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ENVIRONMENTAL ASSESSMENT OF CIRCUMNEUTRAL WETLANDS
WITH SHRUBBY CINQUEFOIL (DASIPHORA FRUTICOSA), HOST
PLANT OF THE ENDANGERD CLAYTON’S COPPER
BUTTERFLY (LYCAENA DORCAS CLAYTONI)
By
Sarah Ann Drahovzal
B.A. Wittenberg University, 1996
A THESIS
Submitted in Partial Fulfillment of the
Requirements for the Degree of
Master of Science
(in Ecology and Environmental Sciences)
The Graduate School
The University of Maine
May 2013
Advisory Committee:
Cynthia Loftin, Associate Professor of Wildlife Ecology and Unit Leader, U.S.
Geological Survey, Maine Cooperative Fish and Wildlife Research Unit,
Co-Advisor
Judith Rhymer, Associate Professor and Department Chair of Wildlife Ecology,
Co-Advisor
Francis Drummond, Professor of Insect Ecology and Entomology, School of
Biology and Ecology
Andrew Reeve, Professor, School of Earth and Climate Sciences
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THESIS ACCEPTANCE STATEMENT
On behalf of the Graduate Committee for Sarah Drahovzal I affirm that this
manuscript is the final and accepted thesis. Signatures of all committee members are on
file with the Graduate School at the University of Maine, 42 Stodder Hall, Orono, Maine.
Cynthia Loftin, Associate Professor of Wildlife Ecology and Unit Leader, U.S.
Geological Survey, Maine Cooperative Fish and Wildlife Research Unit (04/25/2013)
Judith Rhymer, Associate Professor and Department Chair of Wildlife Ecology
(04/25/2013)
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LIBRARY RIGHTS STATEMENT
In presenting this thesis in partial fulfillment of the requirements for an advanced
degree at the University of Maine, I agree that the Library shall make it freely available
for inspection. I further agree that permission for “fair use” copying of this thesis for
scholarly purposes may be granted by the Librarian. It is understood that any copying or
publication of this thesis for financial gain shall not be allowed without my written
permission.
Signature:
Date:
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ENVIRONMENTAL ASSESSMENT OF CIRCUMNEUTRAL WETLANDS
WITH SHRUBBY CINQUEFOIL (DASIPHORA FRUTICOSA), HOST
PLANT OF THE ENDANGERD CLAYTON’S COPPER
BUTTERFLY (LYCAENA DORCAS CLAYTONI)
By Sarah Ann Drahovzal
Thesis Co-Advisors: Dr. Cynthia Loftin
Dr. Judith Rhymer
An Abstract of the Thesis Presented
in Partial Fulfillment of the Requirements for the
Degree of Master of Science
(in Ecology and Environmental Sciences)
May 2013
Clayton’s copper butterfly (Lycaena dorcas claytoni) is a Maine state endangered
species that relies exclusively on shrubby cinquefoil (Dasiphora fruticosa) as its host
plant. This shrub typically is found on the edges of wetlands rich in calcium carbonate or
limestone. Calcareous wetland habitats that support large, persistent stands of D.
fruticosa are rare in Maine (McCollough et al. 2001). Currently only 21 sites in Maine
are known to support large stands of D. fruticosa, and L. d. claytoni populations have
been observed at only nine of these. Because nearly the entire global range of this
butterfly is in Maine, it is critical that Maine assumes the primary role in the conservation
of this rare subspecies.
Conservation of L. d. claytoni depends in part on the ecological integrity of its
habitat. Vegetation structure and hydrological conditions in wetlands may affect the
distribution and robustness of D. fruticosa, which also may influence its use as a host
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plant by L. d. claytoni. I conducted field studies in 2009 and 2010 in ten wetlands in
Maine with robust stands of D. fruticosa to evaluate pore water nutrients, hydrological
conditions, shrub and tree species composition and distribution, and D. fruticosa
distribution, structure, age and condition; seven of these wetlands support populations of
L. d. claytoni, and three of these wetlands are unoccupied. I identified five hydrological
types based on differences in water source and surface and ground water dynamics. Three
wetlands were dominated by groundwater discharge, six wetlands were down-flow
dominant, and one wetland fluctuated between groundwater discharge and recharge. Pore
water analytes reflected hydrogen ion and conductivity gradients among the wetlands and
vegetation community distributions within the wetlands, however, these differences did
not reflect wetland occupation of L. d. claytoni. Dasiphora fruticosa age ranged from 7 to
37 years. Previously reported Lycaena dorcas claytoni encounter rates were greater in
wetlands containing larger D. fruticosa plants of intermediate age and with greater bloom
density. Butterflies are able to differentiate among glucose, fructose and sucrose in
nectar. I found D. fruticosa produces hexose dominant nectar (sucrose/[glucose +
fructose] < 0.1), with only trace amounts of sucrose measured in < 3% of the samples.
Conservation and recovery of L. d. claytoni depends in part on the quality and
distribution of its habitat. Although Maine’s wetlands hosting L. d. claytoni currently
support robust stands of D. fruticosa, their isolation likely limits movement of the
butterfly. Increased connectivity among wetlands containing shrubby cinquefoil may aid
dispersal and improve likelihood of long-term L. d. claytoni population.
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ACKNOWLEDGEMENTS
Funding for this project was provided by the Maine Agricultural and Forest
Experiment Station (MAFES), U.S. Geological Survey, Maine Cooperative Fish and
Wildlife Research Unit, Maine Department of Inland Fisheries and Wildlife (MDIFW),
Maine Outdoor Heritage Fund, The Nature Conservancy (TNC), U.S. Fish and Wildlife
Service (USFWS), and the University of Maine Wildlife Ecology Department. Among
these supporters, I am especially grateful to Beth Swartz (MDIFW), Nancy Sferra (TNC),
and Mark McCollough (USFWS).
Many thanks to my committee for their help throughout this process. Cyndy
Loftin provided a constant source of academic and professional guidance and was a
positive and supportive mentor. Judith Rhymer helped provide many ideas from which
much of this research was based and lent a critical eye and thorough critique of all
products. Andy Reeve and Frank Drummond provided invaluable advice and made this
project possible though their ideas and guidance.
I thank Bill Halteman for the time he spent patiently helping me with statistics;
Mike Day for invaluable advice on annual ring analysis as well as use of his lab; L. Brian
Perkins for the many hours he spent helping me analyze nectar sample and the use of his
lab and HPLC machine; Brad Libby for use of greenhouse space; Dennis Anderson for
advice and supplies for water samples; Lyndsy Shuman for formatting advice, and
George Jacobson and Molly Schauffler for advice regarding vegetation sampling.
I was lucky enough to work with two incredibly intelligent and scrappy field
technicians, Amanda Theriault Fessenden and Katie Chenard, who kept me laughing,
well-fed and in generally good spirits though long days, black fly swarms, flat tires and
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swampy hip waders. I am also indebted to the others who provided (often last-minute)
help in the field: Monika Parsons, Sheryn Olson, Brock Sanborn, Cyndy Loftin, Cory
Michaud, Dawn Morgan, Ian McCullough, Erik Belmer, Dave Ellis, Lindsay Seward, and
Erin Porter. I am very grateful to Josie and Dave Allen at Macannamac Camp Rentals
who generously provided lodging in the Great North Woods.
My time at the university has been much more enjoyable because of the fantastic
graduate students, faculty, and staff in the Wildlife Ecology Department. Everyone in the
department has at one time or another provided academic, professional or moral support.
Rena Carey, Julie Eubanks, Katherine Goodine, and Julie Nowell have provided
administrative help as well as much needed (and well timed) chocolate and coffee breaks.
Nutting room 220 has been my family here in Maine: Kevin Ryan (work spouse), Monika
Parsons, Cory Michaud, Dave Ellis, Vanessa Levesque, Sheryn Olson, Dan Stich, and
Silas Ratten. Special thanks also to Britt Cline, Margarette Guyette, Erynn Call and
Lindsay Seward who have acted as mentors, friends and therapists.
Finally, I am especially grateful to my family and friends (you know who you
are!), who have put up with me through everything. I particularly want to thank my
parents, Jim and Becky Drahovzal, who have been a constant source of strength, support
and inspiration.
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TABLE OF CONTENTS
ACKNOWLEDGEMENTS……………………………………………………………... iii
LIST OF TABLES………………………………………………………………………viii
LIST OF FIGURES……………………………………………………………………... xi
CHAPTER
1. ANALYSIS OF THE HYDROLOGICAL AND CHEMICAL ENVIRONMENT
OF WETLANDS WITH SHRUBBY CINQUEFOIL (DASIPHORA
FRUTICOSA) ......................................................................................................................1
1.1. Introduction .............................................................................................................1
1.2. Methods ...................................................................................................................3
1.2.1. Study species and area ..................................................................................3
1.2.2. Data collection ..............................................................................................6
1.2.2.1. Hydrologic measurements ...................................................................... 6
1.2.2.2. Pore water characterization ..................................................................... 6
1.2.2.3. Peat characterization ............................................................................... 7
1.2.2.4. Woody species characterization .............................................................. 8
1.2.3. Data analysis .................................................................................................8
1.2.3.1. Hydrology ............................................................................................... 8
1.2.3.2. Wetland vegetation structure and chemistry ........................................... 9
1.3. Results .................................................................................................................100
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1.3.1. Wetland hydrological environments .........................................................100
1.3.1.1. Hydrological conditions during L. d. claytoni and D. fruticosa
life history stages ................................................................................ 111
1.3.2. Pore water characterization .........................................................................20
1.3.3. Peat characterization ...................................................................................27
1.3.4. Wetland woody species characterization ....................................................30
1.4. Discussion .............................................................................................................36
1.4.1. Hydrological environments of D. fruticosa ................................................36
1.4.2. Chemical environments of D. fruticosa and relevance to L. d. claytoni .....37
1.4.3. Management implications ...........................................................................41
2. ASSESSMENT OF SHRUBBY CINQUEFOIL (DASIPHORA FRUTICOSA)
AS THE HOST PLANT FOR CLAYTON’S COPPER BUTTERFLY (LYCAENA
DORCAS CLAYTONI) .......................................................................................................42
2.1. Introduction ...........................................................................................................42
2.2. Methods .................................................................................................................45
2.2.1. Study area ....................................................................................................45
2.2.2. Foliar moisture and nitrogen .......................................................................46
2.2.3. Bloom surveys and nectar sampling ...........................................................47
2.2.4. Age structure ...............................................................................................47
2.3. Results ...................................................................................................................49
2.3.1. Foliar nitrogen and moisture .......................................................................49
2.3.2. Nectar sugar composition and bloom surveys ............................................52
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2.3.3. Age structure ...............................................................................................54
2.4. Discussion .............................................................................................................67
2.4.1. Resources for larval L. d. claytoni ..............................................................67
2.4.2. Resources of adult L. d. claytoni .................................................................68
2.4.3. Conservation Implications ..........................................................................72
REFERENCES ..................................................................................................................74
APPENDIX A. SITE MAPS..............................................................................................82
APPENDIX B. HYDROGRAPHS ....................................................................................94
APPENDIX C. COMPONENT LOADINGS ..................................................................112
APPENDIX D. PORE WATER AND PEAT ANALYTES ............................................115
APPENDIX E. SITE MAPS WITH DASIPHORA FRUTICOSA SHRUB
VOLUME ........................................................................................................................117
APPENDIX F. VEGETATION MEASUREMENTS ....................................................124
BIOGRAPHY OF THE AUTHOR ..................................................................................126
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LIST OF TABLES
Table 1.1. Wetland hydrological type, well depth (m), and distance between
screens for deep and shallow wells. ................................................................ 12
Table 1.2. Upwelling (%), mean (± standard deviation, SD) difference (cm)
between water table and deep well, saturation (%), depth (cm) to
water table (mean ±SD), and flood duration (%) for 10 wetlands
monitored June – September 2009. ................................................................. 13
Table 1.3. Upwelling (%), mean (±SD) difference (cm) between water table
and deep well, saturation (%), mean (±SD) depth (cm) to water table,
and flood duration (%) for 10 wetlands monitored June 2009 –
September 2009 in Maine. .............................................................................. 15
Table 1.4. Upwelling (%), mean (± standard deviation, SD) difference (cm)
between water table and deep well, saturation (%), depth (cm) to
water table (mean ±SD), and flood duration (%) for 10 wetlands
monitored May– September 2010 ................................................................... 16
Table 1.5. Upwelling (%), mean (±SD) difference (cm) between water table
and deep well, saturation (%), mean (±SD) depth (cm) to water table,
and flood duration (%) for 10 wetland monitored May 2010 –
September 2010 in Maine. ............................................................................. 18
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Table 1.6. Multiple-response permutation procedure results of water from pore
water samples collected from wetlands in Maine during larval
emergence (24 May -3 June 2010). ................................................................ 24
Table 1.7. Multiple-response permutation procedure results of water analytes from
pore water samples collected from wetlands in Maine during
L. d. claytoni nectaring (12 – 23 July 2010). .................................................. 25
Table 1.8. Multiple-response permutation procedure results of water analytes
from pore water samples collected from wetlands in Maine during
D. fruticosa senescence (17 – 25 August 2010). ............................................ 26
Table 1.9. Multiple-response permutation procedure results of peat analytes
from 10 wetland in northern Maine, 2010. ..................................................... 29
Table 1.10. Multiple-response permutation procedure results of shrub community
from 10 wetlands in mid-state, northwestern and northeastern
Maine, 2010. .................................................................................................. 33
Table 2.1. Pearson correlation coefficients (r) for foliar nitrogen (FN) and
foliar moisture (FM) for 10 wetlands in Maine, USA. ................................... 52
Table 2.2. Pearson correlation coefficients (r) for age vs. length, aboveground
biomass (AGB), and stem diameter of D. fruticosa collected from
10 wetlands in Maine. ..................................................................................... 65
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Table 2.3. Pearson correlation coefficients (r) were calculated for age vs. height
and aboveground biomass (AGB) for upright and prostrate growth
forms for D. fruticosa collected from 10 wetlands in Maine. ......................... 65
Table 2.4. Pearson correlation coefficients (r) for age vs. height and age vs. above
ground biomass (AGB) of D. fruticosa collected from three zones
(water edge, non-forested wetland, and forested) in 10 wetlands
in Maine. ......................................................................................................... 66
Table 2.5. Comparison of growth rates among 10 wetlands............................................. 66
Table C.1. Component loadings of each analyte on the first two principal
components from pore water data collected from 10 wetlands . .................. 112
Table C.2. Component loadings of each analyte on the first three principal
components from peat data collected from 10 wetlands. .............................. 113
Table C.3. Component loadings of shrub species on the first three principal
components from peat data collected from 10 wetlands. .............................. 114
Table D.1. Average pore water analytes for 10 wetlands in mid-state, northwestern
and northeastern Maine, 2010. ...................................................................... 115
Table D.2. Average peat analytes for 10 wetlands in mid-state, northwestern and
northeastern Maine, 2010.............................................................................. 116
Table F.1. Abbreviation (Abbrev.) of shrub and tree species names .............................. 124
Table F.2. Average shrub volumes (m3) for 10 wetlands ............................................... 125
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Table F.3. Basal Area (cm2/transect m) of tree species for 10 wetlands. ....................... 125
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LIST OF FIGURES
Figure 1.1. Locations of wetlands included in our study of shrubby cinquefoil in
Maine ............................................................................................................... 5
Figure 1.2. Running average of inundation and timing of Clayton’s copper (CC)
life history stages during the 2009 and 2010 growing seasons ..................... 19
Figure 1.3. Principal component biplots of water analytes from pore water
samples collected from wetlands in Maine during larval emergence
(24 May -3 June 2010). ..................................................................................................... 21
Figure 1.4. Principal component biplots of water analytes from pore water
samples collected from wetlands in Maine during L. d. claytoni
nectaring (12 – 23 July 2010). ....................................................................... 22
Figure 1.5. Principal component biplots of water analytes from pore water
samples collected from wetlands in Maine during D. fruticosa
senescence (17 – 25 August 2010). .............................................................. 23
Figure 1.6. Principal component biplots of peat analytes at the 10 wetlands. .................. 28
Figure 1.7. Average Dasiphora fruticosa volume (m3). .................................................. 31
Figure 1.8. Shrub coverage (m3/transect m) totaled along transects in 10 wetlands. ....... 32
Figure 1.9. Tree basal area (cm2/ transect m). .................................................................. 35
Figure 2.1. Average foliar moisture (%) for 10 wetlands. ................................................ 50
Figure 2.2. Average foliar nitrogen (%) for 10 wetlands. ................................................. 51
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Figure 2.3. Average concentration (mg/bloom) of fructose, glucose and sucrose
for un-bagged (solid bars) and bagged blooms (hatched bars). ..................... 53
Figure 2.4. Total bloom density (number of blooms/transect m) at 7 wetlands
with L. d. claytoni (solid bars) and 3 wetlands with D. fruticosa but
without L. d. claytoni (hatched bars). ............................................................ 53
Figure 2.5. Box plot of shrub age. .................................................................................... 55
Figure 2.6. Average annual growth rate (mm/year) for 10 wetlands.. .............................. 56
Figure 2.7. Growth curve rates by age (a) and year (b) for all wetlands (n=145)
derived from average stem cross-sectional increment. .................................. 57
Figure 2.8. Growth rates by age (a) and year (b) of shrubs collected from Holt
(n=17), Dwinal (n=29), Pickle (n=11) and Mattagodus (n= 12)
average stem cross-sectional increment. ....................................................... 58
Figure 2.9. Growth curve by age (a) and year (b) for Salmon (n=17) and Crystal
(n=8) derived from average stem cross-sectional increment. ........................ 59
Figure 2.10. Growth curve by age (a) and year (b) for Pillsbury (n=14) and Soper
(n=12) from average stem cross-sectional increment. ................................... 60
Figure 2.11. Growth curve by age (a) and year (b) for Woodland (n=11) and
Portage (n=12) derived from average stem cross-sectional increment. ........ 61
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Figure 2.12. Growth curve by age (a) and year (b) for water edge (n=32),
non-forested (n=60), and forested (n=31) zones derived from average
stem cross-sectional increment. ..................................................................... 62
Figure 2.13. Age structure of Dasiphora fruticosa in ten wetlands ordered from
south to north. ................................................................................................ 63
Figure 2.14. Number of side branches produced by the main stem in each age class
of Dasiphora fruticosa in ten wetlands ordered from south to north.. .......... 64
Figure 2.15. L. d. claytoni nectaring on Solidago uliginosa (Family Asteraceae) ........... 70
Figure A.1. Transect layout at Upper Holt pond, Maine, USA. ....................................... 82
Figure A.2. Transect layout at Lower Holt Pond Maine, USA. ....................................... 83
Figure A.3. Transect layout at Upper Dwinal WMA Maine, USA. ................................ 84
Figure A.4. Transect layout at Lower Dwinal WMA, Maine, USA. ................................ 85
Figure A.5. Transect layout at Pickle Ridge, Maine, USA. .............................................. 86
Figure A.6. Transect layout at Mattagodus WMA, Maine, USA ..................................... 87
Figure A.7. Transect layout at Salmon Stream, Maine, USA. .......................................... 88
Figure A.8. Transect layout at Crystal Fen, Maine, USA. ................................................ 89
Figure A.9. Transect layout at Pillsbury Pond, Maine, USA. ........................................... 90
Figure A.10. Transect layout at Soper Pond, Maine, USA. .............................................. 91
Figure A.11. Transect layout at Woodland Bog, Maine, USA. ........................................ 92
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Figure A.12. Transect layout at Portage Lake, Maine, USA. ........................................... 93
Figure B.1. Hydrograph of water-table well and deep well for Holt, May 2010 –
September 2010. ............................................................................................ 94
Figure B.2. Hydrograph of water-table well and deep well for Dwinal, monitored
June 2009 – September 2009 and May 2010 – September 2010. ................. 95
Figure B.3. Hydrograph of water-table well and deep well for Pickle monitered
May 2010 – September 2010. ........................................................................ 97
Figure B.4. Hydrograph of water-table well and deep well for Mattagodus,
monitored June 2009 – September 2009 and May 2010 –
September 2010. ............................................................................................ 98
Figure B.5. Hydrograph of water-table well and deep well for Crystal, monitored
June 2009 – September 2009 and May 2010 – September 2010. ............... 100
Figure B.6. Hydrograph of water-table well and deep well for Salmon, monitored
June 2009 – September 2009 and May 2010 – September 2010. ............... 102
Figure B.7. Hydrograph of water-table well and deep well for Pillsbury, monitored
June 2009 – September 2009 and May 2010 – September 2010. ............... 104
Figure B.8. Hydrograph of water-table well and deep well for Soper, monitored
June 2009 – September 2009 and May 2010 – September 2010. ............... 106
Figure B.9. Hydrograph of water-table well and deep well for Portage, monitored
June 2009 – September 2009 and May 2010 – September 2010. ............... 108
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Figure B.10. Hydrograph of water-table well and deep well for Woodland,
monitored June 2009 – September 2009 and May 2010 –
September 2010. .......................................................................................... 110
Figure E.1. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transecta at (a) upper and (b) lower Holt Pond, Maine, USA. .......... 117
Figure E.2. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at (a) upper and (b) lower Dwinal, Maine, USA. ............... 118
Figure E.3. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at (a) Pickle Ridge and (b) Mattagodus, Maine, USA. ....... 119
Figure E.4. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at (a) Salmon Stream and (b) Crystal Fen,
Maine, USA. ................................................................................................ 120
Figure E.5. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at Pillsbury Pond Crystal Fen, Maine, USA. ...................... 121
Figure E.6. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at Holt Pond, Maine, USA. ................................................. 122
Figure E.7. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals
along transects at (a) Woodland Bog and (b) Portage Lake, Maine, USA
..................................................................................................................... 123
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CHAPTER 1
ANALYSIS OF THE HYDROLOGICAL AND CHEMICAL ENVIRONMENT OF
WETLANDS WITH SHRUBBY CINQUEFOIL (DASIPHORA FRUTICOSA)
1.1. Introduction
Habitat loss, fragmentation and degradation are primary causes for decline and
extinction of many insect species (Fahrig 2003). Habitat patch size and connectivity may
affect patch occupancy patterns of species that occur in fragmented habitats. However,
habitat patch quality may be as important as habitat loss and fragmentation for
determining patch occupancy (Thomas et al. 2001).
Insect species that are monophagus, habitat specific, and with limited dispersal
ability and short flight periods may have increased vulnerability to extinction, particularly
when habitats are altered or isolated (Hughes et al. 2000, Kotze et al 2003, Kotiaho et al.
2005). Host plant density is a key determinate of population size for monophagus,
habitat-specialized butterflies (Leon-Cortes et al. 2003, Krauss et al. 2005), because their
distribution is limited by the occurrence of their host plant. Knowledge of host plant
habitat requirements is essential for effective conservation of critical habitat for host-
specialist species.
Clayton’s copper butterfly (Lycaena dorcas claytoni) is a Maine state endangered
species that relies exclusively on shrubby cinquefoil (Dasiphora fruticosa) as its host
plant. Lycaena dorcas claytoni was listed as endangered in 1997 under Maine’s
Endangered Species Act (12 MRSA, Section 7753). This subspecies of Dorcus copper
(Lycaena dorcas) has been observed in five wetlands in New Brunswick, Canada, and 10
wetlands in Maine (Webster and Swartz, 2006). A recent extinction of an L. d. claytoni
population at a historically occupied wetland (Crystal Fen) lends urgency to an
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environmental assessment of D. fruticosa habitat in Maine. Although D. fruticosa is not
considered rare, there are few wetlands habitats in Maine that support large, persistent
stands of D. fruticosa (McCollough et al. 2001), and the environmental conditions that
support robust stands of D. fruticosa are not well-defined.
Hydrology is a primary driver of wetland plant community development, structure
and persistence. Water depth, duration and flood frequency, as well as water source
directly affect nutrient availability and peat and pore water pH (Mitsch and Gosselink
2007). Wetlands supporting D. fruticosa in Maine are classified as circumneutral fens and
streamside shrublands and meadows (McCollough et al. 2001). Fens are groundwater-fed
systems (Bedford and Godwin 2003), and the plant rooting zone is supplied with mineral
nutrients from groundwater that has been in contact with mineral substrate (Wassan et al.
1990). Mineral rich groundwater has a near-neutral pH, which leads to faster decay of
organic material than in more acidic conditions. Consequently, greater calcium
concentrations in ground water results in the greater potential release of nutrients through
organic decay, which are then available for uptake by wetland plants. Most fens are
relatively nutrient-poor, however, and vegetation community establishment is affected by
nutrient availability (Keddy 2000). Although peatland nutrient gradients may be tightly
correlated with acidity-alkalinity gradients (Bridgham et al. 1996), hydrology and pH
may have greater influence than nutrient availability on vegetation community
establishment (Vitt 1990; Vitt et al. 1995; Bedford et al. 1999; Bragazza and Gerdol
2002).
Variations in hydrological and chemical gradients in wetlands affect vegetation
establishment and persistence (Mitsch and Gosselink 2007). Knowledge of environmental
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site characteristics that support robust stands of D. fruticosa, such as hydrological
conditions and nutrient availability, is critical for conservation of L. d. claytoni.
Dasiphora fruticosa occurs across 28 wetlands in Maine. Environmental conditions that
affect the range of growth forms, robustness and distribution of D. fruticosa in Maine are
not well understood. The objective of this study was to compare the hydrological
environment, pore water and peat chemistry, and plant community composition of
Maine’s wetlands containing D. fruticosa, and examine conditions in wetlands inhabited
and uninhabited by L. d. claytoni.
1.2. Methods
1.2.1. Study species and area
The nominate species, Dorcus copper, produces one brood per year and deposits
single eggs in August on the underside of cinquefoil leaves near the top of smaller plants
(Scott 1986). Much of what is known of the life history of L. d. claytoni is reported by
McCollough et al. (2001). Eggs drop with the leaves in the autumn and overwinter on the
ground. Clayton’s copper larvae emerge in the spring to feed on cinquefoil leaves,
completing five instars from larva to pupa. Adults nectar on D. fruticosa during late July
through August when shrubby cinquefoil is flowering, and they remain close to
cinquefoil stands throughout the flight season. Emergence and flowering may occur
earlier in hotter, drier years. Conversely, flowering, emergence and egg-laying may
continue into mid-September in cooler years.
Clayton’s copper butterflies occupy wetlands in central (Dwinal Pond Wildlife
Management Area, Mattagodus Meadows, Holt Pond, Pickle Ridge), western (Pillsbury
Pond, Soper Pond, Little Round Pond), and northern (Woodland Bog and nearby
wetland) Maine (Fig.1.1). Moderate numbers of the butterfly were observed at Dwinal,
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Mattagodus, Holt, Pillsbury and Soper in 2007-2008, whereas, few butterflies were
observed during this period at the other occupied wetlands in Maine (Knurek 2010).
Clayton’s copper butterflies have not been observed at Crystal Fen since 2008 (Drahovzal
personal observation; Knurek, 2010), and the wetland is now considered unoccupied.
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Figure 1.1. Locations of wetlands included in this study of shrubby cinquefoil in Maine.
Area (ha)
Holt Pond 2.9
Dwinal ~20.0
Pickle Ridge 0.7
Mattagodus 7.5
Salmon Stream 1.6
Crystal Fen 2.0
Pillsbury Pond 6.3
Soper Pond 4.6
Woodland Bog 0.8
Unnamed Bog ~1.2
Portage Lake ~0.6
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1.2.2. Data collection
1.2.2.1. Hydrologic measurements
I installed hydrologic monitoring wells equipped with continuous water level
recorders in five wetlands hosting L. d. claytoni (Dwinal, Mattagodus, Pillsbury, Soper,
Woodland) and three wetlands with D. fruticosa but without the butterfly (Salmon
Stream, Portage Lake, Crystal Bog) during May - October 2009. I redeployed the
recorders in these wetlands and Pickle Ridge and Holt Pond during May - October 2010.
I inserted two wells (2.5 cm diameter polyvinyl chloride tubes slotted at the lower end)
vertically into the peat, extending from the peat surface and open to the atmosphere. I
placed one well to monitor the water table position and measure the vertical hydraulic
gradient in the near surface sediments and one well at the peat-mineral substrate interface
to monitor groundwater upwelling. Non-vented data logging pressure transducers (Solinst
Model 3001 level logger, Georgetown, Ontario, Canada) were programmed to record
hydraulic head every 30 minutes, and water depths were periodically measured by hand
to assess accuracy of automated measurements. I placed a Model 3001 Barologger
(Solinst, Georgetown, Ontario, Canada) recording every 30 minutes at Dwinal, Salmon,
Pillsbury, and Woodland to correct water level measurements for barometric pressure in
Mattagodous, Pickle and Holt. I used barometric pressure data from Salmon, Pillsbury
and Woodland to correct level logger data from Crystal, Soper, and Portage. Water depth
measurement error was ±0.1cm per meter of water depth above the logger.
1.2.2.2. Pore water characterization
I collected pore water samples during 24 May - 3 June (n=102), 12 - 23 July
(n=102), and 17 – 25 August (n=65) 2010 along transects sampled by Knurek (2010) for
assessment of the taxonomic and population status of Clayton’s copper butterfly, and
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along additional transects I established in the unoccupied wetlands (Appendix A).
Transects traversed the most dense stands of D. fruticosa and captured each wetland’s
vegetation and topographic variation, which I delineated in zones (near water sedge,
shrub-sedge interior, forested wetland; Appendix A). Samples were collected at the base
of randomly selected plants in each transect’s vegetation zones. I collected pore water
with a hand vacuum pump attached to flexible plastic tube placed into a slotted 2.5cm
diameter PVC pipe inserted 10 - 15 cm below the peat surface in the D. fruticosa root
zone. I stored water samples at 4º C in 120 cc glass amber bottles and filtered them
within two hours of collection. Samples were analyzed for NH4-N, NO3-N, phosphorus
(PO4-P), pH and organic conductivity by Maine Soil Testing Service Laboratory at the
University of Maine (AMSTS). NH4/NO3-N in water samples was determined
colorimetrically by an autoanalyzer. The cadmium reduction/sulfanilamide method was
used for NO3-N, and the hypochlorite/salicilate method was used for NH4-N. Phosphorus
was determined with the automated ascorbic acid reduction method (APHA 1995). Water
pH was determined potentiometrically with an electronic pH meter. Conductivity was
measured with a Fisher Scientific Digital Conductivity Meter.
1.2.2.3. Peat characterization
I collected peat samples within the D. fruticosa root zone (up to 30 cm below the
ground surface) in August 2009 (n=48) and May 2010 (n=102) from the base of
randomly selected shrubs where I collected pore water samples. Peat samples were stored
in cardboard sample boxes at 4°C until analyzed by AMSTS. Samples were air dried,
ground, and passed through a 2 mm sieve. Peat pH was determined with a pH meter
inserted in a slurry of peat and deionized water (McLean, 1982). Calcium (Ca), potassium
(K), magnesium(Mg), phosphorus(P), aluminum (Al) , boron (B), copper (Cu), iron (Fe),
Page 27
8
manganese (Mn), sodium (Na), lead (Pb), sulfur (S), and zinc (Zn) were analyzed by
Inductively Coupled Plasma (ICP) emission spectrophotometry after acidifying the peat
samples with pH 4.8 ammonium acetate (Modified Morgan) solution. Extractable
ammonium (NH4-N) and nitrate (NO3-N) in peat were determined in a 1 M KCl extract.
Solution analysis was done colorimetrically by an autoanalyzer. The cadmium
reduction/sulfanilamide method was used for NO3-N, and the hypochlorite/salicilate
method for NH4-N. Organic content of oven dried samples was determined by measuring
loss on ignition (LOI) after combustion at 375º C for two hours.
1.2.2.4. Woody species characterization
During May – August 2010 I recorded the species and intercept length, height,
width and shape (cone, cube, sphere) of each shrub intersecting a line 5 m to the left of
the centerline of the established transects (Mitchell and Hughes 1995). I recorded tree
species and diameter at breast height of all trees > 1m height in belt transects spanning 20
m of the transect centerline.
1.2.3. Data analysis
1.2.3.1. Hydrology
I constructed hydrographs for each study wetland and determined mean difference
between the water table and deep well depths, mean depth to water table, groundwater
upwelling (proportion of water level measurements with upwelling; %), saturation
(proportion of water depth measurements in the rooting zone; %), inundation (proportion
of water depth measurements above the peat surface; %), and flood duration (days, %) for
each wetland and interval [sampling season, life stages (D. fruticosa leaf out, bloom, and
senescence periods; L. d. claytoni egg, larval, and flight periods)]. I interpreted a
downward water flow direction if the water level in the shallow well was higher than that
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in the deep well, a neutral flow direction if the difference between water levels in the
shallow and deep wells was ≤ 0.1 cm, and an upward flow if the water level in the deep
well exceeded that in the shallow well. I calculated a running average of wetland
inundation depth for each wetland and season to describe daily fluctuations in water
levels. I related hydrological patterns to daily precipitation data for the 2009 and 2010
field seasons available from National Weather Service (NWS) stations located in
Millinocket, Clayton Lake, and Caribou, Maine. Data were provided by National Ocean
and Atmospheric Administration Satellite and Information Service from their Web site at
http://www7.ncdc.noaa.gov/CDO/cdoselect.cmd.
1.2.3.2. Wetland vegetation structure and chemistry
I estimated shrub volume and percent linear coverage from height and width
measurements and shape data for each shrub by species intercepting a transect. Shrub
linear coverage was compiled across transects in each wetland. Peat and pore water pH
were converted to H+
ion concentration prior to analysis. I compared pore water, peat
analytes and vegetation volume among wetlands with principal components analysis
(PCA; SYSTAT 12; SYSTAT Soft Inc, Chicago, IL) and scatter plots of the principal
component scores. I compared pore water and peat analytes as well as shrub community
structure among and within wetlands with multi-response permutation procedure (MRPP,
McCune and Grace 2002), with n/sum (n) as the weighting factor and relative Sorensen
distance measure. I compared average volume (m3) of shrubby cinquefoil among
wetlands with Wilcoxon rank sum tests.
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1.3. Results
1.3.1. Wetland hydrological environments
Precipitation did not differ between 2009 (µ=0.37 cm/day, SE=0.07) and 2010
(µ=0.30 cm/day, SE=0.06), however, there were more flooding events during the 2009
growing season (Tables 1.1, 1.2 and 1.3) and more drawdown and variability in water
table levels during the 2010 growing season (Tables 1.4 and 1.5).
I identified five hydrological types based on differences among the study
wetlands in water source and surface and ground water dynamics (Tables 1.1, 1.3 and
1.5; Appendix B). Hydrological type 1 (HT1) includes Woodland, Crystal, and Holt,
which were dominated by groundwater discharge and relatively constant water levels
with little to no inundation. Hydrological type 2 (HT2) includes Mattagodus, Pillsbury
and Soper, which had relatively constant water levels throughout the growing season with
little to no inundation and minimal drawdown below the root-zone at the end of the
season. Hydrological type 3 (HT3) includes Dwinal and Salmon, which were dominated
by surface water inputs and declining water levels during the 2010 growing season.
Inundation or saturation occurred at the beginning of the season; however, the water table
dramatically lowered by the end of the 2010 growing season. Hydrological type 4 (HT4)
includes Pickle, which was dominated by surface water inputs and relatively dynamic
water levels with inundation at the beginning of the season. Although drawdown
occurred at the end of the 2010 season, water levels did not drop below the rooting zone.
Hydrological type 5 (HT5) includes Portage, which fluctuated between groundwater
discharge and groundwater recharge and an inundated and drained root zone several
times during the 2010 growing season.
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1.3.1.1. Hydrological conditions during L. d. claytoni and D. fruticosa life history
stages
Timing of life history stages of L. d. claytoni and D. fruticosa differed between
2009 and 2010. The magnitude and duration of flooding was greatest during the larval
feeding period (Fig.1.2). Portage was an exception during 2009, with the greatest
magnitude and duration of flooding occurring during the D. fruticosa bloom-L. d.
claytoni nectaring period. Mean depth to water table and upwelling generally increased as
drawdown occurred during shrubby cinquefoil senescence (Tables 1.3 and 1.5).
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Table 1.1. Wetland hydrological type, well depth (m), and distance between screens for
deep and shallow wells.
Depth (m)b
Site HTa
shallow deep Distance (cm)c
Holt Pond 1 0.52 3.37 2.54
Dwinal 3 0.97 2.17 0.89
Pickle Ridge 4 1.18 3.94 2.45
Mattagodus 2 0.52 3.60 2.77
Salmon Stream 3 0.62 2.50 1.52
Crystal Fen 1 0.54 0.76 0.00
Pillsbury Pond 2 0.61 1.89 0.97
Soper Pond 2 0.67 2.13 1.15
Woodland Bog 1 0.64 1.86 0.91
Portage Lake 5 0.67 2.61 1.63
a
Hydrological type (HT1= groundwater discharge, constant water levels,
little to no inundation; HT2 = surface water input dominant, constant
water levels, little to no inundation; HT3= surface water input dominant,
inundation at beginning of season, drawdown below rooting zone at end of
season; HT4= surface water input dominant, inundation at beginning of
season, no drawdown below rooting zone; HT5= fluctuated between
groundwater discharge and surface water inputs, inundation and drained
root zone several times during season). b
Depth of well below peat surface. c Distance between shallow and deep well screens.
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Table 1.2. Upwelling (%), mean (± standard deviation, SD) difference (cm) between
water table and deep well, saturation (%), depth (cm) to water table (mean ±SD), and
flood duration (%) for 10 wetlands monitored June– September 2009. Data are reported
for Clayton’s copper (CC) larval feeding (20 June-22 July), shrubby cinquefoil (SC)
bloom and Clayton’s Copper flight (23 July – 19 August), and shrubby cinquefoil
senescence periods in Maine. ND= No Data.
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14
Upwellinga
Mean difference Saturationb
Mean Depth to
Water Table
Flood
durationc
(cm) (±SD) (%) (cm) (±SD) (%)
CC larval feeding
Holt Pond ND ND ND ND ND ND ND
Dwinal 0 5.8 0.83 100 -3.46 1.98 7.3
Pickle Ridge ND ND ND ND ND ND ND
Mattagodus 0 3.7 2.10 100 -6.30 1.16 0
Salmon Stream 5.3 5.4 3.04 100 -1.40 1.16 11
Crystal Fen 100 -6.4 0.29 100 -16.89 0.92 0
Pillsbury Pond ND ND ND ND ND ND ND
Soper Pond 0 18.87 2.84 100 -2.63 1.57 5.9
Woodland Bog 95.13 -0.91 0.38 100 -13.00 2.36 0
Portage Lake 31.8 3.45 4.14 100 -2.30 4.56 36.6
SC bloom/ CC adult nectaring
Holt Pond ND ND ND ND ND ND ND
Dwinal 0 5.9 0.52 100 -6.55 4.10 4.6
Pickle Ridge ND ND ND ND ND ND ND
Mattagodus 0 6.3 1.10 100 -6.75 2.15 0
Salmon Stream 0 5.1 1.21 100 -7.07 1.85 0
Crystal Fen 100 6.6 0.20 100 -15.35 1.49 0
Pillsbury Pond 0 7.0 5.39 100 ND ND ND
Soper Pond 0 21.4 1.56 100 -3.33 2.47 5.5
Woodland Bog 97.8 8.3 0.30 100 -11.46 1.74 0
Portage Lake 11.6 1.0 7.92 100 11.29 10.58 91.5
SC senescence
Holt Pond ND ND ND ND ND ND ND
Dwinal 0 6.5 0.73 100 -11.90 3.50 0
Pickle Ridge ND ND ND ND ND ND ND
Mattagodus 61.2 5.6 2.63 100 -14.90 3.49 0
Salmon Stream 50.1 7.7 1.55 100 -7.40 2.02 0
Crystal Fen 100 -7.5 1.78 100 -21.60 2.54 0
Pillsbury Pond 37.4 ND ND 100 ND ND 0
Soper Pond 0 18.9 4.10 100 -15.10 5.78 0
Woodland Bog 0 1.8 5.21 100 -19.90 5.38 0
Portage Lake 22 -0.8 0.59 100 -8.90 6.58 12.7 a
Proportion of measurements in which water level in the deep well exceeded that in the
water table well. b
Proportion of measurements that fall at or within the rooting zone (30 cm from the peat
surface). c Proportion of measurements in which water levels are above the peat surface.
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Table 1.3. Upwelling (%), mean (±SD) difference (cm) between water table and deep
well, saturation (%), mean (±SD) depth (cm) to water table, and flood duration (%) for 10
wetlands monitored June 2009 – September 2009 in Maine. ND= No Data
Upwellinga
Mean difference Saturationb
Mean Depth to
Water Table
Flood
durationc
(%) (cm) (±SD) (%) (cm) (±SD) (%)
Holt Pond ND ND ND ND ND ND ND
Dwinal 0 5.98 0.95 100 -6.49 7.01 12.72
Pickle Ridge ND ND ND ND ND ND ND
Mattagodus 0 4.39 2.49 100 -9.39 5.12 0.19
Salmon Stream 2.3 7.43 4.45 100 -2.56 7.01 14.46
Crystal Fen 100 -6.85 1.27 100 -18.07 3.48 0
Pillsbury Pond ND ND ND ND ND ND ND
Soper Pond 0 19.75 0.95 100 -7.77 7.14 4.7
Woodland Bog 95.5 -0.88 0.45 100 -14.96 5.41 0
Portage Lake 26.5 4.82 6.36 100 -0.51 10.9 48 a
Proportion of measurements in which water level in the deep well exceeded that in the
water table well. b
Proportion of measurements that fall at or within the rooting zone (30 cm from the peat
surface). c Proportion of measurements in which water levels are above the peat surface.
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Table 1.4. Upwelling (%), mean (± standard deviation, SD) difference (cm) between
water table and deep well, saturation (%), depth (cm) to water table (mean ±SD), and
flood duration (%) for 10 wetlands monitored May– September 2010. Data are reported
for leaf out of the shrubby cinquefoil (SC) and the egg period of the Clayton’s copper
(CC, 12 May- 28 May) Clayton’s copper larval feeding (29 May-14 July), shrubby
cinquefoil bloom and Clayton’s Copper flight (15 July – 19 August), and shrubby
cinquefoil senescence periods in Maine. ND= No Data
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Upwellinga Mean difference Saturationb Mean Depth to
Water Table
Flood
durationc
(%) (cm) (±SD) (%) (cm) (±SD) (%)
SC leaf out/ CC egg
Holt Pond 100 -3.86 0.81 100 -0.71 1.01 25.3
Dwinal 0 8.2 0.71 100 1.4 1.75 78.2
Pickle Ridge 0 n/a - 100 4.9 2.17 100
Mattagodus 0 4.82 1.55 100 -13.5 1.89 0
Salmon Stream 0 15.21 1.96 100 -11.8 1.69 0
Crystal Fen 0 0.99 0.21 100 -9.1 1.62 0
Pillsbury Pond 0 3.58 0.79 100 -7.8 2.55 0
Soper Pond 0 14.79 1.16 100 -5.4 1.31 0
Woodland Bog 97.9 -0.96 0.23 100 -10.4 1.08 0
Portage Lake 71.1 -1.06 1.44 100 1.1 4.29 65.8
CC larval feeding
Holt Pond 95.7 -3.59 1.61 100 -5.3 2.53 16.8
Dwinal 0 7.7 1.57 100 0.5 3.95 61.6
Pickle Ridge 0 n/a - 100 7.2 4.05 97.9
Mattagodus 0 5.11 1.8 100 -11.9 2.8 0
Salmon Stream 0 12.04 1.65 100 -10.4 3.19 0
Crystal Fen 0.3 0.92 0.31 100 -8.1 2.02 0
Pillsbury Pond 5.6 4.48 2.53 100 -8.8 4.59 0
Soper Pond 0 13.94 2.48 100 -7.4 3.56 0
Woodland Bog 92.8 -0.87 0.57 100 -12.5 2.78 0
Portage Lake 34.4 -0.04 4.01 100 -0.9 5.28 58.7
SC bloom/ CC adult nectaring Holt Pond 100 -7.14 2.53 100 -16 6.09 0
Dwinal 0 5.25 1.42 61.6 -24.8 14.36 1.79
Pickle Ridge 24.2 0.63 1.99 100 -7.9 5.36 7.18
Mattagodus 42.3 0.28 2.32 71.2 -26.3 6.73 0
Salmon Stream 29.9 3.32 7.58 73.1 -23.8 10.71 0
Crystal Fen 1.2 0.77 0.44 100 -15.3 2.81 0
Pillsbury Pond 9.9 2.85 2.03 100 -9.1 4.42 0
Soper Pond 0 9.59 5.64 100 -7.1 2.68 1.45
Woodland Bog 9.08 0.37 0.59 100 -15.8 5.46 0
Portage Lake 73.4 -4.79 5.37 99.2 -9.2 9.04 25.8
SC senescence
Holt Pond 49.5 -1.81 5.78 100 -15.8 7.41 0
Dwinal 0 8.18 5.50 48.8 -27.1 22.04 2.4
Pickle Ridge 41.1 -0.48 4.59 100 -15.4 5.25 0
Mattagodus 61.2 -0.54 7.91 24.2 -37.0 7.50 0
Salmon Stream 50.1 -0.05 17.44 48.9 -37.1 18.49 0
Crystal Fen 0 1.50 0.43 100 -16.0 5.79 0
Pillsbury Pond 37.4 1.68 5.41 100 -16.8 10.97 0
Soper Pond 0 4.44 2.06 100 -9.4 4.62 0
Woodland Bog 0 7.62 7.90 56.8 -23.4 11.94 0
Portage Lake 22 1.81 1.33 80.8 -15.9 14.74 16.2 a
Proportion of measurements in which water level in the deep well exceeded that in the
water table well. b
Proportion of measurements that fall at or within the rooting zone (30 cm from the peat
surface). c Proportion of measurements in which water levels are above the peat surface.
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Table 1.5. Upwelling (%), mean (±SD) difference (cm) between water table and deep
well, saturation (%), mean (±SD) depth (cm) to water table, and flood duration (%) for 10
wetland monitored May 2010 – September 2010 in Maine.
Upwellinga
Mean difference Saturationb
Mean Depth to
Water Table
Flood
durationc
(%) (cm) (±SD) (%) (cm) (±SD) (%)
Holt Pond 86.7 -4.2 3.80 100 -10.65 7.40 9
Dwinal 0 7.2 3.17 77.4 -12.84 18.80 33
Pickle Ridge 21.5 3.7 9.04 100 -2.51 10.42 50
Mattagodus 26 2.4 4.92 74.3 -21.97 11.49 0
Salmon Stream 19.9 7.2 11.00 78.8 -21.27 15.27 0
Crystal Fen 0.4 1.0 0.45 100 -12.11 4.98 0
Pillsbury Pond 13.5 3.3 3.40 100 -10.62 7.31 0
Soper Pond 0 9.0 5.56 100 -7.51 3.64 0.4
Woodland Bog 48.5 0.1 1.35 90 -15.66 8.09 0
Portage Lake 47 0.3 6.97 95.3 -6.42 11.23 41
a
Proportion of measurements in which water level in the deep well exceeded that in the
water table well. b
Proportion of measurements that fall at or within the rooting zone (30 cm from the peat
surface). c Proportion of measurements in which water levels are above the peat surface.
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Figure 1.2. Running average of inundation and timing of Clayton’s copper (CC) life
history stages during the 2009 and 2010 growing seasons. SC= shrubby cinquefoil.
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1.3.2. Pore water characterization
Wetlands separated along the electrical conductivity (EC) and hydrogen ion
concentration (H+) gradient in a principal component analysis (PCA) of pore water
analytes, with 63.8-77.6% of the variation explained by the first two dimensions.
Wetlands generally grouped by hydrological types during L. d. claytoni larval emergence
(24 May – 3 June; Fig. 1.3; Appendix B). HT1 wetlands were associated with lower H+
and greater EC; HT2 and HT3 wetlands exhibited intermediate component scores along
both axes; HT4 showed the greatest spread across both axes; and, HT5 separated on the
first axis, indicating greater H+ and lower EC. MRPP confirmed that Portage (HT5)
differed from other wetlands except Mattagodus (HT2; T=-7.4 to -14.3, A-statistics
=0.468 to 0.854; Table 1.6). During D. fruticosa blooming and L. d. claytoni nectaring
(12 July – 23 July 2010) and D. fruticosa senescence (17 – 25 August 2010; Figs. 1.4 and
1.5), pore water analytes did not separate by hydrological types. During D. fruticosa
bloom and L. d. claytoni nectaring, pore water from Pickle (HT4) had the greatest range
along the EC/H+ axis. Despite considerable overlap in pore water PCA scores, Portage
(HT5) differed from the other wetlands (P<0.001, T =-6.85 to -15.23, A= 0.316 to 0.805)
(Table 1.7). Concentrations of pore water analytes from Woodland and Crystal (HT1)
during senescence (late August) were significantly different, according to MRPP
analysis, from all wetlands except Pickle (P≤ 0.001, T= -5.76 to -11.17, A= 0.315 to
0.769) and Holt (P≤ 0.001, T= -5.64 to -11.17, A= 0.187 to 0.700) (Table 1.8).
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Figure 1.3. Principal component biplots of water analytes from pore water samples
collected from wetlands in Maine during larval emergence (24 May -3 June 2010).
Analytes exhibiting heavy loadings (|loading| > 0.5) listed on appropriate axes; the
dominant analyte loading on each axis is indicated in bold. All loadings are listed in
Appendix Table C.1. Within wetland site clusters are enclosed by 68% confidence
ellipses.
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Figure 1.4. Principal component biplots of water analytes from pore water samples
collected from wetlands in Maine during L. d. claytoni nectaring (12 – 23 July 2010).
Analytes exhibiting heavy loadings (|loading| > 0.5) listed on appropriate axes; the
dominant analyte loading on each axis is indicated in bold. All loadings are listed in
Appendix Table C.1. Within wetland site clusters are enclosed by 68% confidence
ellipses.
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Figure 1.5. Principal component biplots of water analytes from pore water samples
collected from wetlands in Maine during D. fruticosa senescence (17 – 25 August 2010).
Analytes exhibiting heavy loadings (|loading| > 0.5) listed on appropriate axes; the
dominant analyte loading on each axis is indicated in bold. All loadings are listed in
Appendix Table C.1. Within wetland site clusters are enclosed by 68% confidence
ellipses.
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Table 1.6. Multiple-response permutation procedure results of water from pore water samples collected from wetlands grouped by
geographic region in Maine during larval emergence (24 May -3 June 2010). T-Statistic (first row) and A-statistic (third row) listed for
each pair with P-value in parentheses (P≤ 0.001 in bold).
Site Dwinal Holt Mattagodus Pickle Salmon Crystal Pillsbury Soper Woodland Portage
Dwinal -2.400 -7.057 -4.786 -10.290 -15.070 0.062 -1.340 -12.290 -14.310
(0.035) (<0.001) (0.003) (<0.001) (<0.001) 0.375 0.096 (<0.001) (<0.001)
0.060 0.256 0.160 0.421 0.502 -0.002 0.040 0.567 0.624
Holt -6.580 -0.917 -3.500 -6.130 -1.030 0.550 -7.250 -12.490
(<0.001) (0.143) (0.011) (<0.001) (0.128) (0.637) (<0.001) (<0.001)
0.239 0.032 0.137 0.179 0.037 -0.017 0.305 0.526
Mattagodus -4.260 -7.460 -11.658 -2.574 3.720 -8.200 -3.210
(0.006) (<0.001) (<0.001) (0.029) (0.010) (<0.001) (0.014)
0.237 0.526 0.597 0.134 0.170 0.603 0.145
Pickle -1.360 -3.690 -2.440 -0.992 -2.030 -7.430
(0.096) (0.006) (0.033) (0.133) (0.047) (<0.001)
0.080 0.136 0.131 0.043 0.131 0.483
Salmon -0.954 -6.520 -3.510 -5.740 -9.380
(0.138) (<0.001) (0.010) (<0.001) (<0.001)
0.050 0.435 0.187 0.491 0.826
Crystal -10.470 -5.890 -10.200 -13.66
(<0.001) (<0.001) (<0.001) (<0.001)
0.492 0.229 0.612 0.861
Pillsbury -0.280 -7.750 -9.040
(0.271) (<0.001) (<0.001)
0.012 0.574 0.365
Soper -7.120 -9.010
(<0.001) (<0.001)
0.365 0.468
Woodland -9.410
(<0.001)
0.854
Portage
Mid-State Northwestern Northeastern
X
X
X
X
X
X
X
X
X
X
23
24
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Table 1.7. Multiple-response permutation procedure results of water analytes from pore water samples collected from wetlands
grouped by geographic region in Maine during L. d. claytoni nectaring (12 – 23 July 2010). T-Statistic (first row) and A-statistic (third
row) listed for each pair with P-value in parentheses (P≤ 0.001 in bold).
Site Dwinal Holt Mattagodus Pickle Salmon Crystal Pillsbury Soper Woodland Portage
Dwinal -4.890 -8.050 -10.640 -10.960 -15.450 -2.880 -2.750 -13.410 -15.230
(0.002) (<0.001) (<0.001) (<0.001) (<0.001) (0.018) (0.018) (<0.001) (<0.001)
0.114 0.303 0.388 0.051 0.648 0.088 0.076 0.778 0.699
Holt -2.510 -2.310 -2.530 -5.800 -0.716 0.603 -6.63 -7.530
(0.031) (0.037) (0.030) (0.001) 0.018 0.674 (<0.001) (<0.001)
0.091 0.080 0.102 0.182 0.025 -0.02 0.271 0.316
Mattagodus -7.070 7.390 -11.720 -0.640 -2.110 -8.660 -6.850
(<0.001) (<0.001) (<0.001) (0.189) (0.460) (<0.001) (<0.001)
0.390 0.488 0.595 0.033 0.094 0.666 0.324
Pickle 0.761 -0.688 -4.840 -3.720 -4.030 -9.930
(0.794) (0.199) (0.003) (0.008) (0.006) (<0.001)
-0.033 0.019 0.264 0.149 0.181 0.618
Salmon 0.069 -5.560 -3.980 -5.210 -9.080
(0.440) (0.001) (0.005) (0.001) (<0.001)
-0.003 0.362 0.189 0.348 0.701
Crystal -9.730 -8.360 -10.610 -13.340
(<0.001) (<0.001) (<0.001) (<0.001)
0.479 0.301 0.491 0.776
Pillsbury -0.024 -7.780 -8.680
(0.356) (<0.001) (<0.001)
0.001 0.607 0.464
Soper -8.020 -7.660
(<0.001) (<0.001)
0.407 0.389
Woodland -9.330
(<0.001)
0.805
Portage
Mid-State Northwestern Northeastern
X
X
X
X
X
X
X
X
X
X
24
25
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26
Table 1.8. Multiple-response permutation procedure results of water analytes from pore water samples collected from wetlands
grouped by geographic region in Maine during D. fruticosa senescence (17 – 25 August 2010) T-Statistic (first row) and A-statistic
(third row) listed for each pair with P-value in parentheses (P≤ 0.001 in bold).
25
26
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1.3.3. Peat characterization
The first three principal components derived from peat analytes explained 56% of
the variation in peat nutrient and mineral composition among wetlands (Fig. 1.6).
Wetlands were loosely grouped by hydrological types. Peat from HT1 wetlands contained
greater concentrations of base cations, whereas peat from HT2 wetlands exhibited
intermediate component scores along the first three axes. Pickle (HT4) peat contained
greater concentrations of Mn, Zn, and Na, and peat from Portage (HT5) was isolated on
the first axis with greater concentrations of Fe, Al, and H+, less LOI, and lower
concentrations of base cations. MRPP results confirmed differences in Portage peat
nutrient and mineral composition compared with the other wetlands (Table 1.9; P<0.001,
T= -9.18 to -15.27). MRPP results also revealed that peat from Holt (HT1) differed from
that collected from all other wetlands (Table 1.9; P<0.001, T= -5.15 to -15.27, A=0.124
to 0.683), although the PCA biplots indicated overlap.
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Figure 1.6. Principal component biplots of peat analytes at the 10 wetlands. Analytes
exhibiting heavy loadings (|loading| > 0.4) listed on appropriate axes; the dominant
analyte loading on each axis is indicated in bold. All loadings are listed in Appendix
Table C.2. Within wetland site clusters are enclosed by 68% confidence ellipses.
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Table 1.9. Multiple-response permutation procedure results of peat analytes from wetlands grouped by geographic region in Maine,
2010. T- Statistic (first row) and A-statistic (third row) listed for each pair with P-value in parentheses (P≤ 0.001 in bold).
Site Dwinal Holt Mattagodus Pickle Salmon Crystal Pillsbury Soper Woodland Portage
Dwinal -6.590 -1.920 -2.500 -2.139 -6.020 -2.156 -2.610 -2.100 -14.400
(<0.001) (0.054) (0.031) (0.043) (<0.001) (0.040) (0.030) (0.050) (<0.001)
0.173 0.060 0.078 0.089 0.170 0.083 0.073 0.095 0.533
Holt 9.090 -10.800 -7.870 -13.100 -5.150 -12.620 -7.820 -15.270
(<0.001) (<0.001) (<0.001) (<0.001) (<0.001) (<0.001) (<0.001) (<0.001)
0.269 0.317 0.226 0.364 0.124 0.337 0.224 0.683
Mattagodus -1.440 -3.310 -3.860 -4.560 -0.920 -3.900 -10.740
(0.088) (0.009) (0.004) (0.003) (0.150) (0.005) (<0.001)
0.048 0.139 0.096 0.168 0.025 0.181 0.563
Pickle -3.640 -9.080 -5.590 -3.470 -4.980 -10.890
(0.009) (<0.001) (0.001) (0.010) (0.002) (<0.001)
0.150 0.220 0.198 0.093 0.218 0.602
Salmon -8.540 -4.080 -4.550 -3.510 -9.180
(<0.001) (0.003) (0.004) (0.005) (<0.001)
0.246 0.143 0.164 0.124 0.675
Crystal -7.790 -8.970 -9.220 -13.40
(<0.001) (<0.001) (<0.001) (<0.001)
0.226 0.214 0.268 0.709
Pillsbury -7.57 -1.87 -10.56
(<0.001) (0.060) (<0.001)
0.239 0.068 0.680
Soper -6.38 -12.97
(<0.001) (<0.001)
0.240 0.588
Woodland -9.22
(<0.001)
0.705
Portage
Mid-State Northwestern Northeastern
X
X
X
X
X
X
X
X
X
X
29
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1.3.4. Wetland woody species characterization
Shrub species composition was similar among the wetlands, and D. fruitcosa was
the dominant shrub species in all sampled wetlands except Pickle Ridge and Portage
Lake (Fig. 1.8). The shrub community was most diverse in Holt, whereas, Woodland
contained the least diverse shrub community (Appendix D). Average D. fruticosa volume
(Fig 1.7, Appendix E and F) and linear coverage (totaled across transects; Fig. 1.8)
differed among wetlands. Dasiphora fruticosa abundance and size generally were greater
in wetlands with consistent water levels (HT1, HT2; Fig. 1.7 and 1.8), however, Dwinal
and Crystal were exceptions to this trend. MRPP confirmed that shrub diversity and
coverage in Portage (Table 1.10; P<0.001, T=-6.23 -11.98, A= -0.148 - 0.497) and Pickle
(P<0.001, T =-5.69 to -12.23, A= 0.156 to 0.381) differed from all other wetlands, and
shrub diversity and coverage in Crystal differed from wetlands other than Salmon and
Woodland (P<0.001, T=-5.38 to -12.21, A=0.127 to 0.342).
The number of tree species in the wetlands ranged 1-6, with the least diversity in
Portage Lake and greatest species diversity in Dwinal (Appendix F). The dominant tree
species was Thuja occidentalis in all wetlands except Pickle, where Betula populifolia
was dominant and Crystal Fen, where Larix laricina was dominant. Total tree basal area
(summed across wetland transects) ranged 0.8 – 400 cm/transect m among wetlands (Fig.
1.9). Dasiphora fruticosa abundance and volume generally were greater in wetlands with
intermediate tree basal area.
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Figure 1.7. Average Dasiphora fruticosa volume (m3). Error bars represent 1 SE. Bars
with the same lowercase letter are not significantly different (P≥0.05). Hatched bars
indicate unoccupied wetlands.
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Figure 1.8. Shrub coverage (m3/transect m) totaled along transects in 10 wetlands.
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Table 1.10. Multiple-response permutation procedure results of shrub community from 10 wetlands in mid-state, northwestern and
northeastern Maine, 2010. T-Statistic (first row) and A-statistic (third row) are listed for each pair with P-value in parentheses (P≤
0.001 in bold).
32 3
3
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Figure 1.9. Tree basal area (cm2/transect m) totaled among transects in sampled wetlands.
Hatched bars indicate unoccupied wetlands.
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1.4. Discussion
1.4.1. Hydrological environments of D. fruticosa
Wetland structure and function are determined in part by the hydrological
environment (Rydin and Jeglum 2006). Water depth, flood duration and frequency, and
flow patterns affect wetland abiotic factors such as nutrient availability and peat
anaerobiosis (Mitsch and Gosselink 2007) as well as cause physical disturbances that can
affect vegetation recruitment (Keddy 2000). These factors in turn directly influence the
establishment of vegetation communities in a wetland (Mitsch and Gosselink 2007).
Dasiphora fruticosa often is found in moist conditions in North America (USDA,
Magee and Ahles 1999) and Britain (Elkington and Woodell 1963). I found the species
occurring in wetlands with a broad range of hydrological regimes. Despite differences in
water sources and variation in inundation frequency and duration, all wetlands were
saturated in the D. fruticosa rooting zone during leaf out. Nearly all the sampled wetlands
were saturated for the majority of the growing season both years, and water pooled on the
peat surface in six of the 10 wetlands during at least one of the study’s two growing
seasons. Dasiphora fruticosa occurs in similar growing season conditions in Britain and
Sweden, where wetlands with the species flood through the winter (Elkington and
Woodell 1963). Although all of the wetlands I studied are snow covered in the winter,
they either are inundated or saturated early in the growing season.
Although Dasiphora fruticosa is a stress-tolerant species (sensu Grime 1979) that
occurs in a variety of hydrological settings, the range of conditions tolerated by D.
fruticosa may not be suitable for L. d. claytoni survival and persistence. Butterfly host
plants found in wetlands may be adapted to flooded conditions, however, butterfly eggs
and larvae may not tolerate submergence (Webb and Pullin 1998, Nicholls and Pullin
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2003, Severns et al 2006). Eggs of Lycaena xanthoides on inundated plants were seven
times less likely to survive as eggs on non-flooded plants (Severns et al. 2006). Wetlands
that had consistent water tables and less growing season inundation in my study had
greater L. d. claytoni encounter rates in 2007 and 2008 (Knurek 2010). Flooding on the
peat surface was longest during May-July in 2009 and 2010, when L. d. claytoni is in the
egg and larval stages. Lycaena dorcas claytoni eggs are in the accumulated leaf litter at
the base of D. fruticosa plants after leaf dehiscence, and fluctuating water levels
following leaf drop (such as those recorded at Salmon, Pickle, and Portage) may drown
the eggs. Similarly, L. d. claytoni larvae have limited mobility and may drown if the host
plant is submerged rapidly or completely. Larvae of Lycaena dispar batavus that are
submerged for 28 days experience increased mortality (Nicholls and Pullin 2003). In
addition to drowning mortality, flooding that makes leaves inaccessible also may cause
larval mortality (Joy and Pullin 1999). Wetland microtopographic variation may provide
habitat for D. fruticosa that is not flooded during the L. d. claytoni egg and larval periods.
Although L. d. claytoni occurs in relative abundance in four of 10 wetlands in Maine,
relationships of wetland inundation depth and timing, quality of D. fruticosa host plants
as larval food, and L. d. claytoni population response to these habitat conditions are
unclear. Similarly, more information is needed about relationships of within-wetland and
landscape-scale habitat composition and arrangement, Clayton’s Copper movement
abilities, and long-term persistence of its populations.
1.4.2. Chemical environments of D. fruticosa and relevance to L. d. claytoni
Although hydrology is considered the primary driver of wetland plant community
establishment and persistence, relationships between wetland plant community
composition and structure and hydrological conditions are complex and not well
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understood (Carter 1986, LaBaugh 1986, Mitsch and Gosselink 2007). Mineral and
oxygen concentrations in a wetland substrate are determined by the source and velocity
of water moving through the wetland (Wassen et al. 1990). The water source and
dynamics of its delivery can influence nutrient availability for wetland plants by
supplying nutrients for direct uptake or by creating conditions that affect the release of
nutrients from the organic soils (Boomer and Bedford 2008) through changes in
carbonate chemistry (Boyer and Wheeler 1989) or redox potential (Smolders et al. 2006).
Vegetation composition in fens in Biebrza, Poland, correlated best with root-zone water
chemistry when weather conditions were extreme (i.e., drought or extremely wet
conditions ; de Mars et al. 1997). Fluctuations in the water table can affect availability of
nutrients in peat and pore water (Carter 1986), which may affect overall productivity and
vegetation composition in a wetland (Mitsch and Gosselink 2007).
Variation in water source and inundation timing complicate characterization of
groundwater-fed wetlands. Pore water chemistry in the D. fruticosa rooting zone in
wetlands monitored in this study generally reflected the wetland’s water source. Wetlands
dominated by groundwater discharge had greater pore water pH and conductivity owing
to mineral nutrients transported in groundwater (Wassan et al 1990). Wetlands dominated
by ground water recharge generally had a lower pH reflecting a reduction in ion
concentration in surface water that dilutes or leaches nutrients from the peat (Mitsch and
Gosselink 2007). Although the sampled wetlands span the pH and conductivity gradient,
there was little separation along the productivity gradient reflecting concentrations of
NH4-N, NO3-N, and PO4-P.
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Plant community types in northern peatlands reflect broad nutrient gradients
(Bedford et al.1999) and nutrient availability (Bragazza and Gerdol 2002). Peat chemistry
in the surveyed wetlands was only weakly related to water source and hydrological types.
Wetlands with groundwater discharge (Holt and Woodland) generally had greater
concentrations of base cations, reflecting the wetland water source, however, peat
nutrient concentrations were similar among wetlands. The minerals and nutrients in peat
generally are bound in organic forms and may not be available for plant uptake (Mitsch
and Gosselink 2007). All of the study wetlands were saturated the majority of the
growing season, and nutrient availability may have been reduced in these wetlands by
anoxic conditions.
Fen vegetation communities are associated with high calcium concentrations, and
P-limitation due to co-precipitation with calcium maintains these plant communities
(Bedford and Godwin 2003). The nutrient profiles among my sampled wetlands were
similar, as was shrub vegetation community composition, and D. fruticosa was the
dominant shrub in all but two of these wetlands. Calcium was the dominant cation in the
peat nutrient profiles, and there were no deficiencies in micro minerals, conditions similar
to those where D. fruticosa is found in Britain (Elkington and Woodell 1963). Phosphate
concentrations in peat samples varied within and across wetlands, indicating that elevated
concentrations of phosphate are not necessary for D. fruticosa, similar to conditions
where D. fruticosa occurs in Britain (Elkington and Woodell 1963). Relationships of
seasonal changes in hydrological conditions, peat nutrients and dynamics of uptake of
those nutrients by D. fruticosa are not known.
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Dasiphora fruticosa size, coverage, and growth form differed among the
wetlands. Dasiphora fruticosa abundance and coverage were greatest at wetlands
(Mattagodus, Pillsbury and Soper) with consistent water tables, little to no flooding but
saturated root zone, and intermediate concentrations of peat and pore water analytes.
Wetlands with a fluctuating water table may receive brief pulses of enrichment or
dilution contributed by overbank flooding, upwelling, or surface water runoff. Flooding
and drawdown events also may create openings in the ground cover to create areas
suitable for shrub seedling recruitment as well as enhance availability of nutrients for
seedlings. Portage Lake experienced dynamic water table fluctuation with frequent
overbank flooding during both growing seasons, and the shrub community is diverse
despite the wetland’s small size. Pickle Ridge experienced extended periods of
inundation through the beginning of the growing season. Dasiphora fruticosa size and
coverage at both of these sites were relatively small while wetlands with more consistent
water levels (Soper and Pillsbury) have larger individual shrubs and more homogenous
stands of D. fruticosa. Little is known about the physiological and metabolic responses of
D. fruticosa to water level variation and relationships to the species’ germination and
growth, which may explain morphological differences found among the wetlands.
Resource quantity is a key predictor of densities of monophagus habitat specialist
butterflies (Krass et al. 2003 León-Cortés et al. 2003, Krauss et al. 2005), and adult
densities may be driven by host plant quantity regardless of plant quality or patch
isolation (Krauss et al. 2005). Dasiphora fruticosa coverage in Maine’s wetlands may
affect population density of L. d. claytoni. Lycaena dorcus claytoni encounter rates
tended to be greater (Kneruk 2010) where D. fruticosa abundance and average size are
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larger. Thomas (1983) found that loss of the larval food plant accounted for
approximately one third of the extinctions of Lysandra bellargus in Britain. Wetlands
occupied by L.d. claytoni occur in three general regions in Maine. I surveyed three
wetlands with robust D. fruticosa unoccupied by L.d. claytoni. It is not known if L.d.
claytoni can disperse to these wetlands without assistance given the distance and lack of
contiguous wetland habitat separating these wetlands; understanding the dispersal ability
of the species is key to identifying wetlands with D. fruticosa for conservation.
1.4.3. Management implications
Although large quantities of larval food plants may be a critical factor for the
persistence of food plant specialists (Krauss et al. 2005), connectivity among locations
with the food resource also may be important (e.g., Fahrig and Merriam 1995). Lycaena
d. claytoni conservation will be enhanced by management strategies that maintain
wetlands with extensive D. fruticosa coverage and that connect wetlands with suitable
dispersal habitat. Conditions that contribute to long-term persistence and quality of D.
fruticosa populations are not known.
Although D. fruticosa can tolerate a range of habitat conditions (Elkington and
Woodell 1963), management strategies that reduce disturbances in the surrounding
watersheds may be critical for maintaining hydrological conditions suitable for the
species. Wetlands with D. fruticosa were classified as seasonally flooded or saturated
circumneutral wetlands; however, hydrological conditions differed among the study
wetlands and between years. Long-term studies of effects of hydrological conditions in
wetlands with D. fruticosa and on the life stages of L.d. claytoni are needed, with
particular emphasis on how spring flooding affects early life stages of L .d. claytoni and
nutrient uptake of D. fruticosa during leaf out.
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CHAPTER 2
ASSESSMENT OF SHRUBBY CINQUEFOIL (DASIPHORA FRUTICOSA) AS
THE HOST PLANT FOR CLAYTON’S COPPER BUTTERFLY (LYCAENA
DORCAS CLAYTONI)
2.1. Introduction
A fundamental goal of species conservation is ensuring accessible habitat,
including adequate food resources. While landscape-scale qualities such as patch
arrangement and dynamics affect the occurrence and persistence of butterfly populations
(Hanski 1998), local patch characteristics and quality and quantity of the host plant
within patches also are important (Thomas et al. 2001, Leon-Cortes et al. 2003, Krauss et
al. 2005). Availability of host plants for larval and adult butterfly food resources is
critical for maintaining viable butterfly populations (Shultz and Dlugosch 1999). The
conditions and quality of the host plant affect survival of eggs and larvae, which
influence butterfly population growth and extinction (Ehrlich and Hanski 2004). Butterfly
population persistence also is affected by quality of nectar plants, which can influence
adult butterfly location and dispersal (Dover 1997). While classic metapopulation
variables such as habitat patch size and isolation may affect butterfly population
occurrence (Hanski 1998), an individual adult butterfly’s use of habitat, including a
female’s ovipostion decision, are influenced by within-patch nectar plant quality.
Successful conservation of rare and endangered butterfly species requires consideration
of within-site host plant quality for all life history stages of the species.
Food resource requirements differ among butterfly life history stages. Egg and
larval survival are affected by adult dispersal and oviposition site selection (García-
Barros and Fartmann 2009), which may be affected by the host plant foliar chemistry
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(e.g., Ehrlich and Raven 1964, Barros and Zucoloto 1999). Resource quantity and quality
for larval butterflies is positively correlated with population size and persistence (Shultz
and Dlugosch 1999). Resource condition during larval development is particularly
important as the juvenile stages typically are the longest life stage (García-Barros and
Fartmann 2009). Phytophagous insects preferentially feed on new, growing leaves,
because young leaves are less fibrous and contain more foliar nitrogen (Marquis and
Braker 1994, Coley and Barone 1996). Leaf nutritional value is affected by nitrogen and
water content (Coley et al. 2006), which can affect caterpillar development rate.
Environmental factors that alter a plant’s physiology and biochemistry potentially change
its nutritional value to herbivores (Bryant et al. 1983, Herms and Mattson 1992, Mattson
and Haack 1987). For example, water stress reduces plant growth, resulting in smaller
plants and plant parts (e.g., leaves, buds) (Kramer 1983). Although host plant quality is
affected by within-site characteristics, environmental conditions at the watershed scale
(e.g., seasonal precipitation patterns) also potentially influence suitability of these
resources for developing larvae (Santiago and Mulkey 2005).
Floral nectar is the most common adult butterfly food and is the primary reward
butterflies acquire from flowers (Gilbert and Singer 1975). Nectar plant availability and
host plant nectar supply and quality affect adult butterfly distribution (Wiklund 1977,
Schneider et al. 2003, Aucklandet al. 2004), population densities, and movement (Britten
and Riley 1994). Greater concentrations of carbohydrates in the adult diet increase
butterfly longevity (Hill and Pierce 1989, Cahenzli and Erhardt 2012) and fecundity (Hill
and Pierce 1989). Nectar is composed primarily of amino acids, lipids, and sugars
(fructose, glucose, sucrose) (Baker and Baker 1983a, Mevi-Schütz and Erhardt 2005).
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Proportions of nectar sugars generally are conserved within host plant species while
differing among species (Percival 1961, Baker and Baker 1979, 1982a, b). The ratio of
sucrose to hexose (fructose and glucose) affects pollinator type, with large ratios of
sucrose to hexose reported in butterfly-pollinated flowers (Erhardt 1991, 1992,
Rusterholz and Erhardt 1997). Suitability of a plant species as a butterfly host may be
determined in part by interspecific differences in butterfly flight distance, life histories,
and reproductive strategies that affect their energy and nutritional needs and nectar
demands (Boggs 1997).
Plant age also may affect nectar quality and availability (Búrquez and Corbet
1998). As perennials age, resource allocation shifts from growth and leaf expansion to
reproduction (Bond 2000). For example, the herbaceous perennial Corydalis intermedia
increases flower production with increasing size and total leaf area until approximately
11 years old, when bloom production becomes constant (Ehlers and Olesen 2004).
Although age-related flower production in woody perennials is not well-studied, flower
bud vigor in the Mediterranean shrub Cistus albidus decreases with increasing plant age
(Oñate and Munné-Bosch, 2010). Similarly, nectar secretion in the annual herb Impatiens
glandulifera (Balsaminaceae) decreases in older plants (Búrquez and Corbet 1998). Age-
dependent nectar quality and quantity potentially can affect butterfly population
persistence, particularly for those reliant on few host plant species.
Shrubby cinquefoil (Dasiphora fruticosa) is the sole host plant of larval and adult
Clayton’s copper butterflies (Lycaena dorcas claytoni), a Maine state endangered species.
The survival and persistence of this butterfly is dependent on the availability and quality
of D. fruticosa for larval and adult food. Although D. fruticosa is not considered rare, few
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wetlands in Maine support large, persistent stands of shrubby cinquefoil (McCollough et
al. 2001), and it is not known if D. fruticosa quality as larval and adult food for L. d.
claytoni varies among these wetlands. The objective of this study was to compare D.
fruticosa leaf quality (i.e., foliar nitrogen, moisture), nectar quality, and shrub age
structure and growth rate among and within Maine’s wetlands.
2.2. Methods
2.2.1. Study area
Clayton’s copper butterfly currently is found in wetlands in central (Dwinal Pond
Wildlife Management Area, Mattagodus Meadows, Holt Pond, Pickle Ridge), western
(Pillsbury Pond, Soper Pond, Little Round Pond), and northern (Woodland Bog and a
nearby wetland) Maine (Fig, 1.1). More butterflies were observed at Dwinal, Mattagodus,
Holt, Pillsbury and Soper in 2007-2008 than were observed in Pickle Ridge and
Woodland Bog during this period (Knurek 2010). All of these wetlands except Little
Round Pond and the wetland near Woodland were included in my study.
Three wetlands (Portage Lake, Salmon Stream, Crystal Fen) not inhabited by
Clayton’s copper were selected for comparison with currently occupied wetlands based
on their proximity to occupied wetlands (Fig. 1.1) and abundance of shrubby cinquefoil
found at these sites. Although a small population of Clayton’s copper previously was
observed at Crystal Fen (McCollough et al. 2001), the butterflies have not been observed
at Crystal Fen since 2008 (E. Knurek 2010, Drahovzal personal observation). Crystal Fen
was considered unoccupied for this study.
Knurek (2010) established transects for assessment of the taxonomic and
population status of Clayton’s copper butterfly; I established additional transects in
wetlands with shrubby cinquefoil and without the butterfly. Transects traversed the most
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dense stands of D. fruticosa and captured each wetland’s vegetation and topographic
variation delineated in zones (near water edge, shrub-sedge interior, forested wetland;
Appendix A).
2.2.2. Foliar moisture and nitrogen
I collected leaves (n=119) during the L. d. claytoni larval growth period (29 May-
14 July) from top branches on randomly selected D. fruticosa plants in forested, non-
forested, and near-open-water zones of each transect. Pre-weighed whirl-pack bags
containing the collected leaves were stored on ice at 4°C and weighed within six hours of
collection to determine initial leaf sample weight. I dried leaves in a drying oven at 105ºC
until the sample reached a consistent weight, from which I calculated the proportion of
water in leaf samples by difference from the initial weight. I ground dried leaf samples
with a mortar and pestle and burned away the sample organic material in a muffle furnace
at 550 °C for 5 hr at the University of Maine’s Analytical Laboratory and Maine Soil
Testing Service Laboratory (ALMSTS). The ALMSTS determined leaf nitrogen (N) with
plasma emission spectrophotometric analysis.
I compared the mean N concentration in leaf samples among wetlands with a one-
way ANOVA. Results were subjected to post-hoc Tukey’s test with significance
determined at α ≤0.05. I compared proportion of foliar moisture among wetlands with a
Kruskal-Wallis test, because foliar moisture data could not be normalized. I evaluated
pairwise differences among the wetlands with follow-up Mann-Whitney U tests. I
assessed correlations between foliar N and foliar moisture with Pearson correlation
coefficients.
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2.2.3. Bloom surveys and nectar sampling
During the adult L. d. claytoni flight period (15 July – 19 August) I counted
blooms on each D. fruticosa shrub intersecting the transect center line and collected
nectar samples (n=503) from randomly selected D. fruticosa shrubs (n=83) in the forested
wetland, non-forested wetland, and near open water zones of transects. I selected six
newly opened blooms on each plant, enclosed three of these blooms in fine-meshed (250-
µm), nylon tulle to exclude nectivores, and left three blooms un-enclosed but marked.
After 24 hours, I placed the individual blooms in a vial with 2 mL of distilled water,
agitated the vial for one minute (Morrant et al. 2009), removed the bloom, and stored the
remaining sample on dry ice during transport to storage within 6 hours in a freezer (-15
°C). I quantified sucrose, fructose, and glucose in 100 µL samples with high performance
liquid chromatography (HPLC) and compared these concentrations to sugar
concentrations in standards.
I compared log-transformed fructose and glucose concentrations with a paired t-
test, and I compared sugar concentrations between enclosed and un-enclosed bloom
samples with two-sample t-tests. Bloom density was determined for each wetland by
calculating the number of blooms per transect meter on D. fruticosa shrubs intersecting
the transect.
2.2.4. Age structure
I randomly selected and harvested shrubs (n=145) from the forested, non-forested,
and near open water zones of each transect in each wetland during mid-September to
mid-October 2010 to examine D. fruticosa age structure in the study wetlands. I collected
all stems, attached branches and roots of individual shrubs; shrubs with a prostrate
growth form were extracted from the peat where adventitious roots originated.
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I dried the main stem and attached branches to a consistent weight to estimate
above ground stem biomass (AGB). I identified the main stem as the largest, central
stem, and secondary stems as branches arising from or near the base of the main stem. I
removed a 2 - 4 cm section from the main stem at the root collar, and I removed similar
sections from four secondary stems to measure secondary stem growth. I sanded the
sections, counted annual rings with the 2010 ring designated as time zero, and measured
ring widths along 4 radii (WINDENDRO™, Guay et al. 1992). I calculated the area of
each growth ring to eliminate the geometric decrease of ring width as stem diameter
increases.
I aggregated growth rate and age data for comparisons of: (i) all individuals
within a wetland, (ii) all individuals within a zone (forested wetland, non-forested
wetland and near open water) of the wetland and among all wetlands, and (iii) all
individuals of a growth form (upright or prostrate). I compared mean age of ramets
among wetlands with Kruskal-Wallis because of non-normality in ages, and I evaluated
pairwise differences among wetlands with post-hoc Mann-Whitney U tests. I examined
Pearson correlations between age and height (primary stem length), age and AGB and
age and the diameter of the main ramet among individuals in the wetlands, among the
three different zones, and among the two growth forms. I normalized growth rate data
with a square root transformation and compared average growth rate among wetlands and
among zones with a one-way ANOVA and post-hoc Tukey’s test with significance for P
<0.05. I created growth curves with the average growth rate by year and age of the shrub,
and I compared growth rates among wetlands and zones within wetlands with
Gleichläufigkeit (GLK) values (Eckstein and Bauch, 1969). The GLK score is based on
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sign tests and is calculated with a classical agreement test. A plus point is assigned when
both shrubs grow similarly compared to the previous year. A minus point is assigned
when one shrub grows more and one shrub grows less compared to the previous year. A
pairwaise GLK score is calculated from the sum of the points divided by the number of
compared years. I visually compared age structure and number of secondary branches
within age classes with histograms.
2.3. Results
2.3.1. Foliar nitrogen and moisture
Percent foliar N (F9,86 = 15.4, P <0.001) and proportion of leaf moisture (Kruskal-
Wallis Χ2
= 63.6 df = 9, P <0.001) differed among wetlands (Fig. 2.1 – 2.2), however,
foliar N did not differ with position (zone) within the wetlands (F2,93 = 1.9, P = 0.144).
Foliar N was correlated with leaf moisture only in Holt and Soper (Table 2.1).
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Figure 2.1. Average foliar moisture (%) for 10 wetlands. Bars with the same letters do not
differ significantly (Mann-Whitney U tests post hoc pairwise comparisons after one-way
ANOVA; threshold for significance P<0.05)
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Figure 2.2. Average foliar nitrogen (%) for 10 wetlands. Bars with the same letters do not
differ significantly (Tukey’s HSD post hoc comparison after one-way ANOVA; threshold
for significance P<0.05)
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Table 2.1. Pearson correlation coefficients (r) for foliar nitrogen (FN) and foliar moisture
(FM) for 10 wetlands in Maine, USA.
FN vs. FM
Wetlands n r
All wetlands 93 0.15
Holt 14 0.73*
Dwinal 15 -0.23
Upper 9 0.13
Lower 6 0.27
Pickle 5 -0.27
Mattagodus 9 0.22
Salmon 6 0.35
Crystal 12 0.57
Pillsbury 8 0.69
Soper 12 0.78*
Woodland 6 0.75
Portage 9 0.31
*p<0.01
2.3.2. Nectar sugar composition and bloom surveys
Dasiphora fruticosa produces hexose-dominant nectar (sucrose/[glucose +
fructose] < 0.1) with only trace amounts of sucrose measured in < 3% of the samples
(Fig. 2.3). The proportion of fructose to glucose was 1:1 for all samples (paired t-test,
enclosed =1.3, p=0.18, df=248; unenclosed=-1.16, p=0.24, df=248; Fig. 2.3). There was a
significantly greater glucose and fructose concentrations in enclosed versus unenclosed
blooms (two sample t-test, fructose t=15.5, p <0.001, df =444, glucose t=16.7, p <0.001,
df=444). Bloom density ranged from 0 (Pickle) to 1.5 (Salmon) blooms/transect m
(Fig.2.4). During L. d. claytoni flight period, 91 % of 349 D. fruiticosa plants contained 0
- 10 blooms, and < 1% had more than 30 blooms.
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Figure 2.3. Average concentration (mg/bloom) of fructose, glucose and sucrose for
unenclosed (solid bars) and enclosed blooms (hatched bars). Error bars represent ±1 SE.
Figure 2.4. Total bloom density (number of blooms/transect m) at 7 wetlands with L. d.
claytoni (solid bars) and 3 wetlands with D. fruticosa but without L. d. claytoni (hatched
bars).
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2.3.3. Age structure
Main stem age ranged 7 to 37 years, and average age differed among wetlands
(Kruskal-Wallis Χ2
=57.4 , df = 9, P <0.01, Fig. 2.5 ). Collected D. fruticosa stems were
oldest in Pillsbury (µ=22.4, SE=1.4) and Portage (µ =22, SE=0.4) and youngest in Pickle
(M =7.4, SE=0.5). Mean stem age did not differ among wetland zones (water edge, non-
forested wetland, forested wetland) (Kruskal-Wallis Χ2
=0.12, df = 2, P =0.94). Stem
height and AGB were not correlated with stem age among wetlands; stem diameter was a
better indicator of shrub age than stem height or AGB (Table 2.2). Within wetland zones,
shrub age, height, and AGB (Table 2.4) were correlated. Height and AGB were
correlated with age in upright shrubs, however, length and AGB were not correlated with
age in prostrate shrubs (Table 2.3).
Although average annual growth rates differed among wetlands (F 9,253 = 8.4,
p<0.001, Fig. 2.6), the trends in periods of increasing and decreasing growth were similar
among the mean growth curves among the wetlands (global GLK= 0.47, Table 2.5);
growth rate declined as shrubs aged and generally increased in the recent decade (Fig. 2.7
– 2.12). Growth curve shapes were also similar among wetland zones (global GLK= 0.57,
Fig. 2.12) with 61%, 57% and 54% agreement between near-water and non-forested,
near-water and forested, and non-forested and forested zones respectively. However,
growth rates differed among zones (F 2, 93 = 5.77, p = .03). Growth rate of D. fruticosa in
the non-forested wetland (M =2.05, SE=0.20) was slower than growth rate of D. fruticosa
collected near the water’s edge (M = 3.23, SE= 0.34; Tukey’s post-hoc p = 0.03),
whereas, growth rate of D. fruticosa in the forested wetland (M = 2.93,SE=0.30) did not
differ from those collected from the non-forested and near water zones.
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Age structure was similar among wetlands, with the predominant age of D.
fruticosa ranging 15 to 20 years (Figure 2.13), however, my sampling approach excluded
shrubs younger than seven years. Most side branches were produced from main ramets
from five to 10 years of age with side branch production decreasing beginning at 10 years
(Fig. 2.14).
Figure 2.5. Box plot of shrub age. Mann-Whitney U tests post hoc pairwise comparisons
after one-way ANOVA, threshold for significance P<0.05. For each box plot, the top bar
is the maximum observation, bottom bar is the minimum observation, top of the box is
the third quartile, bottom of the box is the first quartile, middle bar is the median value
and circles are outliers.
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Figure 2.6. Average annual growth rate (mm/year) for 10 wetlands. Bars with the same
letters do not differ significantly (Tukey’s HSD post hoc comparison after one-way
ANOVA; P<0.05).
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Figure 2.7. Growth rate curves by age (a) and year (b) for all sampled stems pooled over wetlands (n=145) derived from average stem
cross-sectional increment. Bars represent standard error.
a. b.
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0
2
4
6
8
10
12
14
16
18
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
rat
e(m
m^2
)
0
2
4
6
8
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12
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18
1974
1976
1978
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2004
2006
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2010
Gro
wth
rat
e(m
m^2
)
Holt
Dwinal
Pickle
Mattagodus
0
2
4
6
8
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
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e(m
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0
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8
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1976
1978
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1998
2000
2002
2004
2006
2008
2010
Gro
wth
rat
e(m
m^2
)
Holt
Dwinal
Pickle
Mattagodus
Figure 2.8 Growth rates by age (a) and year (b) of shrubs collected from Holt (n=17), Dwinal (n=29), Pickle (n=11) and Mattagodus
(n= 12) derived from average stem cross-sectional increment.
a. b.
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0
2
4
6
8
10
12
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16
18
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
rate
(mm
^2)
0
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4
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wth
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e(m
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Salmon
Crystal
0
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
rate
(mm
^2)
0
2
4
6
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20
02
20
04
20
06
20
08
20
10
Gro
wth
rat
e(m
m^2
)
Salmon
Crystal
Figure 2.9. Growth curve by age (a) and year (b) for Salmon (n=17) and Crystal (n=8) derived from average stem cross-sectional
increment.
a. b.
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59
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0
2
4
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8
10
12
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
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wth
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e(m
m^2
)
0
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Gro
wth
rat
e(m
m^2
)
Pillsbury
Soper
0
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6
8
10
12
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
rat
e(m
m^2
)
0
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8
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1974
1976
1978
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1988
1990
1992
1994
1996
1998
2000
2002
2004
2006
2008
2010
Gro
wth
rat
e(m
m^2
)
Pillsbury
Soper
Figure 2.10. Growth curve by age (a) and year (b) for Pillsbury (n=14) and Soper (n=12) from average stem cross-sectional increment.
a. b.
60
60
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0
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
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wth
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e(m
m^2
)
Woodland
Portage
0
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12
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
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e(m
m^2
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02
20
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20
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Gro
wth
rat
e(m
m^2
)
Woodland
Portage
Figure 2.11. Growth curve by age (a) and year (b) for Woodland (n=11) and Portage (n=12) derived from average stem cross-sectional
increment.
a. b
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0
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1819
74
1976
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e (m
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1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37
Age (years)
Gro
wth
rate
(m
m^2
)
Water edge
Non-forested
Forested
Figure 2.12. Growth curve by age (a) and year (b) for water edge (n=32), non-forested (n=60), and forested (n=31) zones derived from
average stem cross-sectional increment.
a. b.
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Figure 2.13. Age structure of Dasiphora fruticosa in ten wetlands ordered from south to
north. Data are presented in five-year intervals. Sampling approach excluded main ramets
<7 years old. Bars represent the number of individual main raments, and lines represent
number of individual secondary branches in each age interval. First number following the
site name is the number of main ramets; second number is the number of secondary
branches.
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Figure 2.14. Number of side branches produced by the main stem in each age class of Dasiphora fruticosa in ten wetlands ordered
from south to north. First number is the number of side branches. Number in parentheses is the number of main stems.
64
64
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Table 2.2. Pearson correlation coefficients (r) for age vs. length, aboveground biomass
(AGB), and stem diameter of D. fruticosa collected from 10 wetlands in Maine.
Age vs. length Age vs. AGB Age vs. diameter
Wetlands r n r n r n
all wetlands 0.53** 134 0.48** 134 0.64** 138
Holt 0.25 16 0.20 16 0.45 16
Dwinal 0.47* 27 0.29 27 0.40** 27
Upper 0.39 12 0.43 12 0.64* 12
Lower 0.50 15 0.26 15 0.45 15
Pickle 0.29 11 0.48 12
Mattagodus 0.24 12 0.58 12 0.68** 13
Salmon 0.60* 12 0.60* 12 0.61* 13
Crystal 0.49 8 0.72* 8 0.66 8
Pillsbury 0.12 14 0.40 14 0.51 14
Soper 0.69* 12 0.86** 12 0.74** 12
Woodland 0.12 11 0.31 11 0.70** 11
Portage 0.19 11 0.14 11 -0.19 120
*p<0.05; **p<0.01
Table 2.3. Pearson correlation coefficients (r) were calculated for age vs. height and
aboveground biomass (AGB) for upright and prostrate growth forms for D. fruticosa
collected from 10 wetlands in Maine.
Growth form r n
Upright shrubs
ABG vs. age 0.48** 117
Height vs. age 0.51** 117
Prostrate shrubs
ABG vs. age 0.36 17
Height vs. age 0.24 17
*p<0.05; **p<0.01
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Table 2.4. Pearson correlation coefficients (r) for age vs. height and age vs. above ground
biomass (AGB) of D. fruticosa collected from three zones (water edge, non-forested
wetland, and forested) in 10 wetlands in Maine.
Position r n
Water edge
AGB vs. age 0.38* 32
Height vs. age 0.51** 32
Non-forested wetland
AGB vs. age 0.54** 66
Height vs. age 0.52** 66
Forested wetland
AGB vs. age 0.57** 36
Height vs. age 0.60** 36
*p<0.05; **p<0.01
Table 2.5. Comparison of growth rates among 10 wetlands. Cell value is the
Gleichläufigkeit (GLK) score, which is the agreement between the trends of the mean
growth curve.
Dwinal Pickle Mattagodus Salmon Crystal Pillsbury Soper Woodland Portage
Holt 0.51 0.44 0.47 0.42 0.40 0.60 0.46 0.43 0.46
Dwinal 0.57 0.54 0.51 0.56 0.61 0.50 0.53 0.67
Pickle 0.36 0.39 0.18 0.57 0.54 0.40 0.40
Mattagodus 0.44 0.46 0.60 0.46 0.38 0.43
Salmon 0.40 0.43 0.54 0.49 0.40
Crystal 0.50 0.44 0.39 0.39
Pillsbury 0.67 0.44 0.58
Soper 0.50 0.44
Woodland 0.39
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2.4. Discussion
As the sole host plant of L. d. claytoni, D. fruticosa provides the primary food
source for both larval and adult life history stages. Quality (e.g., foliar N and moisture
levels) of larval leaf forage and bloom densities differed among wetlands, however,
nectar sugar composition was relatively consistent among and within the wetlands.
Although some butterflies are reported to prefer sucrose-dominant nectars, D. fruticosa
produces hexose-dominant nectar. Dasiphora fruticosa age structure was similar among
wetlands, however, the differences in shrub size and coverage among wetlands indicates
a morphological plasticity to proximate environmental cues.
2.4.1. Resources for larval L .d. claytoni
Lycaena dorcus claytoni spend the majority of their lifetime in juvenile life
stages, making the quality of the larval resources particularly important. While differing
among wetlands, foliar N and moisture levels were not correlated in my study. My
sampling procedure pooled young and mature leaves, which may have confounded
estimates of leaf N available in larval food if L .d. claytoni feed preferentially on new
leaves. Larval insect herbivore growth and survival may increase with foliar N content of
the host plant, although there are exceptions to this trend. For example, Lycaena tityrus
growth varied inconsistently with leaf N of its host plant (Fisher and Fiedler 2000).
Although larval growth and developmental rates of L. tityrus increased with foliar N,
greater pupal mortality and reduced adult size were observed in individuals reared on
food plants with N concentration.
Lepidopteron larvae prefer young, emerging leaves, which may be in response to
decreasing leaf N and moisture concentrations as leaves age (Mattson 1980, Meyer and
Montgomery 1987). Little is known about patterns of host plant use and searching
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behavior of L. d. claytoni larvae, however, larvae were observed on the undersides of D.
fruticosa leaves at the end of branches in areas of new leaf growth (C. Michaud, personal
communication). Several of the sampled wetlands were flooded during the larval period
(Chapter 1). New growth at the tops of the shrubs may provide refuge from flooding as
well as a quality food source. Feeding behavior of L. d. claytoni larvae as well as
relationships among L.d. claytoni life stage, D. fruticosa leaf age, and changes in foliar N
are not well-studied.
2.4.2. Resources of adult L. d. claytoni
Carbohydrate availability can significantly increase longevity, fecundity (Hill and
Pierce 1989, Hill 1989), and flight distance in adult Lepidoptera. Flight is an
energetically expensive activity that may be limited by carbohydrate availability (O’Brien
1999). Dasiphora fruticosa produces small amounts of nectar (VanOverbeke et al 2007).
Adult Clayton’s Copper butterflies do not fly far from their host plants (Layberry et al.
1998), which may reflect limitations of the energy source provided by D. fruticosa.
Concentrations of nectar sugars were greater in enclosed versus unenclosed blooms
indicating nectar consumption by nectivores. Dasiphora fruticosa are pollinated by
Hymenoptera, Diptera, and Hemiptera in addition to Lepidoptera (Guillén et al. 2005,
Elkington and Woodell 1963), however, little is known about competition for nectar
among these species and L .d. claytoni.
Butterflies are able to differentiate between glucose, fructose and sucrose in
nectar, and many studies have demonstrated a preference of sucrose over hexose
(fructose and glucose; Erhardt 1991, 1992, Rusterholz and Erhardt 1997). Dasiphora
fruitcosa produces hexose-dominant nectar. The relative ratio of fructose to glucose in
sampled D. fruticosa blooms was relatively constant among and within wetlands, and
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sucrose was absent in all but a few of the nectar samples. VanOverbeke et al. (2007)
documented the use of D. fruticosa as a nectar source for 10 species of butterflies in the
Jemez Mountains, New Mexico. Although carbohydrate composition was not evaluated
in the New Mexico D. fruticosa, nectar sugar ratios often are relatively constant within a
species (Percival 1961, Baker and Baker 1979, 1982a, b).
Flowers of D. fruticosa have shallow nectaries. Flowers with this morphology
typically produce hexose-rich nectar (Percival 1961), which does not evaporate as readily
as sucrose-rich nectar (Corbet 1979). Hexose-dominate nectar also is less viscous than
sucrose-dominate nectar containing the same weight of sugar (Weast 1980), however,
other non-sugar constituents also may affect nectar viscosity (Heyneman 1983). Nectars
with high viscosity may limit energy intake in butterflies (Baker 1975). Although I
measured sugar concentration per bloom, I did not measure nectar viscosity and sugar
concentration per volume of nectar. Additional information is needed about D. fruticosa
nectar sugar concentration, viscosity, and concentrations of non-sugar solutes such as
amino acids, and relationships with L. d. claytoni nectaring behavior.
Despite differences in chemical structure, sucrose, fructose and glucose contain
roughly the same energetic content per unit gram (16.48 x 103J/g, Weast 1980), and sugar
preferences may not necessarily be correlated to the nutritional value for the butterfly
(Nicolson 2007). Carbohydrate composition may impart a distinct nectar taste or odor
that is recognized by the pollinator (Baker and Baker 1982a). Although D. fruitcosa is
considered the sole adult nectar plant (McCollough et al 2001), I observed L. d. claytoni
nectaring on Solidago uliginosa (Fig. 2.15). Asteraceae typically have hexose dominant
nectar (Percival 1961), suggesting that L. d. claytoni may be able to use other hexose-
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dominant nectar sources. Solidago uliginosa was not abundant in the study wetlands, and
it is not likely to be a primary nectar source for L. d. claytoni.
Figure 2.15. L. d. claytoni nectaring on Solidago uliginosa (Family Asteraceae).
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Flower density as well as nectar availability may affect butterfly foraging
efficiency (May 1985). Foraging rates of Phoebis sennae and Agraulis vanillae in Florida
increased and flight time decreased where bloom availability was greater (May 1985).
Shrubby cinquefoil bloom density varied among wetlands, and wetlands with greater
bloom densities during 2010 generally had greater L. d. clatonyi encounter rates during
2008. Dasiphora fruticosa bloom densities in Maine wetlands were considerably less
than bloom densities of the species in New Mexico (VanOverbeke et al. 2007), with 43%
of the New Mexico shrubs containing more than 50 blooms, whereas, <1% of the Maine
shrubs contained more than 30 blooms. The availability of D. fruticosa shrubs with
abundant blooms may affect wetland site suitability for L. d. claytoni. Investigations of
the adult feeding behavior of L. d. claytoni may prove useful for determining foraging
rate and efficiency and wetland suitability as relates to D. fruticosa bloom availability.
Age-related differences in the quality of shrubs as a food resource may affect
butterfly abundance. Juvenile shrubs typically do not flower or produce seeds (Bond
2000), and bloom vigor may decrease in older shrubs (Oñate and Munné-Bosch 2010),
potentially limiting resources for nectaring butterflies. Wetlands with the greatest L. d.
claytoni encounter rates in 2008 contained primarily intermediate-aged shrubs and the
greatest range in shrub age. Most sampled shrubs were 15-20 years old. Dasiphora
fruticosa regenerates clonally and through recruitment of seedlings (Lent and Reier 1999,
Elkington and Woodell 1963). My sampling methods excluded shrubs < 7 years old, so
seedling recruitment in these wetlands is unknown. Despite differences in size and
coverage (Chapter 1), D. fruticosa age structure and side stem production were similar
among wetlands, and growth rates declined with stem age in all the wetlands. Age-
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dependent patterns of growth have been observed in other woody plants (White 1980). As
multi-stemmed shrubs age, productivity and growth rates decline (Yoda and Suzuki 1993,
Ishii and Takeda 1997; Aikawa and Hori 2006), and more growth is allocated to below
than above ground (Kawamura and Takeda 2008). Age-related changes in D. fruticosa
nectar and leaf quality and effects on food resource availability for L. d. claytoni are
unknown.
Dasiphora fruticosa demonstrates morphological plasticity within wetlands that
reflects environmental conditions (Chapter 1), despite common growth rates among
wetlands. Growth rate was greater at the wetland perimeter (near water and forested
zones) than the center (non-forested) of the wetland. Further, shrub age was correlated
with shrub ABG and height within these zones, although not when age was pooled across
the wetland zones. Sites near open water experience more inundation and dynamic
hydrological conditions, whereas, sites in the forested zone are drier and more shaded.
Dasiphora fruticosa may respond to this variable hydrological environment by increasing
growth rate. The observed range of variability in the phenotypic response of this species
to environmental conditions may provide insight into its tolerance of changes created by
wetland vegetation or water management.
2.4.3. Conservation Implications
Dasiphora fruticosa provides primary larval and adult food resources for L. d.
claytoni, and management strategies that promote conditions favorable for the continuing
regeneration and expansion of D. fruticosa patches are critical for maintaining viable L.
d. claytoni populations. Wetlands are inherently dynamic systems, and D. fruticosa
demonstrates a relatively high level of phenotypic plasticity in these wetlands. The depth
and duration of inundation varied seasonally and among the wetlands where D. fruticosa
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occurs; however, all of the wetlands were saturated within 30 cm of the rooting zone for
the majority of growing season (Chapter 1). Management strategies that provide a
diversity of microclimates and hydrologic conditions may promote D. fruticosa
regeneration while providing flood refugia for the juvenile stages of L. d. claytoni. Age-
related changes in host plant quality may affect butterfly abundance. Therefore, periodic
disturbances that mimic the natural hydrological dynamics may be important for
promoting shrubby cinquefoil reproduction and maintaining robust habitat patches with a
diversity of age classes.
Little is known about the distance that L. d. claytoni is able to travel or if dispersal
occurs among occupied wetlands. Preliminary genetic studies have shown genetic
differentiation among the three geographical regions (central, western, and northern) and
suggest that between region gene flow is limited (C. Michaud, unpublished data),
however, relatedness among wetlands within these regions is unknown. Robust stands of
D. fruticosa currently are restricted to the isolated wetlands surveyed in this study.
Increased connectivity among wetlands containing shrubby cinquefoil may aid dispersal
and increase likelihood of long-term L. d. claytoni population persistence. Additionally,
suitable habitat maintained along waterways connecting occupied wetlands that includes
plant species producing hexose-dominant nectar (such as species in the Asteracae family)
during L. d. claytoni flight period also may enhance L. d. claytoni population persistence.
Page 93
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APPENDIX A. SITE MAPS
Figure A.1. Transect layout at Upper Holt pond, Maine, USA. National Wetland
Inventory (NWI; Maine Office of GIS) Classifications: SS= Scrub-shrub, FO= Forested,
UB=Unconsolidated bottom.
Upper Holt Pond b.
a.
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Figure A.2. Transect layout at Lower Holt Pond Maine, USA. National Wetland
Inventory (NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
UB=Unconsolidated bottom.
Lower Holt Pond b.
a.
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Figure A.3. Transect layout at Upper Dwinal WMA Maine, USA. National Wetland
Inventory (NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
UB=Unconsolidated bottom, EM=Emergent.
Upper Dwinal b.
a.
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Figure A.4. Transect layout at Lower Dwinal WMA, Maine, USA. National Wetland
Inventory (NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
UB=Unconsolidated bottom, EM=Emergent. Shrubby cinquefoil extent is an estimated
edge.
Lower Dwinal b.
a.
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Figure A.5. Transect layout at Pickle Ridge, Maine, USA. National Wetland Inventory
(NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub.
Pickle Ridge b.
a.
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Figure A.6. Transect layout at Mattagodus WMA, Maine, USA. National Wetland
Inventory (NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent.
Mattagodus b.
a.
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Figure A.7. Transect layout at Salmon Stream, Maine, USA. National Wetland Inventory
(NWI) classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent.
Salmon Stream b.
a.
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Figure A.8. Transect layout at Crystal Fen, Maine, USA. National Wetland Inventory
(NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent.
Crystal Fen b.
a.
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Figure A.9. Transect layout at Pillsbury Pond, Maine, USA. National Wetland Inventory
(NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, UB=Unconsolidated
bottom.
Pillsbury Pond b.
a.
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Figure A.10. Transect layout at Soper Pond, Maine, USA. National Wetland Inventory
(NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent, UB=Unconsolidated bottom.
Soper Pond b.
a.
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Figure A.11. Transect layout at Woodland Bog, Maine, USA. National Wetland
Inventory (NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent, UB=Unconsolidated bottom.
Woodland b.
a.
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Figure A.12. Transect layout at Portage Lake, Maine, USA. National Wetland Inventory
(NWI) Classifications (Maine Office of GIS): SS= Scrub-shrub, FO= Forested,
EM=Emergent, UB=Unconsolidated bottom.
Portage Lake b.
a.
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APPENDIX B. HYDROGRAPHS
Figure B.1. Hydrograph of water-table well and deep well for Holt, May 2010 –
September 2010. Hydrograph separated into four time periods: Shrubby cinquefoil leaf
out stage and Clayton’s Copper egg period (12 May -28 May) larval period of the
Clayton’s Copper (29 May – 14 July), shrubby cinquefoil blooming period and Clayton’s
Copper adult fight period (14 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Also plotted is daily rainfall for
sampling period. CC= Clayton’s copper, SC= Shrubby Cinquefoil.
2010
-70
-50
-30
-10
10
300
5/1
2/2
01
0 2
1:0
0:0
0
05
/16
/20
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17
:30
:00
05
/20
/20
10
14
:00
:00
05
/24
/20
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10
:30
:00
05
/28
/20
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07
:19
:48
06
/01
/20
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03
:49
:48
06
/05
/20
10
00
:19
:48
06
/08
/20
10
20
:49
:48
06
/12
/20
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17
:13
:06
06
/16
/20
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13
:43
:06
06
/20
/20
09
10
:13
:06
06
/24
/20
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06
:13
:06
06
/28
/20
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02
:43
:06
07
/01
/20
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23
:35
:41
07
/05
/20
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20
:05
:41
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/09
/20
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16
:05
:41
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/13
/20
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12
:35
:41
07
/17
/20
09
09
:05
:41
07
/21
/20
09
05
:35
:41
07
/25
/20
09
02
:05
:41
07
/28
/20
09
22
:35
:41
08
/01
/20
09
19
:05
:41
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/05
/20
09
15
:35
:41
08
/09
/20
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12
:05
:41
08
/13
/20
09
08
:35
:41
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/17
/20
09
05
:05
:41
08
/21
/20
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01
:35
:41
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/24
/20
09
22
:05
:41
08
/28
/20
09
18
:35
:41
09
/01
/20
09
15
:05
:41
09
/05
/20
09
11
:35
:41
09
/09
/20
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08
:05
:41
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/13
/20
09
04
:55
:13
09
/17
/20
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01
:25
:13
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/20
/20
09
21
:55
:13
09
/24
/20
09
18
:25
:13
Deep wellWater table well
Peat surface
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Figure B.2. Hydrograph of water-table well and deep well for Dwinal, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
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:06
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:43
:06
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:35
:41
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/05
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:05
:41
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:05
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:35
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:05
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/24
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18
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:13
Deep wellWater table well
Peat surface
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Figure B.3. Hydrograph of water-table well and deep well for Pickle monitored May
2010 – September 2010. Hydrograph separated into four time periods: Shrubby
cinquefoil leaf out stage and Clayton’s Copper egg period (12 May -28 May) larval
period of the Clayton’s Copper (29 May – 14 July), shrubby cinquefoil blooming period
and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of
shrubby cinquefoil senescence (20 August – 27 September). Also plotted is daily rainfall
for sampling period. CC= Clayton’s copper, SC= Shrubby Cinquefoil.
2010
-70
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10
300
5/1
2/2
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0 2
1:0
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:00
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/01
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:41
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/05
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20
:05
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/20
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:05
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12
:35
:41
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:05
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:35
:41
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/20
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02
:05
:41
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:35
:41
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/01
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:05
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:35
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18
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:41
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/01
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15
:05
:41
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/05
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11
:35
:41
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/09
/20
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08
:05
:41
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/13
/20
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04
:55
:13
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/17
/20
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01
:25
:13
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/20
/20
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21
:55
:13
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/24
/20
09
18
:25
:13
Deep wellWater table well
Peat surface
Page 117
98
Figure B.4. Hydrograph of water-table well and deep well for Mattagodus, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
998
98
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-70
-50
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-10
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0 2
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21
:30
:00
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/20
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:00
:00
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:48
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/28
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23
:19
:48
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/01
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/06
/20
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00
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:43
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:13
:06
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:06
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/04
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:35
:41
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/08
/20
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03
:35
:41
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:05
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:05
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:05
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:35
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:05
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:05
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:35
:41
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/29
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10
:05
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/02
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:35
:41
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/06
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:05
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/10
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/14
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:25
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/18
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12
:55
:13
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/22
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13
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:13
Deep Water table
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-70
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:19
:48
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/08
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20
:49
:48
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/12
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17
:13
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/16
/20
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13
:43
:06
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/20
/20
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10
:13
:06
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/24
/20
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:13
:06
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/28
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02
:43
:06
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/01
/20
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23
:35
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/05
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20
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16
:05
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/13
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12
:35
:41
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/20
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09
:05
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/21
/20
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05
:35
:41
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/25
/20
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02
:05
:41
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/28
/20
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22
:35
:41
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/01
/20
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19
:05
:41
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/05
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15
:35
:41
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/09
/20
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12
:05
:41
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/13
/20
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08
:35
:41
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/20
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05
:05
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/21
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01
:35
:41
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/20
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22
:05
:41
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/28
/20
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18
:35
:41
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/01
/20
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15
:05
:41
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/05
/20
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11
:35
:41
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/09
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08
:05
:41
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/13
/20
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04
:55
:13
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/17
/20
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01
:25
:13
09
/20
/20
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21
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:13
09
/24
/20
09
18
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:13
Deep wellWater table well
Peat
surface
99
99
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100
Figure B.5. Hydrograph of water-table well and deep well for Crystal, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
100
100
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101
2010
0
1
2
3
4
5
6
5/12
5/26 6/
96/
23 7/7
7/21 8/
48/
18 9/1
9/15
2009
-70
-50
-30
-10
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30
05
/12
/20
10
21
:00
:00
05
/16
/20
10
17
:30
:00
05
/20
/20
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14
:00
:00
05
/24
/20
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10
:30
:00
05
/28
/20
10
07
:19
:48
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/01
/20
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03
:49
:48
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/05
/20
10
00
:19
:48
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/08
/20
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20
:49
:48
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/12
/20
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17
:13
:06
06
/16
/20
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13
:43
:06
06
/20
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10
:13
:06
06
/24
/20
09
06
:13
:06
06
/28
/20
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02
:43
:06
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/01
/20
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23
:35
:41
07
/05
/20
09
20
:05
:41
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/09
/20
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16
:05
:41
07
/13
/20
09
12
:35
:41
07
/17
/20
09
09
:05
:41
07
/21
/20
09
05
:35
:41
07
/25
/20
09
02
:05
:41
07
/28
/20
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22
:35
:41
08
/01
/20
09
19
:05
:41
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/05
/20
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15
:35
:41
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/09
/20
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12
:05
:41
08
/13
/20
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08
:35
:41
08
/17
/20
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05
:05
:41
08
/21
/20
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01
:35
:41
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/20
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22
:05
:41
08
/28
/20
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18
:35
:41
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/01
/20
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15
:05
:41
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/05
/20
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11
:35
:41
09
/09
/20
09
08
:05
:41
09
/13
/20
09
04
:55
:13
09
/17
/20
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01
:25
:13
09
/20
/20
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21
:55
:13
09
/24
/20
09
18
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Deep wellWater table well
Peat surface
101
101
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102
Figure B.6. Hydrograph of water-table well and deep well for Salmon, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
102
102
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103
2010
0
1
2
3
4
5
6
5/12
5/26 6/
96/
23 7/7
7/21 8/
48/
18 9/1
9/15
2009
-70
-50
-30
-10
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30
05
/12
/20
10
21
:00
:00
05
/16
/20
10
17
:30
:00
05
/20
/20
10
14
:00
:00
05
/24
/20
10
10
:30
:00
05
/28
/20
10
07
:19
:48
06
/01
/20
10
03
:49
:48
06
/05
/20
10
00
:19
:48
06
/08
/20
10
20
:49
:48
06
/12
/20
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17
:13
:06
06
/16
/20
09
13
:43
:06
06
/20
/20
09
10
:13
:06
06
/24
/20
09
06
:13
:06
06
/28
/20
09
02
:43
:06
07
/01
/20
09
23
:35
:41
07
/05
/20
09
20
:05
:41
07
/09
/20
09
16
:05
:41
07
/13
/20
09
12
:35
:41
07
/17
/20
09
09
:05
:41
07
/21
/20
09
05
:35
:41
07
/25
/20
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02
:05
:41
07
/28
/20
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22
:35
:41
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/01
/20
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19
:05
:41
08
/05
/20
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15
:35
:41
08
/09
/20
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12
:05
:41
08
/13
/20
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08
:35
:41
08
/17
/20
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05
:05
:41
08
/21
/20
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01
:35
:41
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/24
/20
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22
:05
:41
08
/28
/20
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18
:35
:41
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/01
/20
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15
:05
:41
09
/05
/20
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11
:35
:41
09
/09
/20
09
08
:05
:41
09
/13
/20
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04
:55
:13
09
/17
/20
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01
:25
:13
09
/20
/20
09
21
:55
:13
09
/24
/20
09
18
:25
:13
Deep wellWater table well
Peat surface
103
103
Page 123
104
Figure B.7. Hydrograph of water-table well and deep well for Pillsbury, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
104
104
Page 124
105
2009 2010
-70
-50
-30
-10
10
30
05
/12
/20
10
21
:00
:00
05
/16
/20
10
17
:30
:00
05
/20
/20
10
14
:00
:00
05
/24
/20
10
10
:30
:00
05
/28
/20
10
07
:19
:48
06
/01
/20
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03
:49
:48
06
/05
/20
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00
:19
:48
06
/08
/20
10
20
:49
:48
06
/12
/20
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17
:13
:06
06
/16
/20
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13
:43
:06
06
/20
/20
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10
:13
:06
06
/24
/20
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06
:13
:06
06
/28
/20
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02
:43
:06
07
/01
/20
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23
:35
:41
07
/05
/20
09
20
:05
:41
07
/09
/20
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16
:05
:41
07
/13
/20
09
12
:35
:41
07
/17
/20
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09
:05
:41
07
/21
/20
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05
:35
:41
07
/25
/20
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02
:05
:41
07
/28
/20
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22
:35
:41
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/01
/20
09
19
:05
:41
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/05
/20
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15
:35
:41
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/09
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12
:05
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/13
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08
:35
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05
:05
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/21
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01
:35
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22
:05
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18
:35
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/01
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15
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/05
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11
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/09
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Deep wellWater table well
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Figure B.8. Hydrograph of water-table well and deep well for Soper, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
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2009 2010
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Deep wellWater table well
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Figure B.9. Hydrograph of water-table well and deep well for Portage, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
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2009 2010
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Deep wellWater table wellPeat surface
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Figure B.10. Hydrograph of water-table well and deep well for Woodland, monitored June 2009 – September 2009 and May 2010 –
September 2010. Hydrograph separated into three time periods for 2009: larval period of the Clayton’s Copper (20 June – 22 July),
shrubby cinquefoil blooming period and Clayton’s Copper adult fight period (23 July – 19 August), and at the beginning of shrubby
cinquefoil senescence (20 August – 27 September). Hydrograph separated into four time periods for 2010: Shrubby cinquefoil leaf out
stage and Clayton’s Copper egg period (12 May -28 May) larval period of the Clayton’s Copper (29 May – 14 July), shrubby
cinquefoil blooming period and Clayton’s Copper adult fight period (14 July – 19 August), and at the beginning of shrubby cinquefoil
senescence (20 August – 27 September). Also plotted is daily rainfall for sampling period. CC= Clayton’s copper, SC= Shrubby
Cinquefoil.
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APPENDIX C. COMPONENT LOADINGS
Table C.1. Component loadings of each analyte on the first two principal components
from pore water data collected from 10 wetlands (see Fig. 1.3 -1.5).
Analyte
First principal
component
Second principal
component
1st Sampling period
Nitrate (NO3-N) 0.769 -0.053
Ammonium (NH4-N) 0.409 0.790
Phosphorus (PO4-P) 0.300 0.866
Hydrogen ion (H+) 0.843 -0.375
Electrical conductivity (EC) -0.854 0.265
2nd Sampling period
Nitrate (NO3-N) 0.264 0.702
Ammonium (NH4-N) 0.047 0.788
Phosphorus (PO4-P) 0.737 0.196
Hydrogen ion (H+) -0.848 0.150
Electrical conductivity (EC) 0.831 -0.288
3rd Sampling period
Nitrate (NO3-N) 0.666 0.279
Ammonium (NH4-N) 0.826 0.313
Phosphorus (PO4-P) 0.667 0.194
Hydrogen ion (H+) 0.554 -0.681
Electrical conductivity (EC) -0.224 0.877
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Table C.2. Component loadings of each analyte on the first three principal components
from peat data collected from 10 wetlands (see Fig. 1.6).
Analyte
First principal
component
Second principal
component
Third principal
component
Hydrogen ion [] (H ion) -0.722 0.325 -0.004
Loss on ignition (LOI) 0.658 0.500 -0.225
Nitrate (NO3-N) 0.205 -0.682 0.003
Ammonium (NH4-N) 0.256 0.473 0.127
Calcium (Ca) 0.801 -0.394 0.190
Potassium (K) 0.658 0.325 -0.091
Magnesium (Mg) 0.546 -0.248 0.238
Phosphorus (P) 0.785 0.093 -0.122
Aluminum (Al) -0.805 0.088 -0.044
Copper (Cu) 0.438 0.595 0.106
Iron (Fe) -0.805 0.315 0.107
Manganese (Mn) -0.152 -0.115 0.765
Sodium (Na) -0.130 -0.045 0.565
Sulfur (S) 0.273 -0.066 -0.284
Zinc (Z) 0.404 0.364 0.639
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Table C.3. Component loadings of shrub species on the first three principal components
from peat data collected from 10 wetlands. Only species that occurred in more than 20
sampling sites included.
Species name
First
principal
component
Second principal
component
Third
principal
component
Dasiphora fruticosa -0.345 0.006 0.101
Alnus incana 0.049 -0.122 -0.017
Andromeda polifolia -0.047 0.078 -0.075
Betula pumila 0.388 0.119 -0.381
Chamaedaphne calyculata 0.166 -0.81 0.291
Cornus sericea 0.528 0.329 0.721
Ledum groenlandicum -0.234 0.138 0.077
Lonicera oblogifolia 0.009 -0.15 0.346
Myrica gale 0.678 -0.082 -0.459
Photinia melanocarpa -0.062 0.088 -0.034
Rhamnus alnifolia 0.607 0.04 -0.416
Rosa palustris 0.763 0.334 0.398
Spirea alba 0.313 -0.823 0.077
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APPENDIX D. PORE WATER AND PEAT ANALYTES
Table D.1. Average pore water analytes for 10 wetlands in mid-state, northwestern and
northeastern Maine, 2010. Data are reported for leaf out of the shrubby cinquefoil (SC)
and the egg period of the Clayton’s copper (CC, 24 May- 3 June) Clayton’s copper larval
feeding (12 - 23 July), shrubby cinquefoil bloom and Clayton’s Copper flight (17 – 25
August). ND= No Data.
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Table D.2. Average peat analytes for 10 wetlands in mid-state, northwestern and northeastern Maine, 2010. ND= No Data.
123
116
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APPENDIX E. SITE MAPS WITH DASIPHORA FRUTICOSA SHRUB
VOLUME
Figure E.1. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at (a) Upper and (b) Lower Holt Pond, Maine, USA.
a.
b.
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Figure E.2. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at (a) Upper and (b) Lower Dwinal, Maine, USA. Shrubby cinquefoil extent is
an estimated edge for Lower Dwinal.
a.
b
.
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Figure E.3. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at (a) Pickle Ridge and (b) Mattagodus, Maine, USA.
a.
b
b.
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Figure E.4. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at (a) Salmon Stream and (b) Crystal Fen, Maine, USA.
b.
a.
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Figure E.5. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at Pillsbury Pond Crystal Fen, Maine, USA.
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Figure E.6 Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at Holt Pond, Maine, USA.
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Figure E.7. Dasiphora fruticosa volume (m3) per transect meter at 10m intervals along
transects at (a) Woodland Bog and (b) Portage Lake, Maine, USA.
a
b.
a.
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APPENDIX F. VEGETATION MEASUREMENTS
Table F.1. Abbreviation (Abbrev.) of shrub and tree species names.
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Table F.2. Average shrub volumes (m3) for 10 wetlands.
Table F.3. Basal Area (cm2/transect m) of tree species for 10 wetlands
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BIOGRAPHY OF THE AUTHOR
Sarah Drahovzal was born in Tuscaloosa, Alabama, in 1974. She graduated from
Tates Creek High School in Lexington, Kentucky, in 1992. She attended Wittenberg
University in Springfield, Ohio, where she earned a Bachelors of Art degree in Studio Art
in 1996. She worked for over 9 years at William Russell Pullen Library at Georgia State
University Library in Atlanta, Georgia, while she took post-baccalaureate classes in the
Department of Biology. She is a candidate for the Master of Science degree in Ecology
and Environmental Sciences from the University of Maine in May 2013.