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Ecological Factors, Mixed Breeding System and Population

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Ecological Factors, Mixed Breeding System and Population ECOLOGICAL FACTORS, MIXED BREEDING SYSTEM AND POPULATION GENETIC STRUCTURE IN A SUBTROPICAL AND A TEMPERATE VIOLET SPECIES A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Aurea C. Cortés-Palomec June 2005 This dissertation entitled ECOLOGICAL FACTORS, MIXED BREEDING SYSTEM AND POPULATION GENETIC STRUCTURE IN A SUBTROPICAL AND A TEMPERATE VIOLET SPECIES by AUREA C. CORTÉS-PALOMEC has been approved for the Department of Biological Sciences and the College of Arts and Sciences by Harvey E. Ballard Jr. Associate Professor of Environmental and Plant Biology Leslie A. Flemming Dean, College of Arts and Sciences CORTÉS-PALOMEC, AUREA, C. Ph.D. June 2005. Biological Sciences Ecological factors, mixed breeding system and population genetic structure in a subtropical and a temperate violet species (186 pp.) Director of Dissertation: Harvey E. Ballard, Jr. A mixed breeding system involving the production of chasmogamous and cleistogamous flowers is common in most species of the genus Viola. This system can theoretically affect the patterns of reproduction and distribution of genetic variation in populations. While we understand the theoretical basis behind these expected patterns, there is little empirical evidence comparing the behavior of similar species in widely contrasting environments. To better characterize the effect that this breeding system has on reproduction and genetics a subtropical (Viola grahamii) and a temperate (V. striata) species of violets were compared. V. striata populations were studied in Ohio, USA, and V. grahamii in Michoacán, Mexico during 2002 and 2003. Weekly observations during the reproductive season allowed for the establishment of blooming patterns of the two flower types for each species. The allocation of resources towards flowering was evaluated under natural differential light availability and soil nutrients. The reproductive success of each of the flower types was determined and the genetic diversity in both species assessed. Un sistema reproductivo que incluye la producción de flores casmógamas y cleistógamas en un mismo individuo es común en la mayoría de las especies del género Viola. En teoría, el balance en la producción de estos dos tipos florales afecta dramáticamente la estrategia reproductiva de cada especie y tiene serias repercusiones en su diversidad genética. Para caracterizar este sistema reproductivo, dos especies cercanas de violetas creciendo en ambientes contrastantes fueron escogidas para este estudio: Viola grahamii, una especie subtropical y Viola striata, una especie de climas templados. Las poblaciones de V. grahamii estudiadas se encuentran en el estado de Michoacán en México mientras que las poblaciones de V. striata se localizan en Ohio en Estados Unidos. Censos semanales durante la época reproductiva permitieron establecer los patrones de floración de cada tipo floral para ambas especies. La cantidad de flores producidas de cada tipo fue relacionada con la cantidad de nutrientes y luz en las cuales los diferentes individuos crecían. El éxito de cada tipo floral en producir semillas fue evaluado y la diversidad genética en las distintas poblaciones fue analizada en relación con el sistema reproductivo. Approved: Harvey E. Ballard, Jr. Associate Professor of Environmental and Plant Biology Acknowledgments This work would not have been possible without the assistance of several people. I would like to thank my advisor, Harvey E. Ballard, Jr. for introducing me to the world of violets and for all his help and support during these years. To members of my graduate committee, Kim Brown, Gar Rothwell, Royal Mapes and Kelly Johnson for valuable comments during different stages of the dissertation, in particular to Kim Brown for her encouragement, support and comments during the development of the ecological projects. I extend a special thanks to Gene Mapes for accepting to serve on my committee in place of Gar Rothwell who was unable to attend my final defense on account of a previous foreign travel commitment. I would also like to express my gratitude to the Department of Environmental and Plant Biology, current and past students for all their assistance and advice through my time in the department. I would like to say a special thanks to Theresa Culley from the University of Cincinnati for sharing with me her Viola pubescens microsatellite primers, for her insightful comments about violets and most of all for instructing me in the use of allozyme techniques during two weeks spent working in her lab. I am indebted to the Consejo Nacional de Ciencia y Tecnologia (CONACyT- Mexico) for providing me with graduate program funding under the grant No. 128098 for most of the duration of my program, and to the Department of Environmental & Plant Biology for taking over once my grant was over. Funding for research was provided by grant awards from the Sigma Xi Society (Grant-in-aid of Research), The Botanical Society of America (Karling Award), the Office of Research and Sponsored Programs at Ohio University, and the Wilson Fund from the department of Environmental and Plant Biology. Additional funding in support of field and laboratory work was provided by my advisor Harvey E. Ballard, Jr. Thanks to Sergio Zamudio from the Instituto de Ecología, A.C. Centro Regional del Bajío, in Pátzcuaro, Michoacán for his logistical support during my time completing field work in Mexico. Special thanks to my very good friend José Ricardo Gonzalo Wong from the Universidad Nacional Autónoma de México (UNAM) for his longtime friendship and for helping in the establishment my study sites in Mexico, especially for getting up early with me to go data collecting while being relentlessly pursued by Oso and other unidentified dogs during the summer of 2002. Thanks to Judith Márquez Guzmán from UNAM for all her support and early encouragement to pursue a doctorate degree and for being my academic mentor over the years. I would also like to thank my family for their continued support in every task I have started during my life. And finally, but not least, I would like to thank my husband, Ross A. McCauley for his help in establishing my study sites in Ohio, and in data collection during the summer of 2003 in Mexico. But most of all, I want to thank him for his unconditional love, friendship, support, and for believing in me at every moment along the way. vii Table of Contents Page Abstract............................................................................................................................iii Acknowledgments ............................................................................................................v List of Tables....................................................................................................................x List of Figures ................................................................................................................xiv Chapter 1: Introduction.................................................................................................1 Theoretical background of a Mixed Breeding System in the genus Viola...........1 Research site establishment and description ........................................................9 Species and Sites of Study........................................................................9 Viola grahamii Bentham ..............................................................9 Viola striata Aiton......................................................................10 Viola grahamii Study Site ..........................................................11 Viola striata Study Site ..............................................................20 Quadrat establishment ................................................................28 Chapter 2: Environmental correlates to leaf mass area and leaf nitrogen in the Neotropical violet Viola grahamii....................................................................30 Introduction ........................................................................................................30 Materials and Methods .......................................................................................32 Species of Study .....................................................................................32 Study Sites..............................................................................................33 Sampling, measurements and data handling ..........................................34 Results ................................................................................................................37 Discussion...........................................................................................................46 Chapter 3: Influence of annual fluctuations in nutrient availability on the mixed breeding system of the Neotropical violet species Viola grahamii, and on the chasmogamous flower production in the temperate forest violet Viola striata..................................................................................................................52 Part A. Viola grahamii .......................................................................................52 Introduction ............................................................................................52 Materials and Methods ...........................................................................56 Species of Study .........................................................................56
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