THE ECOLOGICAL STUDY OF THE MARITIME RINGLET BUTTERFLY (COENONYMPHA NIPISIQUIT MCDUNNOUGH) IN DALY POINT, BATHURST, NEW BRUNSWICK. A Dissertation Presented by MAKIRI SEI Submitted to the Graduate School of the University of Massachusetts Amherst in partial fulfillment of the requirements for the joint degree of DOCTOR OF PHILOSOPHY February 2006 Organismic and Evolutionary Biology UMI Number: 3206207 3206207 2006 UMI Microform Copyright All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, MI 48106-1346 by ProQuest Information and Learning Company. © Copyright by Makiri Sei 2006 All Rights Reserved THE ECOLOGICAL STUDY OF THE MARITIME RINGLET BUTTERFLY (COENONYMPHA NIPISIQUIT MCDUNNOUGH) IN DALY POINT, BATHURST, NEW BRUNSWICK A Dissertation Presented by MAKIRI SEI Approved as to style and content by: ____________________________________________ Adam H. Porter, Chair ____________________________________________ Robin A. Harrington, Member ____________________________________________ Paul R. Sievert, Member ____________________________________________ Roy G. Van Driesche, Member _______________________________________ Margaret A. Riley, Director Organismic and Evolutionary Biology DEDICATION To my parents, Toshio and Noriko Sei, who wholeheartedly supported me whiled I studied abroad. v ACKNOWLEDGMENTS I would like to express my thanks to my advisor, Adam H. Porter, for his many years of very positive, thoughtful, and patient guidance and generosity. I would like to extend my gratitude to the members of my committee, Robin H. Harrington, Paul R. Sievert, and Roy G. Van Driesche for their helpful discussions and suggestions on all aspects of this endeavor. My dissertation has improved through communications with Mitchell Baker, Thomas Crist, Christy Hentzler, Daniel P. Jennings, Christopher Majka, Ben Normark, Ann Rypstra, Baiqing Wang, Reginald P. Webster, Arnold D. Well, and Sean Werle, who generously shared their expertise. The maps in Figure 1, 2, and 6 were reproduced with permission from RPW. I am indebted to the Xerces Society, the National Science Foundation, the Graduate Program of Organismic and Evolutionary Biology, and the Department of Natural Resources and Energy of New Brunswick for funding this research. This research was conducted under two unidentified permits issued in 1999 and 2000, #2001-06, #2002-11, and #ES03-014 issued from the Department of Natural Resource and Energy, New Brunswick and #2003-05-15-502-11-S-F issued from Direction de l’Aménagement de la Faune de la Gaspésie-Îles-de-la-Madeleine, Québec. I would like to thank Maureen Toner, Dwayne Sabine, Luc Gagnon, Gilles Godin, Nathalie D’Aoust of the Department of Natural Resource and Energy, New Brunswick, O’Neil Pelletier and Julie Gaudet of Community College of New Brunswick, Bathurst, and Stan Georges of Société de la Faune et des Parcs du Québec for their generous support of my fieldwork in Canada. vi I would like to express my appreciation to Nathalie Arsenault, Christian Brideau, Christopher Drysdall, Paul Ferron, Alain Gouge, Patricia Levigne, Alexandre Haché, and Melanie Boudreau for their participation in this project. Special thanks to Jeremy Houser, Christa Skow, Baiqing Wang, Elizabeth G. Wells, and Clayton B. Winter for the camaraderie over the years. And lastly, a very special thank you to Chad D. Hoefler whose constant support and thoughtful advices helped me in the course of development and progress of this project and enabled me to persevere. vii ABSTRACT THE ECOLOGICAL STUDY OF THE MARITIME RINGLET BUTTERFLY (COENONYMPHA NIPISIQUIT MCDUNNOUGH) IN DALY POINT, BATHURST, NEW BRUNSWICK FEBRUARY 2006 MAKIRI SEI, B.S., UNIVERSITY OF TENNESSEE MARTIN Ph.D., UNIVERSITY OF MASSACHUSETTS AMHERST Directed by: Professor Adam H. Porter I studied the autecology, community ecology, and genetics of an endangered butterfly, the maritime ringlet (Coenonympha nipisiquit McDunnough), that inhabits a limited number of salt marshes in northern New Brunswick and in the Gaspé Peninsula of Québec. I studied the survival rate of first- and second-instar larvae in various microhabitats in a salt marsh at Daly Point Natural Reserve, Bathurst, New Brunswick. I found they survived significantly better in microhabitats dominated by Spartina patens (Aiton) Muhl. at an intermediate elevation. I investigated the tolerance of the maritime ringlet larvae to tidal submergence and compared their performance to a closely related taxon, the inornate ringlet (C. tullia inornata Edwards). The experiments revealed that the maritime ringlet possesses unique adaptations to tidal submergence. I examined the flight and oviposition behaviors of adult females in response to microhabitat. I found that they did not discriminate between microhabitats based upon the likelihood of larval survival as long as S. patens or other potential hosts were abundant. I explored the correlation between predator species richness and abundance with the larval survival rate in microhabitats. I found that predator abundance and species richness often responded viii negatively to increasing tidal flooding, suggesting that high larval mortality at high elevation sites can be caused by high predation pressure. Lastly, I investigated the possibility of genetic introgression between the maritime ringlet and inornate ringlet and reconstructed the phylogeny of the C. tullia-group taxa in North America. The genetic evidence did not support the possibility of large-scale genetic introgression and raised the taxonomic status of the maritime ringlet from a subspecies of holarctic C. tullia to a full species. The phylogenetic analyses suggested that the divergence of the maritime ringlet was much earlier than previously believed. My results will aid in protection and recovery of this endangered species. ix TABLE OF CONTENTS Page ACKNOWLEDGMENTS v ABSTRACT vii LIST OF TABLES xi LIST OF FIGURES xiv CHAPTER 1. INTRODUCTION 1 Description of the salt marsh in the Daly Point Natural Reserve 4 2. MICROHABITAT-SPECIFIC EARLY LARVAL SURVIVAL OF THE MARITIME RINGLET (COENONYMPHA NIPISIQUIT) 8 Introduction 8 Materials and Methods 10 Results 15 Discussion 17 3. LARVAL ADAPTATION OF THE MARITIME RINGLET TO A SALINE WETLAND HABITAT 26 Introduction 26 Materials and Methods 29 Results 32 Discussion 34 4. FLIGHT AND OVIPOSITION BEHAVIOR OF THE ADULT MARITIME RINGLET FEMALES IN RESPONSE TO MICROHABITAT 41 Introduction 41 Materials and Methods 43 Results 48 Discussion 50 5. BETWEEN THE DEVIL AND THE DEEP BLUE SEA: PREDATION AND FLOODING MAY DELIMIT LARVAL MICRODISTRIBUTION OF THE MARITIME RINGLET 69 [...]... adressing the underlying processes that control the population size By the time one sees the sign of decline in population size, it may be too late to rescue the population by investigating the cause of decline in order to amend the problem Thus, there is a need to study the maritime ringlet population biology in detail I chose a population of the maritime ringlet that occurs in a salt marsh within the Daly. .. SURVIVAL OF THE MARITIME RINGLET (COENONYMPHA NIPISIQUIT) Introduction Maritime ringlet butterflies (Coenonympha nipisiquit McDunnough, Satyridae) are a rare subspecies of the widespread holarctic common ringlet butterflies (Coenonympha tullia Müller) Found only in salt marshes in maritime Canada, maritime ringlets are subjected to periodic tidal submergence in their pre-adult stages, and they are one of the. .. need to identify the critical microhabitat requirements of this butterfly The natural history of the maritime ringlet differs in several ways from that of the inornate ringlet, Coenonympha tullia inornata Edwards, the sister North American subspecies that occurs in the adjacent uplands (Webster 1998) The occurrence of the maritime ringlet coincides with salt meadow cordgrass, Spartina patens (Aiton)... second instar in late September, and they resume growth and development in late May of the following year The allele frequency data in Wiernasz (1989) suggest that the maritime ringlet may be only distantly related to all of the inornate ringlet populations sampled from the Northeastern United States and Canada It is speculated that the maritime ringlet evolved from a small coastal population of the common... attention in Europe partly because of their low vagility (Joy & Pullin 1997; Lhonore & Lagarde 1999; Cassel et al 2001) With so few extant maritime ringlet populations in such a strongly delimited and vulnerable 1 habitat type, there is considerable need to study the ecology of this butterfly in depth The natural history of the maritime ringlet differs in several ways from that of the inornate ringlet, ... be the primary nectar source for the adult maritime ringlets and a good indicator of the maritime ringlet occurrence, although the adults can readily use other nectar sources The maritime ringlet is univoltine, and the adults fly from late July to mid August, a month later than common ringlet populations (Wiernasz 1989) that occur within a kilometer of the salt marshes Larvae enter diapause as the. .. reconstructed the phylogeny of the C tulliagroup taxa in North America to help guide conservation efforts concordant with evolutionary processes Description of The Salt Marsh in the Daly Point Natural Reserve The salt marsh is located in the Daly Point Natural Reserve in Bathurst, New Brunswick, Canada (Figure 1) The natural reserve was owned by Noranda Inc., a mining corporation, and leased to the Department of. .. test whether microhabitats chosen for oviposition overlaps with microhabitats suitable for the larvae In addition, I explored the cause of larval mortality by surveying predator communities across the salt marsh and by studying the effects of tidewater submergence of larvae in laboratory Lastly, I investigated the possibility of genetic introgression between the maritime ringlet and inornate ringlet. .. Primulaceae), and Virginia rose (Rosa virginiana Mill., Rosaceae) The maritime ringlet is univoltine, and the adults fly from late July to mid August, a month later than adults of the common ringlet (Wiernasz 1989) from populations within a kilometer of the salt marshes Larvae enter diapause during the second instar in late September, and they resume growth and development in late May of the following year (Webster... manipulation of the salt marshes, including the building of dykes and ditches, grazing, exploitation for salt hay, and burning, all of which used to be prevalent in the northern Atlantic coast (Roberts & Robertson 1986; Adam 1990) The general isolation of the salt marshes, as well as the philopatric nature of maritime ringlets, seems to preclude this subspecies from naturally colonizing other salt marshes . THE ECOLOGICAL STUDY OF THE MARITIME RINGLET BUTTERFLY (COENONYMPHA NIPISIQUIT MCDUNNOUGH) IN DALY POINT, BATHURST, NEW BRUNSWICK. A Dissertation. to study the ecology of this butterfly in depth. The natural history of the maritime ringlet differs in several ways from that of the inornate ringlet, Coenonympha tullia inornata Edwards, the. ProQuest Information and Learning Company. © Copyright by Makiri Sei 2006 All Rights Reserved THE ECOLOGICAL STUDY OF THE MARITIME RINGLET BUTTERFLY (COENONYMPHA NIPISIQUIT MCDUNNOUGH) IN DALY POINT,