ABSTRACT POST, ANGELA R. a Systematic
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ABSTRACT POST, ANGELA R. A Systematic Study of Cardamine Weed Species in United States Container Nurseries. (Under the direction of Dr. Joseph C. Neal). Cardamine species present in the United States nursery trade were identified and characterized based on morphological and molecular evidence. The genus Cardamine L., or the bittercresses, contains some of the most prolific weed species in the container nursery industry. Though most bittercress in the U.S. nursery industry was thought to be hairy bittercress (Cardamine hirsuta), we suspected other species to be present due to the global movement of nursery stock. We surveyed 21 nurseries in California, Mississippi, Missouri, New York, North Carolina, and Oregon to collect a representative sample of bittercress species occurring in the U.S. nursery industry. Type material and other herbarium samples were examined for all species encountered in order to identify each accession and determine which characters were most important for identification. From these nursery surveys four Cardamine species were collected: Cardamine corymbosa, C. hirsuta, C. flexuosa and C. oligosperma, each representing 90%, 6%, 2%, and 2% of collections respectively. C. flexuosa was originally described in Europe but also has distribution throughout Asia. European and Asian populations may represent separate taxa based on molecular evidence. Here we treat them as the same taxon. Type material no longer exists for C. flexuosa as it was described in Europe. A lectotype is designated here. Cardamine corymbosa, native to New Zealand, is a newly reported species for the United States. Cardamine hirsuta has worldwide distribution and C. oligosperma is native to the United States. Representatives of each species collected, as well as select herbarium specimens, were chosen for a molecular study to examine the relationships between these species. Sequence was generated for the nrDNA Internal Transcribed Spacer region (ITS) and for the single copy nuclear COP1-Interacting Protein 7 region (CIP7). Sequence data was aligned and subjected to maximum parsimony and bayesian inference. When comparing the phylogenies created by ITS and CIP7 datasets, there were areas of congruence and incongruence. The ITS data resolved C. corymbosa as sister to C. flexuosa whereas the CIP7 data resolved C. corymbosa sister to C. hirsuta. The combined alignment resolved the Cardamine accessions with higher bootstrap (BS) support and posterior probability (PP) than either of the separate matrices alone. In the combined matrix C. hirsuta and C. corymbosa resolve as sister taxa. However, no analysis fully resolved the relationships within this closely related species complex. A Systematic Study of Cardamine Weed Species in United States Container Nurseries by Angela Rose Post A thesis submitted to the Graduate Faculty of North Carolina State University In partial fulfillment of the Requirements for the degree of Master of Science Horticultural Science Raleigh, North Carolina June 23rd 2008 APPROVED BY: _______________________________ ______________________________ Dr. Joseph C. Neal Dr. Alexander Krings Committee Chair _______________________________ ______________________________ Dr. Bryon R. Sosinski Dr. Qiu-Yun (Jenny) Xiang BIOGRAPHY Angela R. Post was born Angela Rose Richardson in Asheboro North Carolina and grew up in the nearby rural town of Franklinville. From childhood Angela had a love of plants and always enjoyed flower and vegetable gardening with her grandmother. Her high school horticulture program set her on the path to become a plant scientist and in 2001 she began an undergraduate program in Horticultural Science at North Carolina State University. One year later Angela changed her major to Botany for a more rigorous math and science base. She completed her B. S. Botany in 2003 followed by a B. S. in Biological Sciences in 2004 and a B.A. in Multidisciplinary Studies with a concentration in Medicinal Ethnobotany in 2005. Angela returned to North Carolina State as a teaching assistant and research technician in Fall 2006 before starting her M. S. program in Horticultural Science. Under the direction of Dr. Joseph Neal Angela studied the systematics and taxonomy of weedy Cardamine species in United States container nurseries. During her time as a master’s student Angela traveled to five major herbaria to study her group and presented research at nine local, regional, and national meetings. Just before her program began she was married to John Richard Post and a year and a half later they welcomed their first child Liana Rose Post to the family. Upon completion, Angela will move on to Cornell University to pursue Ph.D. in Plant Biology with a focus on systematics. In the future Angela hopes to become a university professor with a teaching and research appointment. ii TABLE OF CONTENTS List of Figures …………………………………………………………………… v Abbreviations ……………………………………………………………………. vii Scope and Justification ………………………………………………………….. 1 Literature Cited ………………………………………………………… 9 Chapter 1. On the Identity of the weedy Cardamine (Brassicaceae) in United States nurseries: Evidence from molecules and morphology……………………… 13 Abstract …………………………………………………………………. 15 Introduction ……………………………………………………………… 15 Materials and Methods ………………………………………………….. 22 Results ……………………………………..……………….…………… 26 Discussion ……………………………………………………………….. 27 Taxonomic Treatment……………………………………………………. 30 Acknowledgements ……………………………………………………... 37 Literature Cited …………………………………………………………. 38 Chapter 2. Lectotypification of Cardamine flexuosa Withering (Brassicaceae)….. 61 Abstract …………………………………………………………………. 63 Introduction ……………………………………………………………... 63 Materials and Methods ………………………………………………….. 63 Results …………………………………………………………………… 63 Discussion ……………………………………………………………….. 64 Acknowledgements …………………………………………………….. 65 Literature Cited …………………………………………………………. 66 Chapter 3. Cardamine corymbosa (Brassicaceae): New to the United States….. 71 Abstract …………………………………………………………………. 73 Introduction ……………………………………………………………... 73 Description………………………………………………………………. 74 Discussion ……………………………………………………………….. 75 Acknowledgements ……………………………………………………… 76 Literature Cited …………………………………………………………. 77 Discussion and Conclusions……………………………………………………... 81 iii Literature Cited …………………………………………………………. 85 Appendix A. Exsiccate…………………………………………………………. 86 Appendix B. Photographs………………………………………………………. 88 iv LIST OF FIGURES Page Chapter 1. Figure 1.1 Strict consensus of 700,000 most parsimonious trees resulting from parsimony analysis of nuclear gene region CIP7 for Cardamine species in United States nurseries (Length=59; CI= 0.814; RI=0.875). Bootstrap support (BS) is given above branches and Bayesian posterior probabilities (PP) are given below branches. Species identifications shown are based on morphological examinations. State abbreviations are given in parenthesis next to accessions. UK denotes fresh accessions from England and H denotes herbarium specimens.………………………………………….……….. 45 Figure 1.2 Majority Rule consensus for ITS for Cardamine species in United States nurseries including our fresh accessions labeled with state acronyms and selected ones, labeled “Lih”, from Lihova et al. (2006). UK denotes fresh accessions from England and OLK and HIR are herbarium sheets. Bayesian posterior probabilities (PP) are given above branches. Species identifications shown are based on morphological examinations……………………………….……………………………………… 47 Figure 1.3. Strict consensus of 13 most parsimonious tree resulting from parsimony analysis for combined matrix of CIP7and ITS for Cardamine species in United States nurseries (Length=73; CI= 0.922; RI=0.978). Bootstrap support (BS) is given above branches and Bayesian posterior probabilities (PP) are given below branches. Species identifications shown are based on morphological examinations. Morphological characters are given under species identification. State abbreviations are given in parenthesis after each accession. UK denotes fresh accessions from England and H denotes herbarium specimens. ………………………………………...…………… 49 Figure 1.4. Illustration of Cardamine corymbosa Hook.f. a) habit, b) basal leaf, c) petaliferous flower-cutout, d) petaliferous flower-top view, e) apetalous flower, f) apetalous flower-top view, g) apetalous flower-cutout, h) stamen, i) silique, j) silique- cutout, k) seed. [Illustrations by Nancy C. Routh] …...…………………………… 51 Figure 1.5. Illustration of Cardamine flexuosa With. a) habit, b) basal leaf, c) cauline leaf, d) flower- cutout, e) flower-side view, f) flower-top view, g) petal, h) stamens, i) seed. [Illustrations by Nancy C. Routh] …………………………..... 53 Figure 1.6. Illustration of Cardamine hirsuta L. a) habit, b) basal leaf, c) cauline leaf, d) flower-cutout, e) flower-side view, f) flower-top view, g) petal, h) stamens, i) silique, j) silique-cutout, k) seeds. [Illustrations by Nancy C. Routh]……………….………………………………………………………..…… 55 v LIST OF FIGURES (continued) Page Figure 1.7. Illustration of Cardamine oligosperma L. a) habit, b) basal leaf, c) cauline leaf, d) flower-cutout, e) flower-side view, f) flower-top view, g) petal, h) stamen, i) silique. [Illustrations by Nancy C. Routh]………………………………...……. 57 Figure 1.8. Illustration of Cardamine scutata Thunb. a) habit, b) basal leaf, c) cauline leaf, d) flower-cutout, e) flower-side view, f) flower-top view, g) petal, h) stamen, i) silique, j) seed. [Illustrations by Nancy C. Routh]…...……………………..… 59 Chapter 2. Figure 2.1. Digital image of the lectotype for Cardamine flexuosa in W. Curtis’ Flora