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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

ii

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)

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:

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

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.

iii

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

iv

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.

v

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

vi

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

vii

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

viii

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

ix

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

x

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

xi

Table F.3. Basal Area (cm2/transect m) of tree species for 10 wetlands. ....................... 125

xii

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


samples collected from wetlands in Maine during L. d. claytoni

nectaring (12 – 23 July 2010). ....................................................................... 22


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

xiii

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

xiv

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

xv

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


Figure B.7. Hydrograph of water-table well and deep well for Pillsbury, monitored


Figure B.8. Hydrograph of water-table well and deep well for Soper, monitored


Figure B.9. Hydrograph of water-table well and deep well for Portage, monitored


xvi

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


along transects at (a) upper and (b) lower Dwinal, Maine, USA. ............... 118


along transects at (a) Pickle Ridge and (b) Mattagodus, Maine, USA. ....... 119


along transects at (a) Salmon Stream and (b) Crystal Fen,

Maine, USA. ................................................................................................ 120


along transects at Pillsbury Pond Crystal Fen, Maine, USA. ...................... 121


along transects at Holt Pond, Maine, USA. ................................................. 122


along transects at (a) Woodland Bog and (b) Portage Lake, Maine, USA

..................................................................................................................... 123

1

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

2

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

3

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,

4

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.

5

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

6

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

7

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),

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

9

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.

http://www7.ncdc.noaa.gov/CDO/cdoselect.cmd

10

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.

11

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).

12

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.

13

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.

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


Dwinal 0 5.9 0.52 100 -6.55 4.10 4.6


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


Dwinal 0 6.5 0.73 100 -11.90 3.50 0


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.

15

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 Depth to

Water Table

Flood

durationc

(%) (cm) (±SD) (%) (cm) (±SD) (%)


Dwinal 0 5.98 0.95 100 -6.49 7.01 12.72


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


water table well. b



16

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

17

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


water table well. b



18

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 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


water table well. b



19

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.

20

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).

21

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.

22


collected from wetlands in Maine during L. d. claytoni nectaring (12 – 23 July 2010).




ellipses.

23


collected from wetlands in Maine during D. fruticosa senescence (17 – 25 August 2010).




ellipses.

24

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

25

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).


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


X

X

X

X

X

X

X

X

X

X

24

25

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

27

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.

28

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.

29

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).


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


X

X

X

X

X

X

X

X

X

X

29

30

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.

31

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.

32

Figure 1.8. Shrub coverage (m3/transect m) totaled along transects in 10 wetlands.

33

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

34

33

34

35

Figure 1.9. Tree basal area (cm2/transect m) totaled among transects in sampled wetlands.

Hatched bars indicate unoccupied wetlands.

36

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

37

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

38

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.

39

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.

40

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

41

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.

42

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

43

(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).

44

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

45

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

46

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.

47

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.

48

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

49

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).

50

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)

51

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)

52

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.

53

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).

54

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.

55

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.

56

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).

57

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.

57

57

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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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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.

59

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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

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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

61

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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.

62

62

63

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.

64

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

65

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

66

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


Non-forested wetland

AGB vs. age 0.54** 66


Forested wetland

AGB vs. age 0.57** 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

67

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

68

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

69

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-

70

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).

71

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-

72

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

73

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.

74

REFERENCES

Aikawa, S and Y. Hori. 2006. Effect of a multi-stemmed growth form on matter

production of an understory shrub, Stephanandra incise. Plant Species Biology

21:31–39.

APHA. 1995. American Public Health Association, Standard Methods for the

Examination of Water and Wastewater, 19th

edition. American Public Health

Association, Washington, DC, USA.

Auckland, J.N., D.M. Debinski and W.R. Clark. 2004. Survival, movement, and resource

use of the butterfly Parnassius clodius.Ecological Entomology 29:139–149.

Baker, H.G. 1975. Sugar concentrations in nectars from hummingbird flowers.

Biotropica 7: 37–41.

Baker, H. G. and I. Baker 1979. Sugar ratio in nectars. Phytochemical Bulletin 12: 43–

45.

Baker, H. G. and I. Baker 1982a. Floral nectar sugar constituents in relation to pollinator

types. In C. E. Jones and R. J. Little, eds., Handbook of Experimental Pollination

Biology. New York: Van Nostrand-Reinhold.

Baker, I. and H.G. Baker 1982b. Chemical constituents of nectar in relation to pollination

mechanisms and phylogeny. In M.H. Nitecki, ed., Biochemical Aspects of

Evolutionary Biology, pp. 131-171, Chicago: University of Chicago Press.

Baker, H. G. and I. Baker, 1983a. A brief historical review of the chemistry of floral

nectar. In: Bentley, B. and Elias, T. (eds), The biology of nectaries. Columbia

University Press, pp. 126–152.

Baker, H. G. and I. Baker, 1983b. Floral nectar sugar constituents in relation to pollinator

type. In: Jones C. E. and Little R. J. (eds), Handbook of experimental pollination

biology. Van Nostrand Reinhold, New York, pp. 117-140.

Barros, H. C. H. and F. S. Zucoloto. 1999. Performance and host preference of Ascia

monuste (Lepidoptera, Pieridae). Journal of Insect Physiology 45: 7-14.

Bedford, B., M.R. Walbridge and A. Aldous.1999. Patterns in nutrient availability and

plant diversity of temperate North America wetlands. Ecology 80: 2151-2169.

Bedford, B. L. and K. S. Godwin. 2003. Fens of the United States: Distribution,

characteristics, and scientific connection versus legal isolation. Wetlands 23: 608-

629.

Boggs, C. L., 1997. Reproductive allocation from reserves and income in butterfly

species with differing adult diets. Ecology 78: 181-191.

75

Bond, B.J. 2000. Age-related changes in photosynthesis of woody plants. Trends in Plant

Science 5:349–353.

Boomer, K.M.B. and B. L. Bedford. 2008. Groundwater-Induced Redox-Gradients

Control Soil Properties and Phosphorus Availability across Four Headwater

Wetlands, New York, USA. Biogeochemistry 90:259-274.

Boyer, M.L.H. and B.D. Wheeler. 1989. Vegetation patterns in spring-fed calcareous

fens: calcite precipitation and constraints on fertility. Journal of Ecology 77: 597–

609.

Bragazza, L. L. and R.R. Gerdol. 2002. Are nutrient availability and acidity-alkalinity

gradients related in Sphagnum-dominated peatlands? Journal of Vegetation

Science 13: 473-482.

Bridgham, S.D., J. Pastor, J.A. Janssens, C. Chapin, and T.J. Malterer. 1996. Multiple

limiting gradients in peatlands: A call for a new paradigm. Wetlands 16: 45–65.

Britten, Hugh B. and Lynn Riley. 1994. Nectar Source Diversity as an indicator of

Habitat Suitability for the endangered Uncompahgre Fritillary, Boloria

acrocnema (Nymphalidae). Journal of the Lepidopterists’ Society 48(3):173-179.

Bryant J.P., F.S.I. Chapin, D.R. Klein. 1983. Carbon/nutrient balance of boreal plants in

relation to vertebrate herbivory. Oikos 40: 357-368.

Búrquez A. and S.A. Corbet. 1998 Dynamics of production and exploitation of nectar:

lessons from Impatiens glandulifera Royle. In: Bahadur B, editor. Nectary

biology. Nagpur: Dattsons; pp. 130–152.

Cahenzli, F. and A. Erhardt. 2012. Host plant defense in the larval stage affects feeding

behaviour in adult butterflies. Animal Behaviour 84:995-1000.

Carter, V. 1986. An overview of the hydrologic concerns related to wetlands in the

United States. Canadian Journal of Botany 64:364-374.

Coley, P.D. and J.A. Barone. 1996. Herbivory and plant defenses in tropical forests.

Annual Review of Ecology, Evolution, and Systematics 27:305-335.

Coley, P.D., M.L. Bateman, and T.A. Kursar. 2006. The effects of plant quality on

caterpillar growth and defense against natural enemies. OIKOS 115:219-228.

Corbet, S. A., P. G.Willmer, J. W. L. Beament, D. M. Unwin, and O. E. Prys-Jones.

1979. Post-secretory determinants of sugar concentration in nectar. Plant, Cell and

Environment 2: 293–308.

De Mars, H., M.J. Wassen, and H. Olde Venterink. 1997. Flooding and groundwater

dynamics in fens at Biebrza (Poland). Journal of Vegetation Science 8: 319–328.

76

Dover J.W. 1997. Conservation headlands: effects on butterfly distribution and

behaviour. Agric.Ecosyst. Environ. 63: 31–49.

Eckstein, D. and J. Bauch.1969 .Beitrag zur Rationalisierung eines

dendrochronologischen Verfahrens und zur Analyse seiner Aus-sagesicherheit.

Forstwissenschaftliches Centralblatt 88:230–250.

Ehlers, B.K. and J.M. Olesen. 2004. Flower production in relation to individual plant age

and leaf production among different patches of Corydalis intermedia. Plant

Ecology 174: 71-78.

Ehrlich P.R. and I. Hanski. 2004. On the Wings of Checkerspots: A Model System for

Population Biology. Oxford University Press,Oxford, UK.

Ehrlich, P.R. and P. H. Raven. 1964. Butterflies and Plants: A Study in Coevolution.

Evolution 18: 586-608.

Elkington, T. T. and S.R.J. Woodell. 1963. Potentilla fruticosa L. Journal of Ecology

51:769-781.

Erhardt, A. 1991. Nectar sugar and amino acid preferences of Battus philenor

(Lepidoptera, Papilionidae). Ecological Entomology 16: 425– 434.

Erhardt, A. 1992. Preferences and non-preferences for nectar constituents in Ornithoptera

priamus poseidon (Lepidoptera, Papilionidae). Oecologia (Berlin) 90: 581-585.

Fahrig, L., 2003. Effects of habitat fragmentation on biodiversity. Annual Review of

Ecology, Evolution, and Systematics 34: 487–515.

Fahrig, L., and G. Merriam. 1985. Habitat patch connectivity and population survival.

Ecology 66, 1762–1768.

Fischer, K. and K. Fiedler. 2000. Response of the copper butterfly Lycaena tityrus to

increased leaf nitrogen in natural food plants: evidence against the nitrogen

limitation hypothesis. Oecologia 124: 235-241.

García-Barros, E and T. Fartmann. 2009. Butterfly oviposition: sites, behavior and

modes. In: J. Settele, TG Shreeve, M. Konvicka, and H. Van Dyck (eds) Ecology

of butterflies in Europe. Cambridge University Press. Cambridge, pp. 29-42.

Gilbert L.E. and M.C, Singer.1975. Butterfly ecology. Annual Reviw of Ecology,

Evolution, and Systematics 6:365–397

Grime J. P. 1979. Plant Strategies and Vegetation Processes. John Wiley and Sons, New

York. 222 pp.

77

Guillén, Antonio, Enrique Rico and Santiago Castroviejo. 2005. Reproductive biology of

the Iberian species of Potentilla L. (Rosaceae). Anales del Jardín Botánico de

Madrid 62(1): 9-21.

Hanski, I. 1998. Metapopulation dynamics. Nature 396: 41-49.

Herms D.A., W.J. Mattson. 1992. The dilemma of plants: to grow or defend. Quarterly

Review of Biology 67: 282-335

Heyneman, A.J. 1983. Optimal sugar concentrations of floral nectars: dependence on

sugar intake efficiency and foraging costs. Oecologia 60: 198–213.

Hill, C.J. 1989. The effect of adult diet on the biology of butterflies 2. The common crow

butterfly, Euploea core corinna. Oecologia 81: 258-266

Hill, C.J. and N.E. Pierce. 1989. The effect of adult diet on the biology of butterflies: 1.

The common imperial blue, Jalmenus evagoras. Oecologia 81: 249-257.

Hughes, J. B., G.C. Daily, and P.R. Ehrlich. 2000. Conservation of insect diversity: a

habitat approach. Conservation Biology 14:1788-1797.

Ishii H. and H. Takeda. 1997. Effects of the spatial arrangement of aerial stems and

current-year shoots on the demography and growth of Hydrangea hirta in a light-

limited environment. NewPhytologist 136: 443–453.

Joy J. and A.S. Pulllin. 1999. Field studies on flooding and survival of overwintering

large heath butterfly Coenonymha tullia larvae on Fenn’s and Whixall Mosses in

Shrophire and Wrexham, U.K. Ecological Entomology 24: 426-431.

Kawamura. K. and H. Takeda. 2008. Developmentally programmed and plastic processes

of growth in the multistemmed understory shrub Vaccinium hirtum (Ericaceae).

Botany 86: 268-277.

Keddy, P. A. 2000. Wetland Ecology: Principles and Conservation. Cambridge

University Press, Cambridge, UK. 614pp.

Knurek, E.S. 2010. Taxonomic and population status of the Clayton’s Copper butterfly

(Lycaena dorcas claytoni). (Unpublished thesis) The University of Maine, Orono,

ME.

Kotiaho, J., V. Kaitala, A. Komonen, and J. Päivinen. 2005. Predicting the risk of

extinction from shared ecological characteristics. Proceedings of the National

Academy of Sciences of the United States of America 102:1963–1967.

Kotze, D. J., J. Niemelä, R.B. O'Hara, and H.Turin. 2003.Testing abundance-range size

relationships in European carabid beetles (Coleoptera, Carabidae). Ecography

26: 553–566.

78

Kramer, P. J. 1983. Water Relations of Plants. Academic Press, Orlando, FL.

Krauss, J., I. Steffan-Dewnter, and T. Tscharntke. 2003. Landscape occupancy and local

population size depends on host plant distribution in the butterfly Cupido

minimus. Biological Conservation 120: 355-361.

Krauss, J., I. Steffan-Dewnter, C.B. Müller and T. Tscharntke. 2005. Relative importance

of resource quantity, isolation and habitat quality for landscape distribution of a

monophagous butterfly. Ecography 28: 465-474.

LaBaugh, J. W. 1986. Wetland ecosystem studies from a hydrologic perspective. Water

Resources Bulletin 22:1-10.

Layberry, R.A., P.W. Hall, and J.D. LaFontaine. 1998. The butterflies of Canada.

University of Toronto Press. Toronto. 280pp.

Lent, M. and Ü.Reier. 1999. Origin, chromosome number and reproduction biology of

Potentilla fruticosa (Rosaceae) in Estonia and Latvia. Acta Botanica Fennica 162:

191-196.

León-Cortés, J. L., J. J. Lennon, and C. D. Thomas. 2003. Ecological dynamics of extinct

species in empty habitat networks. 2. The role of host plant dynamics. Oikos. 102:

465-477.

Magee, D.W. and H.E. Ahles. 1999. Flora of the Northeast: a manual of the vascular flora

of New England and adjacent NewYork. University of Massachusetts Press,

Amherst. 1077 pp.

Maine office of GIS.(n.d.) National Wetlands Inventory. In Maine office of GIS Data

Catalog. Retrieved from http://www.maine.gov/megis/catalog/.

Marquis, R.J and H. E. Braker. 1994. Plant-herbivore interactions: diversity, specificity

and impact. In: L.A. McDade, K.S. Bawa, H.A. Hespenheide, G.S. Hartshorn.

(eds.),University Chicago Press. La Selva - Ecology andnatural history of a

Neotropical rain forest. Chicago (USA): University of Chicago. p. 261-281.

Mattson, W. J., 1980. Herbivory in relation to plant nitrogen content. Annual Review of

Ecology, Evolution, and Systematics 11: 119-161.

Mattson W.J., R.A. Haack. 1987. The role of drought in outbreaks of plant-eating insects.

Bioscience 37: 110-118.

May, P.G. 1985. Foraging selectivity in adult butterflies: Morphological, ecological, and

physiological factors affecting flower choice. Ph.D. dissertation, University of

Florida, Gainesville.

79

McCollough, M., M. Siebenmann, and B. I. Swartz. 2001. Clayton’s Copper Assessment.

Maine Department of Inland Fisheries and Wildlife. Wildlife Division. Resource

Assessment Section. Endangered and Threatened Species Program. 1-37.

McLean, E. O.1982. Soil pH and lime requirement. In Page, A. L., R. H. Miller and D. R.

Keeney (eds.) Methods of soil analysis. Part 2 - Chemical and microbiological

properties. (2nd Ed.). Agronomy 9:199-223.

Mevi-Schütz J, Erhardt A. 2005. Amino acids in nectar enhance butterfly fecundity: A

long-awaited link. The American Naturalist 165: 411-419.

Meyer, G. A. and M. E. Montgomery. 1987. Relationships between leaf and the food

quality of cottonwood foliage for the gypsy moth Lymantria dispar. Oecologia

72: 527- 532.

Mitchell, W. A., and H.G. Hughes. 1995. “Line Intercept: Section 6.25, U.S. Army Corps

of Engineers Wildlife Resources Management Manual,” Technical Report EL-95-

22, U.S. Army Engineer waterways Experiment Station, Vicksburg, MS.

Mitsch, W. J. and J. G. Gosselink. 2007. Wetlands 4th

Edition, Hoboken, New Jersey:

John Wiley and Sons, Inc. 582 pp.

Morrant D.S., R. Schumann, and S. Petit. 2009. Field sampling and storing nectar from

flowers with low nectar volumes. Annals of Botany 103: 533–542.

Mortelliti A., G. Amori, and L. Boitani. 2010. The role of habitat quality in fragmented

landscapes: a conceptual overview and prospectus for future research. Oecologia

163: 535-547.

Nicholls, C.N. and A.S. Pullin. 2003. The effects of flooding on survivorship in

overwintering larvae of the large copper butterfly Lycaena dispar batavus

(Lepidoptera: Lycaenidae), and its possible implications for restoration

management. European Journal of Entomology 100: 65-72.

Nicolson, S.W. 2007. Nectar consumers. In: S.W. Nicolson, M. Nepi, and E. Pacini

(eds.), Nectaries and Nectar.

O'Brien, D.M. 1999. Fuel use in flight and its dependence on nectar feeding in the

hawkmoth Amphion floridensis. The Journal of Experimental Biology 202: 441-

451.

Oñate, M. and S. Munné-Bosch. 2010. Loss of flower bud vigour in the Mediterranean

shrub, Cistus albidus L. at advanced developmental stages. Plant Biology 12:

475–483.

80

Pennekamp, F., E. Monteiro, and T. Schmitt. 2013. The larval ecology of the butterfly

Euphydryas desfontainii (Lepidoptera: Nymphalidae) in SW-Portugal: food plant

quantity and quality as main predictors of habitat quality. Journal of Insect

Conservation 17:195-206.

Percival, M.S. 1961. Types of nectar in angiosperms. New Phytologist 60:235 -281.

Rusterholtz, H. and A. Erhardt. 1997. Preferences for nectar sugars in the peacock

butterfly, Inachis io. Ecological Entomology 22: 220–224.

Rydin, H. and J. Jeglum. 2006. The biology of peatlands, Oxford University Press. 343

pp.

Santiago, L.S. and S.S. Mulkey. 2005. Leaf productivity along a precipitation gradient in

lowland Panama: patterns from leaf to ecosystem. Trees 19:349-356.

Schneider, C., J. Dover, and G.L.A. Fry. 2003.Movement of two grassland butterflies in

the same habitat network: the role of adult resources and size of the study area.

Ecological Entomology 28:219–227.

Schultz, C.B. and K.M. Dlugosch. 1999. Nectar and hostplant scarcity limit populations

of an endangered Oregon butterfly. Oecologia 119: 231-238.

Scott, J. A. 1986. The butterflies of North America. Stanford University Press. Stanford,

California. 583 pp.

Severns, P.M., L. Boldt, and S. Villegas. 2006. Conserving a wetland butterfly:

quantifying early lifestage survival through seasonal flooding, adult nectar, and

habitat preference. Journal of Insect Conservation 10:361-370.

Smolders A.J.P., L.P.M Lamers, E.C.H.E.T Lucassen, G. Van der Velde, and J.G.M.

Roelofs. 2006. Internal eutrophication: how it works and what to do about it –a

review. Chemistry and Ecology, 22, 93–111.

Thomas, J.A. 1983. The ecology of Lysandra Bellargys (Lepidoptera: Lycaenidae) in

Britain. Journal of Applied Ecology 20: 59-83.

Thomas, J.A., N. A. D. Bourn, R. T. Clarke, K. E. Stewart, D. J. Simcox, G. S. Pearman,

R. Curtis and B. Goodger. 2001. The quality and isolation of habitat patches both

determine where butterflies persist in fragmented landscapes. Proceedings of the

Royal Society B: Biological Sciences 268: 1791-1796.

United States Department of Agriculture, Agricultural Research Service, Beltsville Area

Germplasm Resources Information Network (GRIN) http://www.ars-grin.gov/cgi-

bin/npgs/html/taxon.pl?413500.

http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?413500

http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?413500

81

VanOverbeke, D. R., P. K. Kleintjes Neff and S. M. Fettig. 2007. Potentilla fruiticosa

(Rosaceae) as a Nectar Plant for Butterflies. Journal of the Lepidopterists’ Society

61: 222-227.

Vitt, D.H. 1990. Growth and production dynamics of boreal mosses over climatic,

chemical and topographic gradients. Bot. J. Linn. Soc. 104: 35-59.

Vitt, D.H., S.E. Bayley, and T. L. Jin. 1995. Seasonal variation in water chemistry over a

bog-rich fen gradient in Continental Western Canada. Can. J. Fish. Aquat. Sci. 52:

587-606.

Wassan, M. J., A Barendregt, A. Palczynski, J.T. De Smidt and H. De Mars. 1990. The

relationship between fen vegetation gradients, groundwater flow and flooding in

an undrained valley mire at Biebrza, Poland. Journal of Ecology 78: 1106-1122.

Weast, R.C., (Ed.) 1980. CRC handbook of chemistry and physics, 60th edn. Boca Raton,

Florida: CRC Press.

Webb, M.R. and A.S. Pullin. 1998. Effects of submergence by winger flooding on

diapusing caterpillars of a wetland buttery, Lycaena dispar batavus. Ecological

Entomology 23: 96 – 99.

Webster, R. P. and B. Swartz. 2006. Population studies of the Clayton’s copper at Dwinal

Pond Wildlife Management Area (Lee/Winn, Penobscot County, Maine). Maine

Department of Inland Fisheries and Wildlife, Bangor, ME 04401.

White, J. 1980. Demographic factors in populations of plants. In: O.T. Solbring (ed.)

Demography and Evolution in Plant Populations. Botanical Monographs, Vol. 15.

Blackwell Scientific Publications, Oxford, pp. 21-48.

Wiklund C. 1977. Oviposition, feeding and spatial separation of breeding and foraging

habitats in a population of Leptidea sinapis (Lepidoptera). Oikos 28:56–68.

Yoda K. and M. Suzuki. 1993. Quantitative analysis of major axis development in

Viburnum dilatatum and V. wrightii (Caprifoliaceae). Journal of Plant Research

106: 187–194.

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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.

83

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.

84

Figure A.3. Transect layout at Upper Dwinal WMA Maine, USA. National Wetland


UB=Unconsolidated bottom, EM=Emergent.

Upper Dwinal b.

a.

85

Figure A.4. Transect layout at Lower Dwinal WMA, Maine, USA. National Wetland


UB=Unconsolidated bottom, EM=Emergent. Shrubby cinquefoil extent is an estimated

edge.

Lower Dwinal b.

a.

86

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


EM=Emergent.

Mattagodus b.

a.

88

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.

89

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.

90

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.

91

Figure A.10. Transect layout at Soper Pond, Maine, USA. National Wetland Inventory


EM=Emergent, UB=Unconsolidated bottom.

Soper Pond b.

a.

92

Figure A.11. Transect layout at Woodland Bog, Maine, USA. National Wetland



Woodland b.

a.

93

Figure A.12. Transect layout at Portage Lake, Maine, USA. National Wetland Inventory



Portage Lake b.

a.

94

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

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Deep wellWater table well

Peat surface

95

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.

95

95

96

0

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96

96

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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

-50

-30

-10

10

300

5/1

2/2

01

0 2

1:0

0:0

0

05

/16

/20

10

17

:30

:00

05

/20

/20

10

14

:00

:00

05

/24

/20

10

10

:30

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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

09

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

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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

08

/05

/20

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15

:35

:41

08

/09

/20

09

12

:05

:41

08

/13

/20

09

08

:35

:41

08

/17

/20

09

05

:05

:41

08

/21

/20

09

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

09

08

:05

:41

09

/13

/20

09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat surface

98

Figure B.4. Hydrograph of water-table well and deep well for Mattagodus, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

998

98

99

-70

-50

-30

-10

10

300

5/1

2/2

01

0 2

1:0

0:0

0

05

/16

/20

10

21

:30

:00

05

/20

/20

10

22

:00

:00

05

/24

/20

10

22

:49

:48

05

/28

/20

10

23

:19

:48

06

/01

/20

10

23

:49

:48

06

/06

/20

10

00

:19

:48

06

/10

/20

09

00

:43

:06

06

/14

/20

09

01

:13

:06

06

/18

/20

09

01

:43

:06

06

/22

/20

09

01

:43

:06

06

/26

/20

09

02

:13

:06

06

/30

/20

09

02

:43

:06

07

/04

/20

09

03

:35

:41

07

/08

/20

09

03

:35

:41

07

/12

/20

09

04

:05

:41

07

/16

/20

09

04

:35

:41

07

/20

/20

09

05

:05

:41

07

/24

/20

09

05

:35

:41

07

/28

/20

09

06

:05

:41

08

/01

/20

09

06

:35

:41

08

/05

/20

09

07

:05

:41

08

/09

/20

09

07

:35

:41

08

/13

/20

09

08

:05

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08

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/20

09

08

:35

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08

/21

/20

09

09

:05

:41

08

/25

/20

09

09

:35

:41

08

/29

/20

09

10

:05

:41

09

/02

/20

09

10

:35

:41

09

/06

/20

09

11

:05

:41

09

/10

/20

09

11

:35

:41

09

/14

/20

09

12

:25

:13

09

/18

/20

09

12

:55

:13

09

/22

/20

09

13

:25

:13

09

/26

/20

09

13

:55

:13

Deep Water table

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

2010 2009

-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

10

03

:49

:48

06

/05

/20

10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

/20

09

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

09

02

:05

:41

07

/28

/20

09

22

:35

:41

08

/01

/20

09

19

:05

:41

08

/05

/20

09

15

:35

:41

08

/09

/20

09

12

:05

:41

08

/13

/20

09

08

:35

:41

08

/17

/20

09

05

:05

:41

08

/21

/20

09

01

:35

:41

08

/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

09

08

:05

:41

09

/13

/20

09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat

surface

99

99

100

Figure B.5. Hydrograph of water-table well and deep well for Crystal, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

100

100

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

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

10

03

:49

:48

06

/05

/20

10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

/20

09

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

09

02

:05

:41

07

/28

/20

09

22

:35

:41

08

/01

/20

09

19

:05

:41

08

/05

/20

09

15

:35

:41

08

/09

/20

09

12

:05

:41

08

/13

/20

09

08

:35

:41

08

/17

/20

09

05

:05

:41

08

/21

/20

09

01

:35

:41

08

/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

09

08

:05

:41

09

/13

/20

09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat surface

101

101

102

Figure B.6. Hydrograph of water-table well and deep well for Salmon, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

102

102

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

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

10

03

:49

:48

06

/05

/20

10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

/20

09

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

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07

/21

/20

09

05

:35

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07

/25

/20

09

02

:05

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07

/28

/20

09

22

:35

:41

08

/01

/20

09

19

:05

:41

08

/05

/20

09

15

:35

:41

08

/09

/20

09

12

:05

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08

/13

/20

09

08

:35

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08

/17

/20

09

05

:05

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08

/21

/20

09

01

:35

:41

08

/24

/20

09

22

:05

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08

/28

/20

09

18

:35

:41

09

/01

/20

09

15

:05

:41

09

/05

/20

09

11

:35

:41

09

/09

/20

09

08

:05

:41

09

/13

/20

09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat surface

103

103

104

Figure B.7. Hydrograph of water-table well and deep well for Pillsbury, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

104

104

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

10

03

:49

:48

06

/05

/20

10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

/20

09

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

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07

/21

/20

09

05

:35

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07

/25

/20

09

02

:05

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07

/28

/20

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22

:35

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08

/01

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09

19

:05

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08

/05

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09

15

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08

/09

/20

09

12

:05

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08

/13

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09

08

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08

/17

/20

09

05

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08

/21

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09

01

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08

/24

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22

:05

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08

/28

/20

09

18

:35

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09

/01

/20

09

15

:05

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09

/05

/20

09

11

:35

:41

09

/09

/20

09

08

:05

:41

09

/13

/20

09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat surface

105

105

106

Figure B.8. Hydrograph of water-table well and deep well for Soper, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

106

106

107

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

10

03

:49

:48

06

/05

/20

10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

/20

09

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

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07

/13

/20

09

12

:35

:41

07

/17

/20

09

09

:05

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07

/21

/20

09

05

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07

/25

/20

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02

:05

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07

/28

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22

:35

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08

/01

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09

19

:05

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08

/05

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15

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08

/09

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12

:05

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08

/13

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09

08

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09

05

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08

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01

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08

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22

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08

/28

/20

09

18

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09

/01

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09

15

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09

/05

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09

11

:35

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09

/09

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09

08

:05

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09

/13

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09

04

:55

:13

09

/17

/20

09

01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13


Peat surface

107

107

108

Figure B.9. Hydrograph of water-table well and deep well for Portage, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

108

108

109

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

10

03

:49

:48

06

/05

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10

00

:19

:48

06

/08

/20

10

20

:49

:48

06

/12

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17

:13

:06

06

/16

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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

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07

/09

/20

09

16

:05

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07

/13

/20

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12

:35

:41

07

/17

/20

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09

:05

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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

08

/05

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09

15

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08

/09

/20

09

12

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08

/13

/20

09

08

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08

/17

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05

:05

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08

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01

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08

/24

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22

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08

/28

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09

18

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09

/01

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09

15

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09

/05

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09

11

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09

/09

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09

08

:05

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09

/13

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09

04

:55

:13

09

/17

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01

:25

:13

09

/20

/20

09

21

:55

:13

09

/24

/20

09

18

:25

:13

Deep wellWater table wellPeat surface

109

109

110

Figure B.10. Hydrograph of water-table well and deep well for Woodland, monitored June 2009 – September 2009 and May 2010 –







Cinquefoil.

110

110

110

111

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

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Peat surface

111

111

112

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


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


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

114

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

115

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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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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transects at (a) Pickle Ridge and (b) Mattagodus, Maine, USA.

a.

b

b.

120


transects at (a) Salmon Stream and (b) Crystal Fen, Maine, USA.

b.

a.

121


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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transects at (a) Woodland Bog and (b) Portage Lake, Maine, USA.

a

b.

a.

124

APPENDIX F. VEGETATION MEASUREMENTS

Table F.1. Abbreviation (Abbrev.) of shrub and tree species names.

125

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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125

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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.

ENVIRONMENTAL ASSESSMENT OF CIRCUMNEUTRAL WETLANDS … · environmental assessment of circumneutral...

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