 E Ro

Appendix 4 continued.

OTUs PetD* PetlleL* PetlleD* PeduncD* PedicD* BPetShp‡ WPetLamBas§ WPetLamL* KPetClwW* WPetLamMar§ WPetLamApx§ P83 0.48 0.8 0.36 0.72 0.2 Oc Sa 5.0 0.5 E Ro F91 0.60 0.9 0.40 0.60 0.2 Oc Sa 4.8 0.5 E Ro F104 0.34 0.9 0.36 0.68 0.2 Oc Sa 4.9 0.6 E Ro MO37 0.44 0.9 0.28 0.40 0.2 Oc Sa 4.1 0.5 E Ro F114 0.40 0.8 0.40 0.50 0.2 Oc Sa 4.2 0.5 E Ro F126 0.42 0.8 0.44 0.46 0.2 Oc Sa 4.6 0.5 E Ro F109 0.40 1.0 0.52 0.40 0.2 Oc Sa 5.0 0.5 E Ro US131 NA NA NA NA NA NA NA NA NA NA NA F122 NA NA NA NA NA NA NA NA NA NA NA F111 0.64 0.8 0.40 0.76 0.2 Oc Sa 6.5 0.6 E Ro F110 0.68 0.7 0.40 0.60 0.2 Oc Sa 5.8 0.6 E Ro B11 0.40 0.7 0.40 0.96 0.2 Oc Sa 5.0 0.5 E Ro NY122 0.56 1.0 0.40 0.76 0.2 Oc Sa 5.4 0.7 E Ro US13 0.40 0.6 0.30 0.60 0.2 Oc Sa 4.4 0.5 E Ro US26 0.34 0.9 0.26 0.53 0.2 Oc Sa 3.8 0.5 E Ro F124 0.44 0.9 0.34 0.56 0.2 Oc Sa 4.8 0.5 E Ro US20 0.46 0.9 0.42 0.52 0.3 Oc NA 4.9 0.5 E NA US21 0.48 0.9 0.40 0.44 0.2 Oc Sa 4.3 0.5 E Ro US129 0.52 0.7 0.36 0.64 0.2 Oc Sa 3.7 0.5 E Ro F133 0.64 1.1 0.52 0.62 0.2 Oc NA 4.7 0.5 E Ro MO32 0.64 0.8 0.38 0.92 0.2 Oc Sa 4.8 0.6 E Ro MO19 0.68 0.9 0.44 0.68 0.2 Oc Sa 5.0 0.6 E Ro US34 0.46 0.7 0.40 0.76 0.2 Oc Tr 5.1 0.6 E Ro F98 0.64 1.2 0.48 0.96 0.2 NA NA 4.5 0.4 E NA US132 0.44 0.8 0.40 0.52 0.2 Oc Sa 3.6 0.5 E Ro LPB3 0.60 0.6 0.38 0.65 0.2 Oc Tr 4.7 0.5 E Ro NY97 0.52 1.0 0.40 0.68 0.2 Oc Sa 5.3 0.6 E Ro P8 0.66 0.9 0.36 0.65 0.2 Oc Sa 4.2 0.5 E Ro MU19 0.52 7.3 0.42 0.58 0.3 NA NA 4.2 0.4 E NA NY63 0.64 1.1 0.48 0.92 0.2 Oc Tr 4.4 0.5 E Ro SI6 0.44 0.9 0.40 0.46 0.2 Oc Sa 3.5 0.5 E Ro LPB22 0.56 0.9 0.40 0.76 0.2 Oc Sa 4.3 0.5 E Ro US19 0.6 1.0 0.4 0.6 0.3 Oc Sa 4.2 0.5 E Ro MU21 NA NA NA NA NA NA NA NA NA NA NA NY24 0.56 1.1 0.38 0.60 0.2 Oc Sa 3.8 0.5 E Ro

Appendix 4 continued.

OTUs PetD* PetlleL* PetlleD* PeduncD* PedicD* BPetShp‡ WPetLamBas§ WPetLamL* KPetClwW* WPetLamMar§ WPetLamApx§ MO3 NA NA NA NA NA NA NA NA NA NA NA F2 0.44 1.1 0.34 0.72 0.2 Oc Sa 4.2 0.5 E Ro F90 NA NA NA NA NA NA NA NA NA NA NA MU34 0.48 0.8 0.24 0.48 0.2 Oc Sa 3.2 0.5 E Ro

Appendix 4 continued.

OTUs KPetLamBas§ KPetLamMar§ KPetLamApx§ OvPub† StylPub† SeedShp SeedCol SepLobApxAng WPetLamArcApxAng WPetClwW* KPetLamL* US14 Tr E A S G NA NA 15.0 90.0 0.4 3.6 NY51 Tr E Ro G G NA NA 10.0 160.0 0.5 3.1 F121 Tr E AcA G G NA NA 10.0 140.0 0.5 3.2 NY66 Tr E AcA S G NA NA 10.0 105.0 0.5 3.1 MU27 Tr E AcA G G NA NA 10.0 120.0 0.5 2.7 US171 NA NA NA NA NA NA NA NA NA NA NA US169 Tr E A S G NA NA 10.0 120.0 0.4 2.4 F118 Tr E Ro G G Mt BY* 10.0 80.0 0.4 2.7 MO44 NA NA NA NA NA NA NA NA NA NA NA P97 Tr E Ro M G NA NA 10.0 90.0 0.5 3.1 NY60 Tr E RoA G G NA NA 10.0 110.0 0.4 3.2 MU26 Tr E A S G NA NA 10.0 80.0 0.4 3.2 US32 Tr E RoA S G NA NA 10.0 80.0 0.5 2.9 MO48 Tr E A M G NA NA 10.0 120.0 0.4 2.9 NY119 Tr E A M G Mt LB 10.0 90.0 0.5 3.5 MO49 NA NA NA NA NA NA NA NA NA NA NA MUX NA NA NA NA NA NA NA NA NA NA NA NYX NA NA NA NA NA NA NA NA NA NA NA US29 NA NA NA NA NA NA NA NA NA NA NA NY50 NA NA NA NA NA NA NA NA NA NA NA NY89 Tr E A M G NA NA 10.0 90.0 0.4 2.8 US30 Tr E AcA G G NA NA 10.0 90.0 0.4 3.4 US28 Tr E A S G NA NA 10.0 100.0 0.5 3.7 F48 NA NA NA S G NA NA 10.0 NA NA NA MU9 Tr E RoA G G NA NA 10.0 90.0 0.4 3.1 F131 NA NA NA NA NA NA NA NA NA NA NA NY85 NA NA NA NA NA NA NA NA NA NA NA LPB8 Tr E A S G NA NA 10.0 105.0 0.4 2.5 MU10 NA NA NA NA NA NA NA NA NA NA NA NY104 NA E Ro S G MtO BY* 10.0 86.7 0.3 2.2 LPB11 Sa E A M G NA NA 10.0 60.0 0.3 2.6 LPB19 NA NA NA NA NA NA NA NA NA NA NA NY25 NA NA NA NA NA NA NA NA NA NA NA B4 Tr E A S G NA NA 10.0 85.0 0.3 2.4 LPB18 NA NA NA NA NA NA NA NA NA NA NA

Appendix 4 continued.

OTUs KPetLamBas§ KPetLamMar§ KPetLamApx§ OvPub† StylPub† SeedShp SeedCol SepLobApxAng WPetLamArcApxAng WPetClwW* KPetLamL* MU17 NA NA NA NA NA NA NA NA NA NA NA NY90 Sa E Ro S G Mt DB 10.0 60.0 0.3 2.8 US165 Tr E RoA G G NA NA 10.0 80.0 0.4 3.3 NY79 NA NA NA NA NA NA NA NA NA NA NA LPB4 Tr E Ro S G NA NA 10.0 110.0 0.3 2.2 NY91 Tr E Ro G G NA NA 10.0 65.0 0.3 1.9 F52 Tr E A M G NA NA 10.0 90.0 0.3 1.9 NY21 Sa E A G G NA NA 10.0 40.0 0.3 2.2 MU23 NA NA NA NA NA NA NA NA NA NA NA AHUC2 Tr E Ro GS G Mt LB 10.0 70.0 0.3 1.9 LPB14 Tr E Ro S S NA NA 10.0 95.0 0.4 1.9 F57 Tr E A S G NA NA 10.0 90.0 0.3 2.4 LPB15 Sa E Ro S G NA NA 10.0 60.0 0.3 2.5 LPB17 Sa E RoA S S NA NA 10.0 85.0 0.5 2.2 MU6 Tr E AcA S G NA NA 10.0 100.0 0.4 2.9 MO43 Tr E AcA S G NA NA 10.0 95.0 0.4 3.1 LPB6 NA NA NA NA NA NA NA NA NA NA NA NY98 Sa E A G G NA NA 10.0 70.0 0.5 2.7 US4 Tr E RoA S G NA NA 10.0 120.0 0.4 3.6 P62 Tr E A M S NA NA 10.0 50.0 0.5 3.2 P86 Sa E A M G NA NA 10.0 65.0 0.5 3.9 P6 Tr E A S G NA NA 10.0 95.0 0.5 3.6 NY69 NA E A NA G NA BY 10.0 92.0 0.5 3.6 US8 Sa E A S S NA NA 10.0 85.0 0.5 3.5 US100 Tr E Ro S G Mt LB 10.0 60.0 0.4 2.7 US129 Tr E AcA S G NA NA 10.0 55.0 0.6 4.7 F45 Tr E AcA M G NA NA 10.0 60.0 0.5 3.9 AAU6 Tr E A S G NA NA 10.0 60.0 0.5 3.8 US2 Tr E A M S NA NA 10.0 60.0 0.5 3.5 F44 Tr E A M G Mt LB 10.0 90.0 0.4 3.5 F47 Tr E AcA M S Mt LB 10.0 60.0 0.5 3.6 US35 Tr E A S S Mt LB 10.0 93.3 0.5 3.5 MO10 Tr E A S S NA NA 10.0 63.3 0.5 3.4 NY101 Sa E AcA M S NA NA 10.0 60.0 0.5 3.5 F135 Tr E A S G NA NA 10.0 105.0 0.4 3.6

100

Appendix 4 continued.

OTUs KPetLamBas§ KPetLamMar§ KPetLamApx§ OvPub† StylPub† SeedShp SeedCol SepLobApxAng WPetLamArcApxAng WPetClwW* KPetLamL* P83 Tr E A S G NA NA 10.0 90.0 0.5 3.5 F91 Tr E Ro S G NA NA 10.0 75.0 0.5 3.6 F104 Sa E A S G Mt LB 10.0 52.5 0.5 3.6 MO37 Sa E AcA S G NA NA 10.0 60.0 0.4 3.2 F114 Tr E Ro G G NA NA 10.0 95.0 0.4 3.2 F126 Sa E A S S Mt LB* 10.0 80.0 0.4 3.6 F109 Tr E A S G NA NA 10.0 80.0 0.5 3.7 US131 NA NA NA NA NA NA NA NA NA NA NA F122 NA NA NA NA NA NA NA NA NA NA NA F111 Sa E Ro S S NA NA 10.0 75.0 0.7 4.4 F110 Tr E RoA M G NA NA 10.0 93.3 0.7 4.4 B11 Sa E AcA G G NA NA 10.0 70.0 0.4 3.9 NY122 Sa E RoA M G NA NA 10.0 110.0 0.5 4.0 US13 Tr E AcA M G NA NA 10.0 80.0 0.5 3.2 US26 Tr E AcA M G NA NA 10.0 60.0 0.4 3.1 F124 Sa E AcA M G NA NA 10.0 105.0 0.5 3.8 US20 NA E RoA S S NA NA 10.0 94.0 0.5 3.5 US21 Tr E AcA M G NA NA 10.0 50.0 0.4 3.6 US129 Tr E Ro S G NA NA 10.0 75.0 0.5 2.9 F133 NA E A GS S NA NA 10.0 108.0 0.4 3.7 MO32 Tr E A G G NA NA 10.0 65.0 0.4 3.8 MO19 Sa E A G G NA NA 10.0 140.0 0.5 3.7 US34 Sa E A GS G NA NA 10.0 90.0 0.6 4.0 F98 NA E NA S S NA NA 9.8 120.0 0.4 3.5 US132 Tr E Ro S G NA NA 10.0 85.0 0.5 3.0 LPB3 Tr E A S G NA NA 10.0 90.0 0.4 3.6 NY97 Tr E RoA S G MtO LB 10.0 90.0 0.5 3.6 P8 Tr E A G G Mt LB 10.0 40.0 0.5 3.0 MU19 NA E NA G G NA NA 10.0 102.0 0.4 3.7 NY63 Tr E RoA S G NA NA 10.0 90.0 0.4 3.6 SI6 Tr E AcA M G NA NA 10.0 90.0 0.4 3.0 LPB22 Tr E Ro S G NA NA 10.0 85.0 0.5 3.5 US19 Tr E A S G NA NA 10.0 100.0 0.4 3.3 MU21 NA NA NA NA NA NA NA NA NA NA NA NY24 Tr E Ro S G NA NA 10.0 105.0 0.4 3.0

101

Appendix 4 continued.

OTUs KPetLamBas§ KPetLamMar§ KPetLamApx§ OvPub† StylPub† SeedShp SeedCol SepLobApxAng WPetLamArcApxAng WPetClwW* KPetLamL* MO3 NA NA NA NA NA NA NA NA NA NA NA F2 Tr E AcA S G NA NA 10.0 75.0 0.4 3.1 F90 NA NA NA NA NA NA NA NA NA NA NA MU34 Sa E A G G NA NA 10.0 110.0 0.3 2.7

102

Appendix 4 continued.

OTUs ConFilPcFus FreFilL* FreFilD* OvL* OvW* StylL* StylD* OvlNo FruitL* FruitW* SeedpFruit SeedL* SeedW* US14 29.7 2.7 0.1 2.6 0.6 1.0 0.1 4.0 NA NA NA NA NA NY51 40.3 2.0 0.1 2.2 0.5 0.9 0.1 2.0 NA NA NA NA NA F121 39.7 2.1 0.1 2.4 0.5 0.9 0.1 2.5 NA NA NA NA NA NY66 31.5 2.4 0.1 2.6 0.8 1.1 0.1 2.5 NA NA NA NA NA MU27 34.9 2.1 0.1 2.6 0.8 1.0 0.2 3.0 NA NA NA NA NA US171 NA NA NA NA NA NA NA NA NA NA NA NA NA US169 43.4 2.4 0.1 2.8 0.9 1.0 0.2 2.0 NA NA NA NA NA F118 44.4 2.4 0.2 NA NA NA NA 3.0 4.7 2.9 2.0 1.9 1.7 MO44 NA NA NA NA NA NA NA NA NA NA NA NA NA P97 33.7 2.4 0.1 2.2 0.5 1.3 0.1 3.0 NA NA NA NA NA NY60 42.3 2.3 0.1 4.0 1.6 0.9 0.2 3.5 NA NA NA NA NA MU26 41.3 2.4 0.1 2.7 1.0 1.3 0.2 2.0 NA NA NA NA NA US32 37.3 2.2 0.1 2.6 0.6 1.0 0.2 4.0 NA NA NA NA NA MO48 45.9 2.8 0.1 2.9 1.1 1.1 0.2 2.0 NA NA NA NA NA NY119 39.2 2.5 0.1 2.2 0.6 1.3 0.2 2.0 4.7 2.4 2.0 2.1 1.7 MO49 NA NA NA NA NA NA NA NA NA NA NA NA NA MUX NA NA NA NA NA NA NA NA NA NA NA NA NA NYX NA NA NA NA NA NA NA NA NA NA NA NA NA US29 NA NA NA NA NA NA NA NA NA NA NA NA NA NY50 NA NA NA NA NA NA NA NA NA NA NA NA NA NY89 36.1 2.3 0.1 2.7 0.7 0.8 0.2 4.0 NA NA NA NA NA US30 39.3 2.8 0.1 2.7 1.0 1.4 0.2 2.0 NA NA NA NA NA US28 44.4 2.7 0.1 2.7 0.8 1.4 0.1 3.0 NA NA NA NA NA F48 43.7 2.3 0.1 2.0 0.6 0.9 0.2 2.0 NA NA NA NA NA MU9 33.3 2.2 0.1 1.9 0.6 1.0 0.2 3.0 NA NA NA NA NA F131 NA NA NA NA NA NA NA NA NA NA NA NA NA NY85 NA NA NA NA NA NA NA NA NA NA NA NA NA LPB8 32.9 2.1 0.1 2.1 0.7 0.9 0.2 2.0 NA NA NA NA NA MU10 NA NA NA NA NA NA NA NA NA NA NA NA NA NY104 33.0 2.2 0.4 3.0 0.9 0.9 0.1 2.0 5.4 2.3 2.0 1.9 1.5 LPB11 30.3 2.4 0.1 2.3 0.7 1.0 0.1 2.5 NA NA NA NA NA LPB19 NA NA NA NA NA NA NA NA NA NA NA NA NA NY25 NA NA NA NA NA NA NA NA NA NA NA NA NA B4 30.3 1.9 0.1 1.9 0.7 0.9 0.1 2.0 NA NA NA NA NA LPB18 NA NA NA NA NA NA NA NA NA NA NA NA NA

103

Appendix 4 continued.

OTUs ConFilPcFus FreFilL* FreFilD* OvL* OvW* StylL* StylD* OvlNo FruitL* FruitW* SeedpFruit SeedL* SeedW* MU17 NA NA NA NA NA NA NA NA NA NA NA NA NA NY90 36.1 2.8 0.1 2.7 0.8 1.0 0.1 4.0 5.8 2.3 3.0 1.5 1.3 US165 38.3 2.3 0.1 2.1 0.8 1.9 0.2 2.5 NA NA NA NA NA NY79 NA NA NA NA NA NA NA NA NA NA NA NA NA LPB4 39.4 2.2 0.1 2.1 0.7 0.8 0.2 3.0 NA NA NA NA NA NY91 41.9 1.8 0.1 2.4 1.0 0.9 0.2 2.5 NA NA NA NA NA F52 35.7 1.8 0.1 1.7 0.5 1.0 0.1 2.0 NA NA NA NA NA NY21 37.2 2.3 0.4 1.8 0.5 1.3 0.1 2.2 NA NA NA NA NA MU23 NA NA NA NA NA NA NA NA NA NA NA NA NA AHUC2 42.6 1.9 0.1 2.1 0.9 1.0 0.1 2.0 3.9 2.2 2.0 1.6 1.2 LPB14 47.2 2.2 0.1 2.0 1.0 1.0 0.2 3.0 NA NA NA NA NA F57 33.3 2.4 0.1 1.8 0.6 1.1 0.1 2.0 NA NA NA NA NA LPB15 37.8 2.3 0.1 2.0 0.6 0.9 0.2 3.0 NA NA NA NA NA LPB17 25.4 2.3 0.1 2.3 0.7 1.0 0.1 2.5 NA NA NA NA NA MU6 23.3 2.5 0.1 2.1 0.7 1.4 0.1 3.0 NA NA NA NA NA MO43 27.1 2.7 0.1 2.4 0.8 1.3 0.2 3.0 NA NA NA NA NA LPB6 NA NA NA NA NA NA NA NA NA NA NA NA NA NY98 44.6 2.3 0.1 1.8 0.7 1.1 0.2 2.7 NA NA NA NA NA US4 30.9 2.7 0.1 2.5 0.7 1.6 0.2 2.0 NA NA NA NA NA P62 36.6 2.5 0.1 2.1 0.7 1.5 0.1 2.5 NA NA NA NA NA P86 40.7 3.1 0.1 2.3 0.9 1.5 0.2 2.0 NA NA NA NA NA P6 34.1 2.6 0.1 1.9 0.7 1.2 0.2 2.5 NA NA NA NA NA NY69 43.4 2.9 0.2 1.5 0.4 1.3 0.1 2.0 5.2 1.9 2.0 1.9 1.4 US8 41.3 2.5 0.1 2.1 0.8 1.6 0.2 2.5 NA NA NA NA NA US100 41.0 2.5 0.1 NA NA NA NA 2.0 NA NA 2.0 2.0 1.7 US129 28.2 3.4 0.1 2.5 0.8 2.0 0.2 2.0 NA NA NA NA NA F45 29.4 2.8 0.1 2.4 0.7 1.5 0.2 2.0 NA NA NA NA NA AAU6 39.3 2.5 0.2 2.0 1.1 1.1 0.2 2.5 NA NA NA NA NA US2 42.9 2.9 0.1 2.4 0.8 1.4 0.2 2.0 NA NA NA NA NA F44 35.1 2.4 0.1 1.9 0.8 1.3 0.2 2.0 NA NA NA 2.0 1.6 F47 44.0 2.0 0.1 2.0 0.7 1.2 0.2 2.0 4.7 2.3 2.0 1.8 1.5 US35 43.9 3.5 0.1 3.4 1.8 1.7 0.2 2.0 6.1 3.0 2.0 2.4 1.5 MO10 39.2 2.5 0.1 1.7 0.5 1.3 0.1 2.0 NA NA NA NA NA NY101 35.6 1.9 0.2 1.5 0.6 1.2 0.1 2.0 NA NA NA NA NA F135 41.0 2.3 0.1 2.3 0.8 2.2 0.2 2.5 NA NA NA NA NA

104

Appendix 4 continued.

OTUs ConFilPcFus FreFilL* FreFilD* OvL* OvW* StylL* StylD* OvlNo FruitL* FruitW* SeedpFruit SeedL* SeedW* P83 36.4 3.0 0.1 2.9 0.9 1.3 0.2 4.0 NA NA NA NA NA F91 34.0 2.9 0.1 2.2 0.7 1.9 0.3 3.0 NA NA NA NA NA F104 33.7 2.7 0.1 2.4 0.7 1.4 0.2 4.0 5.3 2.7 2.0 2.0 1.6 MO37 35.6 2.7 0.1 2.0 0.7 1.5 0.2 2.0 NA NA NA NA NA F114 35.2 NA NA 2.1 0.7 1.4 0.1 2.0 NA NA NA NA NA F126 44.5 2.5 0.1 3.1 1.5 1.1 0.1 2.0 6.0 3.2 2.0 2.3 1.9 F109 44.0 2.4 0.2 2.2 0.9 1.4 0.2 2.0 NA NA NA NA NA US131 NA NA NA NA NA NA NA NA NA NA NA NA NA F122 NA NA NA NA NA NA NA NA NA NA NA NA NA F111 38.5 3.1 0.2 2.7 0.9 1.9 0.2 2.5 NA NA NA NA NA F110 34.4 3.0 0.1 2.7 1.0 1.4 0.2 2.7 NA NA NA NA NA B11 36.2 3.1 0.1 2.3 0.7 2.0 0.2 2.0 NA NA NA NA NA NY122 32.0 3.5 0.1 3.5 1.0 2.2 0.2 2.0 NA NA NA NA NA US13 39.3 3.1 0.1 1.7 0.7 1.9 0.2 2.5 NA NA NA NA NA US26 40.0 2.7 0.1 2.6 1.4 1.6 0.2 2.0 NA NA NA NA NA F124 42.4 2.8 0.2 2.2 0.8 1.5 0.2 2.0 NA NA NA NA NA US20 45.6 2.4 0.2 2.4 0.8 1.3 0.2 2.0 3.9 1.3 2.5 1.2 0.9 US21 61.9 1.9 0.1 2.4 0.9 1.3 0.2 2.0 NA NA NA NA NA US129 39.7 2.5 0.1 2.1 0.8 1.3 0.2 2.0 NA NA NA NA NA F133 40.3 2.8 0.1 2.7 0.9 1.6 0.2 2.0 3.8 1.5 2.0 1.8 2.3 MO32 32.3 3.3 0.1 2.4 0.8 2.2 0.2 2.5 NA NA NA NA NA MO19 26.8 3.3 0.1 2.3 0.6 2.3 0.2 3.0 NA NA NA NA NA US34 34.6 2.6 0.1 2.3 0.9 2.0 0.1 2.6 NA NA NA NA NA F98 34.5 3.4 0.1 2.7 1.0 1.7 0.2 4.4 NA NA NA NA NA US132 39.5 2.6 0.1 2.2 0.9 1.2 0.2 2.0 NA NA NA NA NA LPB3 37.9 2.6 0.1 2.3 0.7 1.8 0.1 2.5 NA NA NA NA NA NY97 38.4 3.1 0.2 2.4 0.7 1.6 0.1 3.0 4.1 1.9 1.0 1.5 1.2 P8 35.7 3.9 0.1 2.1 0.5 1.8 0.1 2.0 3.3 2.4 1.0 1.8 1.5 MU19 37.6 2.3 0.5 2.3 0.9 1.5 0.2 3.4 NA NA NA NA NA NY63 32.5 2.5 0.1 3.0 1.1 1.1 0.1 2.5 NA NA NA NA NA SI6 26.6 2.9 0.1 2.7 0.7 1.0 0.2 2.0 NA NA NA NA NA LPB22 33.7 3.2 0.2 2.5 0.6 1.5 0.2 3.0 NA NA NA NA NA US19 34.1 2.7 0.1 2.4 0.7 1.6 0.1 2.0 NA NA NA NA NA MU21 NA NA NA NA NA NA NA NA NA NA NA NA NA NY24 37.3 2.6 0.1 2.3 0.7 1.2 0.1 2.5 NA NA NA NA NA

105

Appendix 4 continued.

OTUs ConFilPcFus FreFilL* FreFilD* OvL* OvW* StylL* StylD* OvlNo FruitL* FruitW* SeedpFruit SeedL* SeedW* MO3 NA NA NA NA NA NA NA NA NA NA NA NA NA F2 38.8 2.4 0.2 2.4 0.9 1.3 0.2 3.0 NA NA NA NA NA F90 NA NA NA NA NA NA NA NA NA NA NA NA NA MU34 37.3 2.2 0.1 2.2 0.7 1.3 0.2 2.0 NA NA NA NA NA

106

Appendix 5. Results from PCS analysis from each species run against each other species. Groups are represented by different colors; T. amabile (blue), T. peruvianum (purple), T. tafiense (black), T. cochabambense (orange), T. andesicola(green), T. australe (brown).

3D PCA Scatterplot of T. amabile vs. T. peruvianum

107

Appendix 5 continued. 3D PCA Scatterplot of T. amabile vs. T. cochabambense

108

Appendix 5 continued. 3D PCA Scatterplot of T. amabile vs. T. andesicola

109

Appendix 5 continued. 3D PCA Scatterplot of T. amabile vs. T. australe

110

Appendix 5 continued. 3D PCA Scatterplot of T. amabile vs. T. tafiense

111

Appendix 5 continued. 3D PCA Scatterplot of T. cochabambense vs. T. australe

112

Appendix 5 continued. 3D PCA Scatterplot of T. cochabambense vs. T. andesicola

113

Appendix 5 continued. 3D PCA Scatterplot of T. cochabambense vs. T. tafiense

114

Appendix 5 continued. 3D PCA Scatterplot of T. cochabambense vs. T. peruvianum

115

Appendix 5 continued. 3D PCA Scatterplot of T. tafiense vs. T. australe

116

Appendix 5 continued. 3D PCA Scatterplot of T. tafiense vs. T. andesicola

117

Appendix 5 continued. 3D PCA Scatterplot of T. tafiense vs. T. peruvianum

118

Appendix 5 continued. 3D PCA Scatterplot of T. peruvianum vs. T. andesicola

119

Appendix 5 continued. 3D PCA Scatterplot of T. peruvianum vs. T. australe

120

Appendix 5 continued. 3D PCA Scatterplot of T. andesicola vs. T. australe

121