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Identifier
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Thesis_MES_2020_KlagG
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Title
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Restoration and enhancement of Viola adunca and associated plant species for larval development of Speyeria zerene hippolyta in Pacific Northwest coastal prairie ecosystems using coconut coir mats
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Date
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2020
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Creator
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Klag, Graham
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extracted text
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RESTORATION AND ENHANCEMENT OF VIOLA
ADUNCA AND ASSOCIATED PLANT SPECIES FOR LARVAL DEVELOPMENT
OF SPEYERIA ZERENE HIPPOLYTA IN
PACIFIC NORTHWEST COASTAL PRAIRIE ECOSYSTEMS USING COCONUT
COIR MATS
by
Graham Klag
A Thesis
Submitted in partial fulfillment
of the requirements for the degree
Master of Environmental Studies
The Evergreen State College
December 2020
©2020 by Graham Klag. All rights reserved.
2
This Thesis for the Master of Environmental Studies Degree
by
Graham Klag
has been approved for
The Evergreen State College
by
________________________
Kathleen Saul, Ph. D.
Member of the Faculty
_______________________
Date
ABSTRACT
Restoration and enhancement of Viola adunca and associated plant species for larval
development of Speyeria zerene hippolyta in Pacific Northwest coastal prairie
ecosystems using coconut coir mats
Graham Klag
Across the coastal prairie grasslands of Washington, Oregon and Northern California, the
decline, extirpation and potential extinction of Oregon silverspot butterfly populations,
Speyeria z. hippolyta, are closely associated with the decline in abundance, density and
extirpation of the butterfly’s larval host plant the Early blue violet, (Viola adunca).
Researchers cite the loss of open low nutrient soil conditions as the number one reason for
the violet and butterfly’s decline. The suppression of historic fire regimes, advancement of
forest succession into the prairies, combined with the introduction of livestock and nonnative invasive pasture grasses, play a role in the loss of interstitial space the violet and
butterfly require. These factors occlude the light and space that Viola adunca needs to
grow, establish and recruit, which subsequently outcompetes the plant. The density and
abundance of Viola adunca must be increased to support the butterfly’s survival and
recovery. Utilizing the traditional botanical restoration technique of plug planting to
enhance violet population sizes has proven to be difficult. This research took an innovative
approach using coconut coir mats that provide a growing substrate that mimics the plants’
historic conditions while also suppressing area non-native invasive plants. The research
results, following a Two-way MANOVA test, indicate a statistically significant difference
in native plant aerial cover between planting types (plugs and the vegetative mats) for
October F(3, 416) = 203.39, p ≤ (0.001). The vegetative mats grew Viola adunca, the
associated native plant species and maintained interstitial space more effectively than the
control plug plots.
4
Table of Contents
INTRODUCTION............................................................................................................. 2
Significance ..................................................................................................................... 5
Practical and Theoretical Application ........................................................................... 11
Research Question ......................................................................................................... 12
Hypothesis ..................................................................................................................... 12
Roadmap of Thesis ........................................................................................................ 12
LITERATURE REVIEW .............................................................................................. 14
Insect Conservation ....................................................................................................... 14
Biosphere Reserves - Cascade Head ............................................................................. 15
Historic Coastal Prairie Environment ........................................................................... 18
Indigenous Prairie Practitioners .................................................................................... 19
Viola adunca - Evolution and Biology .......................................................................... 20
Viola adunca - Microsite Conditions ............................................................................ 23
Speyeria zerene hippolyta- Biology .............................................................................. 25
Modern Habitat Restoration Ecology Techniques ........................................................ 33
Speyeria zerene. hippolyta - Larval Survival ................................................................ 36
METHODS ...................................................................................................................... 37
Overview ....................................................................................................................... 37
Research Study Sites ..................................................................................................... 39
Chinook Territory ...................................................................................................... 39
Tillamook Territory ................................................................................................... 41
Alsea Territory ........................................................................................................... 43
Cold Stratified Germination and Sowing ...................................................................... 45
Experiment Design and Site Treatments ....................................................................... 47
Experiment Design of Study Plots ................................................................................ 48
Experimental Design of Treatment / Planting Areas .................................................... 49
Control Plug and Mat Planting ...................................................................................... 52
iv
Monitoring Data and Analysis ...................................................................................... 53
RESULTS ........................................................................................................................ 55
Willapa National Wildlife Refuge................................................................................. 67
Nestucca Bay National Wildlife Refuge ....................................................................... 70
Rock Creek Siuslaw National Forest ............................................................................ 73
DISCUSSION & CONCLUSION.................................................................................. 76
v
List of Figures
Figure 1: Nestucca study site.Vegetative mat at Willapa. Speyeria z. hippolyta..... 1
Figure 2: Keystone species concept visualized through the Ochre Sea Star. .......... 4
Figure 3: Cascade Head Scenic Research Area historic aerial photography ........... 6
Figure 4: Cascade Head 1910 historic fire, and 1930 forest growth ....................... 7
Figure 5: Oregon silverspot butterfly abundance index at extant sites .................... 9
Figure 6: Oregon silverspot butterfly larva being released. Saddle Mt., Oregon. ... 9
Figure 7: Map of Oregon silverspot populations and research site locations ........ 10
Figure 8: The Cascade Head Biosphere Reserve and its endangered animals ..... 16
Figure 9: Viola adunca biological illustration ....................................................... 20
Figure 10: Silverspot pupae being released on Cascade Head 2015. Students...... 26
Figure 11: Speyeria z. hippolyta’s slow growing life cycle. .................................. 27
Figure 12: Cascade Head small mammal trapping 2016. ...................................... 30
Figure 13: Current and historic habitat of Speyeria z. hippolyta with status ........ 31
Figure 14: Habitat map of Speyeria zenrene spp. Current and historic ................. 32
Figure 15: Mating pair of Speyeria zerene hippolyta. ........................................... 39
Figure 16: Soil removal and Viola adunca planting at Willapa Refuge. ............... 39
Figure 17: Map of Tarlatt at Willapa Restoration Area. ........................................ 40
Figure 18: Restored extant habitat for Speyeria z. hippolyta at Nestucca Bay. ..... 41
Figure 19: Map of Area 3 at South Nestucca Restoration Area. ........................... 42
Figure 20: Speyeria z. hippolyta. habitat Rock Creek Siuslaw National Forest .... 43
Figure 21: Map of Area 8 at Rock Creek Restoration Area. ................................. 44
Figure 22: First mat Viola adunca to flower in trial media test at SCCC. ............. 45
Figure 23: Viola adunca seed direct sow.12.13.18 at SCCC. ................................ 46
Figure 24: Viola adunca survival counts in trial media.3.23.2019 at SCCC. ........ 46
Figure 25: Stratified Viola seed, Festuca seed and Fragaria cuttings at SCCC. .. 46
Figure 26: Experimental planting installation design within treatment types. ...... 48
Figure 27: Scraped area planting designs for control plugs and vegetative mats. . 49
Figure 28: Mowed area planting designs for control plugs and vegetative mats... 50
Figure 29: Scraped and mowed treatments. Nestucca National Wildlife Refuge.. 51
Figure 30: Scraped and mowed treatments. Rock Creek Siuslaw National Forest 51
Figure 31: Plug planting process with control Viola adunca plugs. ...................... 52
Figure 32: Vegetative mat and control plant plugs. ............................................... 52
Figure 33: Coastal Prairie Monitoring application at Nestucca. ............................ 54
Figure 34: Monitoring quadrate and height ruler. .................................................. 54
Figure 35: Field monitoring observations .............................................................. 56
Figure 36: June native cover by site and specie. By scraping and vegmat. ........... 57
Figure 37: June native cover by site and specie. By mowing and vegmat. ........... 57
Figure 38: October native cover by site and specie. By scraping and vegmat. ..... 58
vi
Figure 39: October native cover by site and specie. By mowing and vegmat. ...... 58
Figure 40: January plot 5 planted in scraped area with vegetative mat. ................ 59
Figure 41: June plot 5 planted in scraped area with vegetative mat. ..................... 59
Figure 42: January plot 5 scraped area planted with plant plugs. .......................... 60
Figure 43: June plot 5 scraped area planted with plant plugs. ............................... 60
Figure 44: June percent native cover based on treatment and planting type. ........ 61
Figure 45: June percent non-native cover based on treatment and planting type. . 61
Figure 46: June non-native height based on treatment and planting type.............. 62
Figure 47: June relation of treatment and planting type on native plant cover...... 62
Figure 48: October relation of treatment and planting type on native cover. ........ 63
Figure 49: June total Viola adunca cover by site. .................................................. 63
Figure 50: October total Viola adunca cover by site. ............................................ 64
Figure 51: October Viola adunca with vegmat on scraped areas. All sites. .......... 64
Figure 52: October percent native cover based on treatment and planting type. ... 65
Figure 53: October percent non-native cover based on treatment and planting type.
................................................................................................................................ 65
Figure 54: June relation of treatment and planting type on target native plant
cover. ...................................................................................................................... 66
Figure 55: October relation of treatment and planting type on target native plant
cover. ...................................................................................................................... 66
Figure 56: December 2019 vegetative mats with scraping at Willapa. ................. 67
Figure 57: June 2020 vegetative mats with scraping at Willapa. .......................... 67
Figure 58: October 2020 vegetative mats with scraping at Willapa. ..................... 68
Figure 59: December 2019 vegetative mats with mowing at Willapa. .................. 68
Figure 60: June 2020 vegetative mats with mowing at Willapa ............................ 69
Figure 61: October 2020 vegetative mats with mowing at Willapa. ..................... 69
Figure 62: January 2020 vegetative mats with scraping at Nestucca. ................... 70
Figure 63: June 2020 vegetative mats with scraping at Nestucca. ........................ 70
Figure 64: October 2020 vegetative mats with scraping at Nestucca. ................... 71
Figure 65: January 2020 vegetative mats with mowing at Nestucca. .................... 71
Figure 66: June 2020 vegetative mats with mowing at Nestucca. ......................... 72
Figure 67: October 2020 vegetative mats with mowing at Nestucca. ................... 72
Figure 68: January 2020 vegetative mats with scraping at Rock Creek. ............... 73
Figure 69: June 2020 vegetative mats with scraping at Rock Creek. .................... 73
Figure 70: October 2020 vegetative mats with scraping at Rock Creek................ 74
Figure 71: January 2020 vegetative mats with mowing at Rock Creek. ............... 74
Figure 72: June 2020 vegetative mats with mowing at Rock Creek...................... 75
Figure 73: June 2020 vegetative mats with mowing at Rock Creek...................... 75
Figure 74: June target native cover by planting type. ............................................ 77
Figure 75: October target native cover by planting type. ...................................... 77
vii
Figure 76: Willapa October planted Viola adunca cover. ..................................... 80
Figure 77: Nestucca October planted Viola adunca cover. ................................... 80
Figure 78: Rock Creek October planted Viola adunca cover ................................ 81
Figure 79: Link chart of June native plant species cover and association with
planting type........................................................................................................... 81
Figure 80: Link chart of October native plant species cover and association with
planting type........................................................................................................... 82
Figure 81: Viola adunca ecosystem. The role of structure and plant associations
for insects. .............................................................................................................. 83
Figure 82: Viola adunca mat ecosystem. The role of structure and plant
84
associations for insects. ........................................................................................ 834
Figure 83: Speyeria z. hippolyta ecosystem. Etching, 2014. ................................. 85
viii
List of Tables
Table 1: Seed weight and sowing rates for target plant species ................................ 46
Table 2: Explanatory and response variables of research.......................................... 47
Table 3: Treatment and planting area size and dates ................................................. 47
ix
Acknowledgements
My gratitude to The Evergreen State College Master of Environmental Studies Program,
faculty and my reader Kathleen Saul who believes in creative solutions. To the incarcerated
people and Stafford Creek Corrections Center staff who stewarded the project’s plants from
start into success. Thank you to the MES Thesis Fund, the Sustainability in Prisons Project
and the Siuslaw National Forest for helping to finance and facilitate the project and
fostering people to do good work with plants. To Doug Glavich, Matt Smith, Casey Bruner,
Deanna Williams, Michelle Dragoo, Kami Ellingson, Bill Medlen, the Oregon Silverspot
Working Group and others whose botanical backbone and knowledge helped guide this
project’s outcomes. To William Ritchie and Rebecca Chuck of the US Fish and Wildlife
Service who permitted and aided me in the installation of the project, and to Anne Walker
who authorized the use of precious Viola adunca seed for this research. To the Oregon Zoo
for their continued animal husbandry of Pacific Northwest animals in peril. To my loving
parents Scott and Elaine, my sister Kendra and partner Maura who support me in our
journey towards new knowledge.
x
xi
Figure 1: Nestucca study site. Vegetative mat at Willapa. Speyeria zerene hippolyta.
1
INTRODUCTION
In order to properly introduce this environmental study and fully communicate the research
project’s aim and future implications, I must respect and acknowledge the Alsea,
Tillamook, Chinook and Squaxin people. Without their lands and sovereign land-use
practices, I would never have been able to craft and share this story. Without the gift of
their time and territories, we might never have known the Oregon silverspot butterfly
(Speyeria zerene hippolyta) and its once ubiquitous host plant the Early blue violet (Viola
adunca) (see Figure 1). I myself am not indigenous, and through this work I can add to the
growing gratitude for ancestral embeddedness in ecology. Our cultural landscape has been,
can be, and will be either an incubator for collaboration, diversification and speciation, or
the driver of monoculture, instability and extinction debt1 (Dunn, 2005; Samways, 2020;
Shuey et al., 2016). The loss of Viola adunca represents the loss of indigenous burning
practices that held back the succession of Salal (Gaulthoria shallon), Red Alder (Alnus
rubra), and Sitka Spruce (Picea sitchensis). Speyeria z. hippolyta, like other Speyeria
zerene subspecies, has evolved a close host-plant relationship with Viola adunca (Hill et
al., 2018; Sims, 2017). Viola adunca leaves have high concentrations of nutrients such as
vitamin C that the caterpillar of the butterfly depends on for nutrition to complete its slow
growing life cycle and history (Bierzychudek & Warner, 2015; Hill et al., 2018). In the
best reaches and historic habitat that hosted Speyeria z. hippolyta, Viola adunca densities
can be as much as 100 plants per square meter (McCorkle et al., 1980; Schaeffer, 1992;
Kiser, 1993). Part of this life history depended on human intervention by the native prairie
1
Extinction debt - In ecology, extinction debt is the future extinction of species due to events in the past.
The phrases dead clade walking and survival without recovery express the same idea.
2
dwellers in time immemorial (Schultz et al., 2011; Walsh et al., 2010; Zald, 2009). The
practice of periodically burning the prairies for vegetation management, hunting, trade and
ceremonial purposes also facilitated the ecological needs of Speyeria z. hippolyta and Viola
adunca. Today, the habitat needs of these two species is maintained largely by the passive
environmental forces of slope, aspect, and wind-driven salt spray, and the active forces of
concerned citizens who facilitate restoration ecology (Kaye et al., 2015). The pragmatic
land use practices of indigenous people provide a logical relational framework by which to
better visualize our cultural and ecological landscape: the concept of an ecosystem2
(Tansley, 1935). The impacts humans can provide to biological proliferation from
intermediate disturbance and productivity3 can be observed in example keystone species4
such as the Ochre Sea star (Pisaster ochraceus) (See Figure 2). Here the animal’s predatory
impacts on mussels in the dynamic intertidal area actually leads to greater biological
diversity (Yong, 2013). Might we be able to see humans as a keystone species to the coastal
prairie ecosystem, missing and now returning? Through this research, I planted and
disturbed the earth to share an ancestral idea that can become renewed again as we’ll look
to the future: traditional ecological knowledge (Shelvey & Boyd, 2000; Wilkinson, 2010).
2
Ecosystem - The whole system (in the sense of physics), including not only the organism-complex, but also
the whole complex of physical factors forming what we call the environment of the biome — the habitat
factors in the widest sense. It is the systems so formed which, from the point of view of the ecologist, are the
basic units of nature on the face of the earth. These ecosystems, as we may call them, are of the most various
kinds and sizes. They form one category of the multi-tudinous physical systems of the universe, which range
from the universe as a whole down to the atom (Tansley, 1935, p. 96).
3
The intermediate disturbance hypothesis - which proposes that biodiversity peaks at intermediate levels of
disturbance, is often extended to predict that productivity follows the same response pattern.
4
Keystone species - a species on which other species in an ecosystem largely depend, such that if it were
removed the ecosystem would change drastically.
3
Figure 2: Keystone species concept visualized through the Ochre Sea Star. From: Mayne Island
Conservancy, Ochre Sea Star (2020).
The proliferation of Speyeria zerene spp., such as Speyeria z. hippolyta, throughout the
Pacific Northwest depended on a maintenance regime of fire, climatic and other
environmental conditions that maintained an abundance of early seral stage prairies (Walsh
et al., 2010; Zald, 2009). These dynamically disturbed environments allowed for an
abundance of light and space that Viola adunca, Speyeria z. hippolyta and other plants and
animals needed. The interspecies genocide of the 18th and 19th centuries in the Pacific
Northwest followed the appropriation of land and subsequent invasion of settlers that left
in its wake an extinction debt (Gould & Plew, 1996; Shelvey & Boyd, 2000; Wilkinson,
2010). The disappearance of the Alsea, Tillamook, Chinook and other indigenous denizens
of the region removed the traditional dynamic disturbance regimes that led to creation of
4
suitable habitat for Viola adunca and Speyeria z. hippolyta. In these traditional human
habitats, tribes traveled the land they helped to shape in a collective and collaborative
pragmatic land use regime, that may have been able to last into eternity (Hamman,
Dunwiddie, Nuckols, & McKinley, 2011; Schultz et al., 2011; Shelvey & Boyd, 2000;
Stanley et al., 2011; Walsh et al., 2010; Wilkinson, 2010; Zald, 2009). We have an
opportunity to develop projects of reciprocity through restoration ecology for the
rehabilitation of this habitat with the help of people, plants and a specie in rehabilitation
(Kaye et al., 2015). To know and embrace the human gifts we still can give, to reform a
relationship value of survival (Oren Lyons, Indigenous voice).
Significance
Grasslands are endangered ecosystems (Hughes et al., 2000; Kocher & Williams, 2000;
Schultz et al., 2011; Sims, 2017). North American native grassland ecosystems have
experienced species extirpation and decline in richness over the past century due to
agricultural conversion, fire suppression and development, with the loss of these
ecosystems as high as 99% (Noss et al., 1995; Hixon et al., 2010). Today, the coastal
prairie grassland is rare and in rapid decline (Ceballos et al., 2010) (see Figures 3 and 4).
Based on historic aerial photography from 1952 the map below highlights some of the
modern losses of the coastal prairie of one of the butterfly’s and violet’s former strongholds
the Oregon coastal headland; Cascade Head.
In 2015, the Siuslaw National Forest
contracted with me to conduct a series of community workshops with the help of area high
school students to generate a restoration design solution for the recovery of Speyeria z.
hippolyta within the Cascade Head Scenic Research Area: The Cascade Head Coastal
5
Prairie Charrette.5 Following a summer of experience working with scientists, Forest
Service biologists, and graduate students, the high school students gained an understanding
of the coastal prairie ecosystem that was then shared back with the community through the
workshop series. This research builds on the results of the workshop series and goals set
by the community—to come up with a non-herbicidal solution for the restoration and
enhancement of Viola adunca for the larval development of Speyeria z. hippolyta. This
research represents some of the voices, ideas and experiences of the denizens of the
Cascade Head Scenic Research Area and how they see the future of recovery for Speyeria
z. hippolyta in their community.
Figure 3: Cascade Head Scenic Research Area overlay of historic aerial photography. Maps with current
satellite imagery used to delineate the historic prairie extent.
5
Charrette - a meeting in which all stakeholders in a project attempt to resolve conflicts and map solutions.
6
Figure 4: Cascade Head Scenic Research Area fire. 1910 historic grass mountain fire, and 1930 forest growth.
From: Tooze Family/Frank Boyden (2015)
The Cascade Head Coastal Prairie Charrette referenced some of the contemporary coastal
prairie restoration in the Pacific Northwest that I will also explore through the literature
review. Research points to many tested techniques that could provide valuable
contributions to restoring these remnant prairie grassland ecosystems for species, such as
Viola adunca and Speyeria z. hippolyta, by improving the size and quality of habitat at
scale (Hughes et al., 2000; Petix et al., 2018). Topsoil removal, herbicide application and
prescribed fire all have costs and benefits (Jones et al., 2010; Russell & Schultz, 2010;
Sivakoff et al., 2016). In some cases, restoration treatments themselves can affect host plant
quality or survival (Awmack & Leather, 2002). This research aims to tackle some of these
restoration dilemmas and provide solutions for host plant quality and survival. The growing
global body of research on insect conservation points increasingly to the precautionary
principle and the need for the protection, restoration and enhancements of private and
public lands still available to buffer the unknown and unforeseen variables of climate
change (Bauerfeind & Fischer, 2013; Menéndez et al., 2007; Samways, 2020; Thorpe &
Stanley, 2011). In 1980, the Oregon silverspot butterfly (Speyeria zerene hippolyta), was
7
federally listed as a threatened species due to the loss of the butterfly’s host plant Viola
adunca (Speyeria zerene hippolyta W.H. Edwards; USFWS 2001) (see Figure 5). Today,
the caterpillar of the butterfly still depends on the leaves of Viola adunca as its staple food
source during its 6 stages of larval development (Crone et al., 2007) (see Figure 6).
However, after decades of Viola adunca depletion and difficulties with plant
reestablishment and quality, innovation is needed to address the index counts of this
increasingly threatened and likely to now be endangered species (USFWS staff discussion).
In fact, according to the Oregon silverspot butterfly recovery plan, “the butterfly has only
four populations left in the world, with three in Oregon and a small disjunct population in
northern California” (See Figure 7) (Speyeria zerene hippolyta W.H. Edwards; USFWS
2001). Coastal prairie ecosystems like Mt. Hebo and Rock Creek, which still provide
habitat for both the violet and the butterfly include shallow and rocky soils, that support
assemblages of sparse, low growing native plants adapted to low nutrient growing
conditions (McCorkle et al, 1980). These are the ideal conditions for Viola adunca
germination, establishment and recruitment (Almasi & Kollmann, 2007), as well as for
other nectar source species preferred by Speyeria z. hippolyta, like Canadian goldernrod
(Solidago elongate). Sadly, most of the prairies along the coast have lost these qualities,
due to landscape level conversion to non-native pasture grasses for and by livestock (Petix
et al., 2018). In order to address these issues and add to the ecological restoration toolkit,
there is a need for research that explores horticultural methodology and restoration
techniques that achieve rapid restoration targets at less cost while also taking into account
micro and macro site conditions including roads and other obstacles to insects (Littlejohn,
2012; Petix et al., 2018; Shuey et al., 2016; Stanley et al., 2008).
8
Figure 5: Oregon silverspot butterfly abundance index at extant sites in Oregon and California 1990 to 2018.
From A. Walker, pers. comm. (2018).
Figure 6: Oregon silverspot butterfly larva being released into suitable Viola adunca habitat. Saddle
Mountain, Oregon. From: Oregon Zoo (2018).
9
Figure 7: Map of Oregon silverspot populations and research site locations. Source map from: Andersen et
al., (2010).
10
Practical and Theoretical Application
Federal, state and non-governmental organizations such as the Xerces Society agree that
invertebrate and pollinator conservation is an important aspect for not only the agricultural
production of the United States but also for understanding how invertebrates act as
indicator species reflecting the health of our habitats, both human and non-human (Thorpe
& Stanley 2011). This research will present a creative solution that addresses a specific
species’ life history needs, while also taking into account some of the structural diversity
and ecosystem complexity required for functioning ecosystems (Haaland et al., 2011;
Hughes et al., 2000; Petix et al., 2018; Thorpe & Stanley, 2011). By creating ecological
functionality both structurally and biologically for Viola adunca and Speyeria z. hippolyta
within coconut coir6 mats, this research will showcase how restoration ecology can work
to make localized structural changes which can lead towards larger global conservation
goals (Foster et al., 2007; Glaeser & Schultz, 2014; Samways, 2020). The enhancement of
new and former habitat areas can build on this work (Maiti & Maiti, 2015). In areas where
habitat does not exist—rooftop gardens, roadsides, plazas—the mats can create habitats for
insects and humans, proximal to places of development and busy human existence and
activity (Haaland et al., 2011; Littlejohn, 2012). These plant building products can also
lead to the growth of industries in restoration ecology and emerging markets such as land
systems architecture (Systems & Interdisciplinary, 2007).
6
Coconut coir - coir, or coconut fibre, is a natural fibre extracted from the outer husk of coconut.
11
Research Question
Will out-grown and out-planted vegetative mats of Viola adunca, Festuca romeri, and
Fragaria chiloensis made from coconut coir and Alnus rubra chips promote Viola adunca
and associated plant species ability to establish and maintain interstitial spacing within the
mat more effectively than the traditional plug planting techniques?
Hypothesis
This research project will take an innovative non-herbicidal approach to ecological
restoration by utilizing a common environmental engineering and soil conservation tool coconut coir mats ( Abad et al., 2002; Maiti & Maiti, 2015). I hypothesize that using the
mats as a medium to grow the Viola adunca will create an environment suitable to the
violet’s historical habitat, while inhospitable to non-native invasive plants and their seed
above and within the soil. If the vegetative mats are more effective than traditional plug
planting, they will provide a new restoration tool to support the threatened Speyeria z.
hippolyta larval development by suppressing non-native invasive plants, retaining soil
moisture and creating a low nutrient growing media and recruitment substrate for Violas.
Roadmap of Thesis
My thesis research problem will explore the primary literature available to frame my
research question within the contemporary context of restoration ecology and the known
historic ecological conditions of the Viola adunca and the Speyeria z. hippolyta ecosystem.
This thesis begins by exploring the global-to-local implications of insect conservation and
12
the invaluable role it can play in shaping a human value system of true sustainability the
concept of the ecosystem (Tansley, 1935). It will then explore the Cascade Head Biosphere
Reserve and how the Biosphere Reserve concept and designation can help to communicate
and achieve habitat for insects and human conservation efforts. After an exploration of the
future of the concept of sustainability, the discussion will turn back to the ecological
baseline of the historic coastal prairie--what it must have looked like and how indigenous
peoples saw their land through the lens of traditional ecological knowledge. The concept
of sustainability was once known by the indigenous and their territories, it is up to this
thesis and future research to continue the eclipsing of western science with this knowledge.
This thesis taps into this knowledge, exploring the biology and ecology of Speyeria z.
hippolyta and its once abundant host plant Viola adunca while confronting some of the
hurdles, limitations and successes for the restoration of the coastal prairie and the recovery
of Speyeria z. hippolyta (Crone et al., 2007; Petix et al., 2018; Russell & Schultz, 2010;
Service, 2013; Stanley et al., 2008). Based on insect conservation work, this thesis engages
with research on a larval level to model the exploration and herbivory of the caterpillar’s
early instars and what a meter square of high-quality habitat could and should look like.
This project builds off prior research ideas and professional experiences in restoration
ecology and horticulture, growing emergent vegetative mats of various native Carex and
Juncus species, with coconut coir in an aquaponics7 system with the Sustainability in
Prisons Project (SPP) for the Oregon Spotted Frog (Rana pretiosa) at Stafford Creek
Corrections Center in Aberdeen, Washington. This thesis aims to achieve this high-quality
7
Aquaponics - is a combination of aquaculture, which is growing fish and other aquatic animals, and
hydroponics which is growing plants without soil.
13
low growing larval habitat in the coconut coir mats to add a new tool into the coastal prairie
restoration toolkit (Glaeser & Schultz, 2014; Petix et al., 2018; Stanley et al., 2011).
LITERATURE REVIEW
Insect Conservation
Insects along with other invertebrates, play a vital role in many terrestrial ecosystem
processes. Conserving insects is conserving what they do as much as conserving
them for their own sakes, yet the task is vast and fraught with both challenges and
opportunities we will explore here. (Samways, 2020, p. Half title).
Undoubtably, we face dilemmas in assigning value to the conservation of insects, while
solitary insects such as Speyeria z. hippolyta are aesthetically beautiful and visually
elusive, ants invade our homes and eat food while displaying elaborate behavioral
cooperation (Pearce & Wilson, 1990). Specialized plant feeding insects have faced habitat
loss from logging, agriculture, infrastructure development and an increasingly urbanizing
environment for too long (Dunn, 2005). This puts insects at a particularly high risk of
extinction globally (Samways, 2020). Our landscape is fragmented (Schtickzelle et al.,
2007). As habitat corridors continue to shrink in volume and increase in perimeter, it will
continue to drive invasive species movement, and further threatened endemic species
(Dunn, 2005; Littlejohn, 2012; Zielin et al., 2016). Still, researchers debate how much loss
has occurred, as the baseline of ecosystems have shifted rapidly and in some cases without
proper observation or documentation (Samways, 2020). The silent vector forces of exotic
and invasive plants continue to increase pressure on species resources and the habitat
14
structure that buffers extinction debt (Stanley et al., 2011; Samways, 2020). The idea of
land sharing and the concept of biosphere reserves provide one of these global-to-local
opportunities to advance our land systems knowledge and architecture in a way that
protects the animals and insects dwelling there through best management and restoration
practices directed by science (Systems & Interdisciplinary, 2007).
Biosphere Reserves - Cascade Head
The United Nations Educational Scientific and Cultural Organization, UNESCO,
developed the concept of biosphere reserves to bring together ideas of conservation of
species and sustainability. In this global initiative, 714 active biosphere reserves around
the world have been designed around three major areas: 1. Conservation of biodiversity
and cultural diversity; 2. Socio-culturally and environmentally sustainable economic
development; and 3. Logistical support and development through research, monitoring,
education and training (UNESCO 2020, Biosphere Reserves, about). The support structure
of a biosphere reserve helps to facilitate the graded intensity of the land-sharing-landsparing spectrum (Samways, 2020; Van Cuong et al., 2017).
This begins with the core area of a biosphere, focused on conservation of the landscape,
ecosystems, species and humans within by preserving genetic variation within species
(Ishwaran et al., 2008). Unlike early attempts at creating reserves, the biosphere reserves
do not take people out of the picture—humans, plants, and animals live and work together,
not at odds with one another (Bridgewater, 2002). Designated in 1971, The Cascade Head
Biosphere Reserve (see Figure 8) hosts ample opportunities to work with this community
15
ecosystem concept, in an area that through human activities could potentially restore the
resources and habitat for Speyeria z. hippolyta (Speyeria zerene hippolyta W.H. Edwards;
USFWS 2001). Biosphere Reserves offer populations one of the best methods to critically
calibrate value, resources and economic development through a proactive conservation lens
for insects and the humans they serve (Bridgewater, 2002; Samways, 2020).
Figure 8: Map of the Cascade Head Biosphere Reserve and its endangered animals
The growing body of research on coastal prairie restoration in the Pacific Northwest points
to many tested techniques that could provide valuable contributions to restoring these
prairie grassland ecosystems for species such as Speyeria z. hippolyta (Bennett, Thomsen,
& Strauss, 2011; Petix et al., 2018; Russell & Schultz, 2010). By improving the size and
quality of habitat we might begin to see the role we can play and did historically (Awmack
16
& Leather, 2002; Hughes et al., 2000; Zald, 2009). However, many of these tested
techniques have drawbacks. For example, while topsoil removal appears to be one of the
most successful techniques in reestablishing native forb and grass cover, the cost and
feasibility of soil removal continues to be a hurdle for large scale and site-specific
restoration (Petix et al., 2018; Russell & Schultz, 2010; Schultz & Ferguson, 2020; Sims,
2017; Thorpe & Stanley, 2011). Moreover, the most adopted landscape level alterative—
the use of herbicide—has significant stakeholder push back and growing animal and human
health concerns (Gillam, 2017). To address these concerns and add to ecological function
during restoration there is a need for research into treatment combinations and the
innovation of novel techniques. But first, we must understand the dramatic changes to these
coastal prairie ecosystems over time (Schultz et al., 2011; Stanley et al., 2011). These
changes have informed our understanding of the role geographic variability and
disturbance regimes play in the maintenance of plant communities and indicator species
(Foster et al., 2007; Jermy, 1984; Sims, 2017). These include Viola adunca and the
subspecies of Speyeria zerene, whose larva depend on the leaves of the violet. This review
will explore the role of micro and macro site conditions in combination with historic and
modern land use practices that aid in maintaining a seral stage of herbivory for the butterfly
in coastal prairie ecosystems. Framed within the contemporary context of restoration
ecology and the known historic ecological conditions of Viola adunca and Speyeria z.
hippolyta, this literature review will provide the foundation for research into the question:
Will planted native vegetative mats growing Viola adunca, made from coconut coir and
Alnus rubra chips promote Viola adunca’s ability to establish and maintain interstitial
spacing within the mat more effectively than traditional plug planting techniques?
17
Historic Coastal Prairie Environment
To understand how to establish and maintain density and abundances of Viola adunca and
other plant species native to Pacific Northwest coastal prairies, we must first understand
the mechanisms and conditions, both biotic and abiotic, that created the historic prairie and
caused its decline. “One of the most important conservation issues in ecology is the
imperiled state of grassland ecosystems worldwide due to land conversion, desertification,
and the loss of native populations and species” (Ceballos et al., 2010, p. 2). Across the
globe, temperate grasslands and meadows have sharply declined in spatial extent relative
to other grassland types such as savannahs and tropical grasslands (Kruess & Tscharntke,
2002; Sims, 2017; Zald, 2009). Temperate grasslands create hotbeds of primary
production, biodiversity, and carbon storage comparable to that of tropical rainforest, and
can even assist with climate change mitigation by affecting the earth's surface albedo,
deflecting some of the sun's energy which darker colored forests absorb (Foster et al., 2007;
Van Geel et al., 1999; Walsh et al., 2010). In addition to providing ecosystem services like
those just mentioned, these coastal prairies contain rare endemic plant species and disjunct
populations of relict botanical communities (Zald, 2009). Ultimately, the limited
availability and the high level of speciation and numbers of rare species present in prairies
contribute to their societal and ecological importance (Dunn, 2005; Hughes et al., 2000).
Zald (2009) recounts that the current extent and spatial distribution of coastal prairies
relates largely to tree encroachment. Over the past hundred years, in locations such as Mt.
Hebo where the last population of Speyeria z. hippolyta currently thrives, as much as (96%)
of the historic coastal prairie extent has been lost to forest encroachment (Bachelet et al.,
2011; Zald, 2009).
18
Indigenous Prairie Practitioners
Fire severity is a measure of the effects of fire on the environment—both in damage
to vegetation and impacts on the soil. Fire severity is driven by weather conditions,
the topography of the landscape, and the fuels that are present. Of these, weather is
the overriding factor (OSU Extension Service, 2012. Fire severity).
The large and high severity wildfires that created the important disturbance agents in the
coastal prairie environment became considerably disconnected over time. The earth's last
major climatic change occurred approximately 2770 years ago (Van Geel et al., 1999).
During this time the coastal prairies experienced stand replacing fire regimes every 140 to
170 years. The last 2770 years has been what is referred to as the "Little Ice Age" and saw
the frequency of these fire regimes shift to every 240 to 270 years. Indigenous people
thrived as active users shaping their landscape to maintain the coastal prairies by lighting
more frequent and less severe fires (Hamman et al., 2011; Zald, 2009). Indigenous people
incorporated fire as part of ceremonial practice, burning blankets to welcome the salmon
home, to drive game into the open for hunting, as well as to maintain open prairies for ease
of hunting and gathering activities (Walsh et al., 2010). Additionally, these burning
practices helped to maintain prairie plant assemblages such as camas and other species
which served as sources of food and medicine (Gould & Plew, 1996). Over time, the
intermediate disturbance regimes of the indigenous prairie practitioners created an
ecosystem of high biodiversity and abundance. It was an ecosystem where human land use
management and biodiversity co-evolved (Gould & Plew, 1996; Shelvey & Boyd, 2000).
However, this ecosystem ended beginning in the 1860's when European settlement led to
the proliferation of disease and the subsequent genocide of the region's indigenous people
19
and their land use practices (Zald, 2009). European immigrates settled along the coast to
survive by agrarian and natural resource extraction to find stability in a land fraught with
wind, weather and waves.
Figure 9: Viola adunca biological illustration.
Viola adunca - Evolution and Biology
Viola adunca is a perennial angiosperm forb8, with slender rhizomes9 that evolved out of
one of the largest orders of flowering plants; Malpighiales with over 36 families (Judd et
al., 1999).
8
Forb - a herbaceous flowering plant other than a grass.
Rhizomes - a continuously growing horizontal underground stem which puts out lateral shoots and
adventitious roots at intervals.
9
20
Usually stemless in the early part of the season, later developing aerial stems up to
10 cm tall. Starts to flower early in the growing season. Leaves generally oval to
heart-shaped, hairy to hairless, blades to 3 cm long with fine round-toothed
margins. Stipules reddish-brown or with reddish-brown flecks, narrowly lanceshaped margins slender-toothed or somewhat ragged. Flowers to 1.5 cm long, with
a slender spur half as long as the lowest petal, petals blue to deep violet, the lower
three often white at base and purple-penciled, the lateral pair white-bearded
(MacKinnon et al., 2004, p. 201) (see Figure 9).
Viola adunca’s habitat includes dry to moist meadows, open woods, grasslands and open,
disturbed ground from lowlands to near timberline (Johnston et al., 1974). Its blue flower
primarily attracts bees and flies; once pollinated it produces a capsule of seeds. The primary
mechanism for seed dispersal is during the late summer when the fruit’s small capsules
open and three valves explosively propel the seed outward from the capsules as it dries.
This dispersal can broadcast seed up to approximately 50 feet and into new areas where
the seeds can lie dormant for years or can germinate after approximately 100 days of cool
wet weather if conditions are optimal to initiate germination of the seed (Almasi &
Kollmann, 2007; American & Aug, 2016). Viola adunca can flower multiple times per
year, when punctuated events of moisture and sun coincide (American & Aug, 2016;
Freitas & Sazima, 2003). The evolution of the physical, biological and cultural
environmental influences of Pacific Northwest coastal prairies shaped the ecology that we
witness today (Gould & Plew, 1996; Walsh et al., 2010). The presence and absence of
Speyeria z. hippolyta and Viola adunca offer a clear indicator of the current state of the
ecosystem function of the coastal prairie (Hughes et al., 2000; Jermy, 1984; Kruess &
21
Tscharntke, 2002; Schultz et al., 2011; Sims, 2017; Sivakoff et al., 2016). Together these
two species help us visualize the influence of ecological forces on biological form and
function that has disappeared in many areas today (Freitas & Sazima, 2003; Hill et al.,
2018; Shuey et al., 2016). An animal emblematic and intermediary to the plant and animal
community it co-evolved to. While restoration ecology has often focused on the
reintroduction of a single species, little work has been done to examine the ecology of
restoration itself and its overall effect on plant quality (Almasi & Kollmann, 2007). To
better understand the biological needs of the butterfly and its ability to move into new
habitats, we must examine the life and evolutionary history of Viola adunca (Botanical &
Press, 2016).
Violas have evolved to have three different types of pollination syndromes: sternotribic
(where an insect’s stomach pollinates the flower), nototribic (where an insect’s back
pollinates the flower), and self-pollination (where no insect pollinates the flower, but the
violet reproduces asexually). While yellow violets represent the most ancient of violet
species, blue violets evolved more recently. Eighty-nine percent of Viola adunca
pollination is sternotribic; it is pollinated almost exclusively seventy-two percent by
solitary bees Osmia Andrena, also known as mason bees (Botanical & Press, 2016).
This co-evolution between ecosystem conditions, and a single insect genus is mirrored in
other aspects of Viola adunca’s life history. Research into the germination requirements of
the violet has pointed to soil type and temperature regime as important factors in the
germination success of the species (Almasi & Kollmann, 2007). Other research has shown
that the violet responds well to inadvertent fluctuations in temperature, light, and high soil
22
pH levels (James, 2008). Moreover, Viola adunca sown in soil medias of coarser substrate,
greater structural diversity, and cooler temperatures appear to have the greatest influence
on seed germination (Almasi & Kollmann, 2007; Dunwiddie & Martin, 2016; Sparling,
2020). This points to the applicability of this coconut coir research, focusing on the role
that coconut coir mats and alder chips can play in providing a soil/substrate that mimics
disturbed areas with these microsite conditions. This research can begin to examine the
role of soil structure in the violet’s ability to establish, maintain and recruit in the wild
through mat mimicry. An examination of the role of site soils hosting the plant will address
one of the important gaps that exists in developing a greater understanding of the Viola
adunca’s life history (Kubitzki, 2014).
Viola adunca - Microsite Conditions
Within the realm of soil structure, pH and other substrate variables, research within the
inland prairies of the Salish lowlands and Willamette Valley has pointed to the role of
microsite conditions at valley prairie sites like Mima Mounds, where geological history
and topographic variation are associated with promoting the germination and establishment
of a variety of plants including Viola adunca and the Golden Indian paintbrush (Castilleja
levisecta), (Dunwiddie & Martin, 2016). The Dunwiddie and Martin (2016) paper entitled
“Microsites Matter: Improving the Success of Rare Species Reintroductions” examined the
inland prairie sites of Washington and Oregon at different stages of habitat suitability,
restoration and geologic condition. Research into the role of these different variables on
species presence or absence identified certain microsite conditions, such as loamy soil
types and steeper localized topography, as strong indicators of Golden paintbrush
23
(Castilleja levisecta), Viola adunca and Festuca roemeri (Dunwiddie & Martin, 2016).
Areas in the Salish lowlands, such as at Mima Mounds, presented the greatest abundance
of Castilleja levisecta, Viola adunca and Festuca roemeri with the plant species growing
in close association at the base of mounds.
The three plants’ ability to maintain and recruit themselves within these microsites was
associated with low nutrient and mid-disturbance regimes10 due to the angle of repose11 of
mounds and other topography. Moreover, the authors believe the presence of these
microsite conditions in combination with sub surface nutrient rich soils, such as the
Nisqually loams below the low nutrient surface, was beneficial in promoting plant growth
and survival. The findings of the study also point to the importance of the interaction of
site and topography in affecting the survival and recruitment of Castilleja levisecta, Viola
adunca and Festuca roemeri (Dunwiddie & Martin, 2016; Lawrence & Kaye, 2011;
Sparling, 2020). Turning to this thesis research, the microsite conditions of the prairie may
be mirrored in the conditions of the coconut coir mat planted with the Viola adunca and
Festuca roemeri (Almasi & Kollmann, 2007; Dunwiddie & Martin, 2016; Sparling, 2020).
The vegetative mats’ ability to give these slower growing plants such as Viola adunca and
Festuca roemeri a head start to the existing weed community by providing a low nutrient
surface substrate with access to richer soils below, could provide insights into restoration
succession and greater plant establishment success (Dunwiddie & Martin, 2016; Glaeser
& Schultz, 2014; Hill et al., 2018; Stiling & Moon, 2005).
10
Disturbance regimes - Any of various modes of widespread floral replacement, e.g., flood, fire, disease or
wind.
11
Angle of repose - the steepest angle at which a sloping surface formed of a loose material is stable.
24
Speyeria zerene hippolyta- Biology
Coastal conditions such as fog and wind helped to create a darker smaller Speyeria z.
hippolyta along the coast, also informing a smaller body surface area and a faster ability to
thermoregulate in the early morning light along the forest edge (Speyeria zerene hippolyta
W.H. Edwards; USFWS 2001). Darker wings also help absorb the sunshine the butterfly
needs for thermoregulation in the cool summer temperatures of the coastal prairie were the
butterfly thrives. Both Speyeria z. hippolyta and Viola adunca can often be found in
locations where temperatures from the sun’s energy fluctuates (McCorkle et al., 1980;
Schaeffer, 1992; Kiser, 1993). The Northwest coast’s ambient and often strong winds
shaped the wing dimensions of the coastal Speyeria z. hippolyta that are stout and like their
alpine Speyeria cousins. Persistent northwest summer winds are believed to have played a
role for populations of the butterfly moving between the Clatsop Planes and Saddle
Mountain where coast range meadows provide amble Viola adunca habitat today suitable
for larval release (Petix et al., 2018; Service, 2017). This population migration and gene
flow may have also occurred between Mt. Hebo and proximal historic coastal prairie sites
like Nestucca, Netarts and Cape Mears (see Figure 13).
Speyeria z. hippolyta is a highly localize endemic12 which makes it well adapted to rselection13 and disturbance associations (Sims, 2017; Sivakoff et al., 2016). Viola adunca
population sizes can help provide an estimate of the carrying capacity available for the
12
Endemic - native and restricted to a certain place.
R- Selection - species are those that emphasize high growth rates, typically exploit less-crowded
ecological niches, and produce many offspring, each of which has a relatively low probability of surviving
to adulthood.
13
25
Speyeria z. hippolyta selection and movement into new habitats in combination with
adequate nectar sources for adults (Bierzychudek & Warner, 2015; Hill et al., 2018;
Hughes et al., 2000; Jermy, 1984; Sims, 2017). While male butterflies use the open coastal
meadows for courtship, fertile females are seekers of low vegetation height where Viola
adunca patches can persist for oviposition14 on the violet leaves. Once the females’ eggs
have been laid, the first instar15 of the butterfly typically hatches after approximately 22
days when the larva then eats its egg case, typically between June and July (Kiser, 1993,
Schaeffer, 1992) (see Figure 11).
Figure 10: Oregon silverspot pupae being released on Cascade Head 2015. Charrette students visit Mt.
Hebo to learn about pollinators.
14
Oviposition - means expulsion of the egg from the oviduct to the external environment and is a common
phenomenon in invertebrates, vertebrates and other than eutherian mammals.
15
Instar - a phase between two periods of molting in the development of an insect larva or other
invertebrate animal.
26
Figure 11: Speyeria z. hippolyta’s slow growing life cycle. From: Andersen et al., (2010).
The timing of Speyeria z. hippolyta’s life cycle and history are closely linked to the climatic
and elevational conditions along the coast where localized weather patterns, moisture and
temperature are greatly associated with coastal proximity (McCorkle et al., 1980). The
herbicidal effect of salt spray from summer winds seem to have the greatest effect on these
proximal plant communities, stunting the vertical growth of the plant community. Factors
such as vegetation height, effects adult flight behavior, migration and host plant
availability. Additionally, the phenology16 of the plant community that host the butterfly’s
nectar is closely associate with the coastal prairie summer (Kiser, 1993). Key features to
the habitat needs of Speyeria z. hippolyta is in abundance of late-season nectar sources
such as Canadian goldenrod (Solidago elongate), Pearly Everlasting (Anaphalis
margaritacea), and Pacific aster (Symphyotrichum chilense) (Speyeria zerene hippolyta
W.H. Edwards; USFWS 2001). While the butterfly is currently listed as threatened, an
16
Phenology - the study of cyclic and seasonal natural phenomena, especially in relation to climate and
plant and animal life.
27
endangered classification is now warranted, for the specie based on a lack of funding
available to monitor the species population (Discussions with USFWS).
The caterpillars’ dependence on the leaves of the plant to complete its slow growing life
cycle evolved out of the plant’s once nationwide dispersal and nutritional value
(Bierzychudek & Warner, 2015; Hill et al., 2018). Speyeria z. hippolyta was first described
from three males and one female taken in Oregon with an additional male from Northern
California in 1879. These individuals constitute the classification of the subspecie that we
know today to be true Speyeria z. hippolyta. It is important to note that some biologists
believe the Northern California population represents a convergent subspecie that evolved
through a convergent ecotype derived from adjacent inland Speyeria zerene behrensii or
gloriosa (McCorkle et al., 1980). Alternatively, some hypothesize that the population
represents a divergent relict form of hippolyta now separated from the parent population
by an extensive and dynamic dune complex of the Central Oregon coast (see Figure 13).
As McCorkle (1980) describes in ecological detail; it is important to understand that there
are other Speyeria zerene or Oregon silverspot subspecies that live in Oregon, including
Speyeria z. gloriosa of the Illinois River Valley, and the–now extinct from Oregon– the
Valley silverspot Speyeria z. bremnerii, last observed in the 1970s at Mary’s Peak. Another
more widespread and non-endangered member of the Syperia genus can also be found
within the habitats of Speyeria z. hippolyta; Speyeria hydaspe has cream colored spots
under its wings in contrast to the metallic silver spots of Speyeria z. hippolyta.
The subspecies of zerene Fritillaries once radiated throughout the Pacific
Northwest and into California and illustrated the principles of geographic
variability and subspeciation across the region, with six or more Cascadian
28
subspecies currently or formerly occupying localized habitats as distinct as the
subspecies (Pyle, 2002, p.268) (see Figure 14).
Speyeria zerene hippolyta - occupies the coastal and dune prairies of Washington Oregon
and Northern California. Speyeria zerene zerene (or conchyliatus) - occupies the Southern
Oregon Cascades, the Warner, Klamath, Coast Range Mountains of Northern California
and the Sierra Nevada Mountains extending into Southern California. Speyeria zerene
gloriosa - occupies the riparian areas of the Illinois River, the Siskiyou Mountains, the
Southern Oregon Coast from Coos Bay to Gold Beach and continuing in through the
Northern California coast from Arcata and into the Kings Mountain Range. Speyeria zerene
behrensii - occupies the Mendocino range in Northern California. Speyeria zerene
bremnerii - occupies the Sunshine Coast of British Columbia, the Salish Lowlands of
Washington west to Port Angeles east to the Cascades and historically south throughout
the Willamette Valley. Speyeria zerene unnamed subspecies - occupies the north and
northeast subalpine habitats of the Olympic Mountains in Washington (McCorkle et al.,
1980; Pyle, 2002, p. 268).
Speyeria z. hippolyta’s biology is deeply related to its ecology, which is why there are other
animals that can be used as stand indicators of ecological conditions associated with Viola
adunca and Speyeria z. hippolyta. Animals such as the Pacific jumping mouse (Zapus
trinotatus) and other small mammals can be used as stand-level indicators for ecological
health, in absence of the butterfly. In areas such as Cascade Head, where today the butterfly
has been expatriated, its indication is related to the complex variety of interactions that it
benefits from based on the plants, animals and ecological structure present. In contrast, at
29
Cascade Head, insectivorous Vagrant shrews (Sorex vagrans) have been documented in
small mammal research as having a possible association with mowing regimes on present
pasture grasses as they can still hide in the low vegetation, while larger mammals such as
the jumping mouse are vulnerable to predators (Unpublished research, Wilson 2015). The
Deer mouse (Peromyscus maniculatus) is not an insectivore and thus not a predator of the
Speyeria z. hippolyta larvae, but is a predator of Viola adunca seed (Cor& Bury, 1991;
Wilson & Forsman, 2013).
Figure 12: Pacific jumping mouse (Zapus trinotatus). Vagrant shrew with ticks (Sorex vagrans). Deer
mouse (Peromyscus maniculatus). Cascade Head small mammal trapping 2016.
30
Figure 13: Current and historic habitat map of Speyeria z. hippolyta with locational status.
31
Figure 14: Habitat map of Speyeria zenrene spp. Current and historic in the Pacific Northwest and
California.
32
Modern Habitat Restoration Ecology Techniques
A wealth of research has been conducted on the California coastal prairie complex (Bennett
et al., 2011; Thorpe & Stanley, 2011). While climatically and geologically different than
the coastal prairie ecosystems of Oregon and Washington, there is much we can learn from
this regional research. For example, researchers have examined the interaction between a
native California forb, Seaside fleabane (Erigeron glaucus), and a non-native invasive
grass, Velvet grass (Holcus lanatus), a species that is also of non-native invasive concern
in Pacific Northwest coastal prairie ecosystems. Bennett et al. (2011) found the weeding
of plots to have the most competitive interaction effect on Holcus lanatus, while also
increasing seed bank germination of Erigeron glaucus. The research examined how the
presence and persistence of Holcus lanatus changes the soil community over time, while
also directly competing with the native plant community and the effects of herbivory17 on
the plants (Bennett et al., 2011). They also explored Holcus lanatus competition via
allelopathy.18 Weeded areas led to high rates of germination success for Erigeron glaucus.
Germination rates increased up to (815%) as compared to unweeded plots. Direct
competition of Holcus lanatus and Erigeron glaucus influenced the total survival of
Erigeron glaucus. Mammalian herbivory also played a role in the reduction of Erigeron
glaucus plant size by (71%). Perhaps the most important finding from the study was that
the removal of competitors without the alteration of seed bank resulted in the failure of
native species to re-establish. This points to the role that soil scraping in combination with
17
Herbivory - the state or condition of feeding on plants.
Allelopathy - the chemical inhibition of one plant (or other organism) by another, due to the release into
the environment of substances acting as germination or growth inhibitors.
18
33
vegetative (coconut coir) mats may play in altering the seed legacy of soil monocultures
following degradation and non-native invasion (Bennett, Thomsen, & Strauss, 2011). The
results of the research illustrate the multitude of mechanisms that drive native and nonnative invasive plant interactions in the coastal prairie ecosystem.
In Buisson et al. (2006), research dealing with the California coastal prairies explored the
most appropriate combination of treatments for reintroducing the coastal prairie
bunchgrass California oatgrass (Danthonia californica). In complement to the Holcus
lanatus and Erigeron glaucus study, this research examined the impact of local versus nonlocal seed sources on the long-term survivability of the native bunch grass (Buisson et al.,
2006). The study also examined the misleading nature of short-term plant establishment
studies (research that occurs with the monitoring of one growing season) and the value of
genetically localized seed sources. Plant genetics were of value to the study to trace the
success of local seed sources while also examining restoration treatments, determining that
topsoil removal greatly enhanced both transplanted and seeded Danthonia californica.
Even longer-term research has examined the role of continual disturbance regimes such as
of grazing on Danthonia californica biomass. Grazing was noted to increase the root
growth of Danthonia californica the but also a decrease in the overall plant biomass, which
has also been identified in the Pacific Northwest and other conservation areas (Kruess &
Tscharntke, 2002; Schultz et al., 2011; Dunwiddie et al., 2008). From this California study,
one can conclude that future studies should take a long-term approach to monitoring to
account for restoration success, even in the dynamic and rapidly changing forb
communities of the coastal prairie. The study highlights the beneficial role of local seed
sources, transplanting versus seeding and most importantly the role soil scraping can play
34
in the success of establishing other native species found in the coastal prairie complex with
species such as Viola adunca (Buisson et al., 2006, Thorpe & Stanley, 2011).
Northward to the Pacific Northwest coastal prairie, the most recent restoration efforts for
the violet and butterfly reflect a variety of projects designed to try to re-establish quality
habitat for Speyeria z. hippolyta. Of great significance to this coastal prairie restoration
research is a series of research projects conducted in the Clatsop Plains on the northern
Oregon and southern Washington coast through partnerships with the Institute for Applied
Ecology and the United States Fish and Wildlife Service, which focused on the effect of
scraping and topsoil removal in the restoration treatments of coastal prairie plant
communities at Tarlatt, Willapa (incorporated into this research project) and the Clatsop
Plaines (Petix et al., 2018; Service, 2017) (see Figures 16 & 17). Pertix et al. (2018)
research on the use of grazing, soil impoverishment, and applications of organic herbicide
and heat treatments to sterilize soil of non-native plant seed provides an understanding of
other restoration tools available. These treatments helped to reduce the abundance of
specific groups of non-native invasive plants, they also increased the abundance of seeded
native plant species that continue to outcompete Viola adunca for light (Dover & Settele,
2009; Petix et al., 2018). Ultimately, the study discovered that topsoil removal was most
effective in reestablishing low growing native vegetation in the foredune coastal prairies
of the Clatsop Plains. The field research conducted at the Clatsop Plains in Oregon points
to the current most effective restoration treatments in the Pacific Northwest coastal prairie
complex and suggest that a vegetative mat growing Viola adunca might aid the plant by
suppressing local non-native and even native vegetation from light and topsoil access
(Bennett et al., 2011; Buisson et al., 2006; Jones, Norman, & Rhind, 2010; Jutila & Grace,
35
2002; Petix et al., 2018; Service, 2017). This treatment had the greatest success in native
seeded forb and grass establishment as well as the lowest cover of non-native forbs and
grasses due to removal of a portion of the existing seedbank and exposure of bare soil
(Petix et al., 2018). The research corresponds to the larger study of the role of topsoil
removal in prairies (Buisson et al., 2006; Dunwiddie & Martin, 2016; Jones et al., 2010).
Petix et al. (2018) provides a very site-specific set of tools to consider for restoration
treatments associated with the coastal prairie mat research at Willapa and suggests a role
vegetative mats could play in cutting down on the cost of soil removal and maintaining
interstitial spacing for plants like Viola adunca to grow and possibly recruit.
Speyeria zerene hippolyta - Larval Survival
The paper, “Modeling caterpillar movement to guide habitat enhancement for Speyeria
zerene hippolyta, the Oregon silverspot butterfly”, examines the biology and life history of
the butterfly in relation to Viola adunca density, abundance and location. Bierzychudek &
Warner’s (2015) work highlights the last 20 years of insect conservation research related
to the Speyeria z. hippolyta (Bierzychudek, Warner, McHugh, & Thomas, 2009; Hill et al.,
2018; James, 2008). The article explains the relationship between habitat spatial structure
and the butterfly’s occurrence and abundance as related to the presence of its host plant
Viola adunca. In addition, to identifying the other variables at play related to the butterfly’s
survival; particularly other predators, the article addresses the connection between humancaused landscape alteration and the decline of Viola adunca populations and biodiversity
within regions such as Cascade Head. Using a combination of in the field observations and
computer modeling, the authors explore the role of the butterfly’s larval caterpillar stage
36
in relation to foraging behavior and movement, finding that during the caterpillar’s first
instar phase of development, the insect’s movement is limited to an area of about one meter
square. As the caterpillar develops into its second, third and fourth instar phases, the
insect’s movement based on foraging increased outside of the meter square area in a
random pattern till a Viola adunca plant was reached. Moreover, the research proposes that
with a Viola adunca density of at least four plants per square meter, the butterfly will see
a 10% increase in survivorship from caterpillar to adulthood (Bierzychudek & Warner,
2015). Insect conservation behavior research in this study, provides a clear research
connection for my hypothesis; the role of the vegetative mats in hosting both biological
and structural elements that may help foster the larval development for Speyeria z.
hippolyta, and new Viola mat metrics for measuring survival.
METHODS
Overview
The methods used in this research look to measure the effectiveness of out growing and
out planting experimental coastal prairie vegetative mats as compared to the traditional
restoration planting methods of dibble stick or shovel and plug planting (see Figures 31 &
32). The base substrate of the vegetative mat was made from coconut coir and a coating of
latex for increased durability. The coconut coir was sourced from Sri Lanka and the
company Rolankatm, a layer of Red Alder (Alnus rubra) chips was also placed on top of
the seed sown and coconut coir to retain soil moisture in the mat for sown germinated seed.
Additionally, during the summer, shade cloth was used to cover the hoop house growing
37
the mats and plugs. In contrast to the mats, an equal seed quantity was sown into the
traditional planting plugs filled with soil and topped with a thin layer of granulated granite.
All plant material associated with the mats was sourced from areas in Oregon and
Washington that represent the same genetic region and integrity as the restoration sites.
The project utilized three plant species associated with the life history of Speyeria z.
hippolyta based on ecological field data associated with Speyeria z. hippolyta’s habitat
usage at Mt. Hebo and Rock Creek Siuslaw National Forest, Oregon (Unpublished
research, Glavich 2019). The plant species tested are Viola adunca, Festuca romeri, and
Fragaria chiloensis. Restoration areas prioritized for the planting of plant material include
three sites that spanned the historic geographic distribution of the insect and are currently
undergoing various conservation actions as part the federal recovery plan for the specie
(see Figures 16-21): Willapa Bay and Nestucca Bay National Wildlife Refuges and Rock
Creek Siuslaw National Forest (Speyeria zerene hippolyta W.H. Edwards; USFWS 2001).
Figure 15: Mating pair of Speyeria zerene hippolyta.
38
Research Study Sites
Figure 16: Soil removal and Viola adunca planting for Speyeria z. hippolyta release at Willapa Refuge.
Chinook Territory - Tarlatt Slough on the US Fish & Wildlife Service’s Willapa National
Wildlife Refuge host habitat restoration opportunities for Washington’s coastal prairie. The
site is currently undergoing restoration conversion for Speyeria z. hippolyta from cow
pasture dominated by the grass (Agrostis gigantea), False dandelion (Hypochaeris
radicata), Birdsfoot trefoil (Lotus corniculatus), Lanceolate plantain (Plantago lanceolate)
and other non-native invasive plant species. Large areas of topsoil removal have been out
planted with thousands of Viola plugs (see Figures 16 & 17). Currently, these treatments
and plantings are expanding within the site and are in combination with other treatments
including herbicide, mowing and scraping to deplete the non-native seed bank. The only
natives recorded at the site where those planted. The site is proximal to Sandbar Road.
Based on research at Rock Creek, a hedge row of native shrubs has been planted to buffer
butterflies from the road (Littlejohn, 2012; Zielin et al., 2016).
39
Figure 17: Map of Tarlatt at Willapa Restoration Area.
40
Figure 18: Restored and enhanced extant habitat for Speyeria z. hippolyta at Nestucca Bay Refuge.
Tillamook Territory - Cannery Hill (Area 3 South) on the US Fish & Wildlife Service’s
Nestucca Bay National Wildlife Refuge represents a unique habitat restoration opportunity
for Oregon's coastal prairie (see Figures 18 & 19). In 2013, Cannery Hill was added to the
Refuge. Shortly thereafter habitat restoration work began converting the 1,202 acres from
Reed canary grass (Phalaris arundinacea) and other non-native pasture grasses to native
coastal prairie grasses and forb species. A combination of techniques is still being
experimented with on the grassland, including herbicide application, scraping, mowing and
others (Service, 2013). The work at Nestucca provides case studies and insights into coastal
prairie restoration best practices and techniques including invasive species control, native
seeding proportions, seeding techniques and prescribed burning (Service, 2013). In the
winter of 2014 Viola adunca seed was drilled into the site and in the summer of 2017
captively raised Speyeria z. hippolyta from Mt. Hebo were released to the site to establish
a new population in the renewed habitat.
41
Figure 19: Map of Area 3 South and Nestucca Restoration Area.
42
Figure 20: Enhanced extant habitat for Speyeria z. hippolyta. Rock Creek Siuslaw National Forest.
Alsea Territory - Area 8 of the Rock Creek Site in the Siuslaw National Forest represents
an intact habitat currently and historically harboring Speyeria z. hippolyta. Rock Creek is
located on the east and westsides of highway 101 and is a long-term study site that has
generated a wealth of data on the site-specific behavior of Speyeria z. hippolyta and the
environmental history and impacts to the area, principally the impacts to the Rock Creek
population from highway 101 (Kiser, 1993; Littlejohn, 2012; Zielin et al., 2016) Within
the sandstone cliff road cuts of the highway one can observe an absence of roots and other
historic vegetation preserve that would imply the area has long been prairie (McCorkle et
al., 1980). In other areas of the site, restoration treatments such as soil removal, scraping,
mowing and plug planting is occurring with species including Viola adunca, Yellow-eyed
grass (Sisyrinchium californicum), Beach strawberry (Fragaria chiloensis) and other
coastal prairie pollinator resources.
43
Figure 21: Map of Area 8 and Rock Creek Restoration Area.
44
Figure 22: First mat Viola adunca to flower in trial media test at SCCC.
Cold Stratified Germination and Sowing
At Stafford Creek Corrections Center (SCCC) the initial mat germination test of Viola
adunca seed sown directly into the coconut coir mats and control plug trays with a soil and
granulated gravel cover took place on December 13th, 2018, with ~108 seeds per meter
square meter of mat and two seeds per individual c7 plug with 98 plugs per tray. This
resulted in a (64 %) germination rate in the mat and (82 %) within the plugs respectively
(see Appendix). In March 2019, to ensure consistent plant establishment in the mats and
c7 control plugs, additional Viola adunca, seed was cold stratified in a refrigerator for 100
days at 40o F; this resulted in a (90%) germination rate. The cold-stratified seed was then
directly sown at a rate of ~108 seeds per m2 and two seeds per plug on August 22, 2019.
Festuca romeri was directly sown on August 22, 2019 at nine seeds per m2 and two seeds
per plug (both with an 80% germination rate), and Fragaria chiloensis was planted from
cuttings at two per m2 and one cutting per plug on September 24, 2019 (with a 95% survival
rate). Below (see Figures 23-25) show the process.
45
Table 1
Seed weight and sowing rates for target plant species
Plant Species
Seed grams/
cuttings per
plug tray
Total plug
seed/ cuttings
Seed grams/
cuttings per
1m2 mat
Total mat seed/
cuttings
Sow dates
Viola
adunca,
0.21 grams
98 plugs
5 grams
588 plugs
0.36 grams
13 grams
36 mats
12/13/18
8/22/2019*
Festuca
romeri
Fragaria
chiloensis
0.4 grams
104 plugs
1 cutting
98 plugs
1.6 grams
416 plugs
1 cutting
72 plugs
0.2 grams
7.2 grams
36 mats
72 cuttings
36 mats
8/22/2019*
2 cuttings
9/24/2019
Notes: Sowing details and dates for mat and plug sowing with * = 100-day wet cold stratification.
Figure 23: Viola adunca seed direct sow. December 13, 2018 at SCCC.
Figure 24: Viola adunca survival counts in trial media. March 23, 2019 at SCCC.
Figure 25: Cold stratified Viola seed, Festuca seed and Fragaria cuttings at SCCC.
46
Experiment Design and Site Treatments
Each of the study sites received two types of restoration treatments two weeks prior to the
planting of the plant plugs and vegetative mats—scraping and mowing. The plugs grown
were then out planted randomly within the 12 1 x 1-meter control plots. The vegetative
mats spacing, and planting mirrored the control plots within both treatment types. These
treatments and planting types are summarized in Tables 2 and 3 below (see Appendix for
vegetative mat plant survival table).
Table 2
Explanatory and response variables of research
Explanatory Variables
Response Variables
Scraping
Percent aerial cover and height of Viola adunca,
Festuca romeri, Fragaria chiloensis
Mowing
Percent aerial cover, height and phenology of native
plant species
Control Plugs
Percent aerial cover, height and phenology of nonnative plant species
Vegetative Mats
Percent aerial cover of bare ground and vegetative
mat
Table 3
Treatment and planting area size and dates
Site
Total Treatment Areas
Willapa Bay
Scraped Area 88 m2
National Wildlife Mowed Area 88 m2
Refuge
Total Planted Area
Mats 36 m2
Plugs 36 m2
Date Planted
12/16/19
12/16/19
Nestucca
Scraped area 88 m2
National Wildlife Mowed Area 88 m2
Refuge
Mats 36 m2
Plugs 36 m2
1/24/20
1/20/20
Rock Creek
Scraped area 88 m2
Siuslaw National Mowed Area 88 m2
Forest
Mats 36 m2
Plugs 36 m2
1/24/20
1/19/20
47
Experiment Design of Study Plots
Illustration of study area for each vegetative mat and control plug planting area.
Vegetative Mats
Scraped treatment
Mowed treatment
3m
3m
1m
Coconut Coir/
Viola adunca,
Festuca roemeri
Fragaria
chiloensis
1m
Coconut Coir/
Viola adunca,
Festuca roemeri,
Fragaria
chiloensis
Control Plugs
Scraped treatment
for the Oregon
1m
Mowed treatment
3m
3m
Control Plugs of
22 Viola adunca,
9 Festuca
roemeri
2 Fragaria
chiloensis
1m
Control Plugs of
22 Viola adunca,
9 Festuca
roemeri
2 Fragaria
chiloensis
Figure 26: Study area for vegetative coastal prairie mats plant plug control installation design within
treatment types.
48
Experimental Design of Treatment / Planting Areas
Scraping with Plugs and Mat
11 m
2m
\
8m
11 m
2m
8m
Figure 27: Scraped treatment out planting designs for control plugs and vegetative mats.
(A = Viola adunca, B = Festuca roemeri, C = Fragaria chiloensis).
49
Mowing with Plugs and Mat
11 m
2m
8m
11 m
2m
8m
Figure 28: Mowed treatment out planting designs for control plugs and vegetative mats.
(A = Viola adunca B = Festuca roemeri, C = Fragaria chiloensis).
50
Over the course of the research project’s installation, the time to complete each treatment
method was measured for a cost benefit analysis (see Appendix). For both Willapa and
Nestucca Bay National Wildlife Refuge site treatment prep began on 12/7/19 and 12/17/19
respectively. Here, refuge staff employed the use of a tractor pulled flail mower to scrape,
and mow both sites’ 8 x 22-meter study areas on 1/18/20. At the Rock Creek Siuslaw
National Forest, the site topography made it inaccessible to the tractor. In place of the flail
mower Forest Service staff employed the use of a weed whacker, rakes and a Rototiller to
implement scraping and mowing treatments. While the use of these tools took considerably
longer, treatment results yielded similar outcomes.
Figure 29: Scraped and mowed treatment areas at Nestucca National Wildlife Refuge using a tractor flail
mower 12/17/19.
Figure 30: Scraped and mowed treatment areas at Rock Creek Siuslaw National Forest using a weed
whacker and Rototiller 1/18/20.
51
Control Plug and Mat Planting
The Sustainability in Prisons Project (SPP), a partnership between the Evergreen State
College and the Washington Department of Corrections, grew a total of 36 1 x 1 meter
mats and the study’s control plant plugs within a hoop house at the controlled nursery
environment of Stafford Creek Correctional Facility’s Conservation Nursery in Aberdeen,
Washington. Each vegetative mat was secured using 9 stainless-steel yard staples. All
control plugs were installed using a dibble stick and bisecting hole edge scores by hand
trowel to ensure soil and plug contact (see Figures 31 & 32).
Figure 31: Scraped treatment area at Nestucca National Wildlife Refuge with plug planting process with
control Viola adunca plugs.
Figure 32: Vegetative mat and control plant plugs for Rock Creek Siuslaw National Forest.
52
Monitoring Data and Analysis
ArcGIS Survey123 Connect was used for XLS form design for electronic field data
collection (see Figure 33). The project’s data collection, storage and upload were all done
through the Coastal Prairie Monitoring Application I developed. Data was collected into
three groups (Native, Non-native and Site Details) with local and global IDs. A drop-down
plant list with detailed photo identification for each plant was also created within the
application to assure all plants were correctly identified. The plant list was composed of
native and non-native plants provided by Willapa, Nestucca Bay National Wildlife Refuge,
and the Siuslaw National Forest (see Appendix). Site data collected within the planting
plots included site planting and restoration treatment type, the percent aerial cover and
height of native plant species, the percent aerial cover and height of non-native plant
species and the plants’ phenology. For the purpose of the study, the percent aerial cover of
bare ground and vegetative mats (coconut coir) were included in the native grouping, as
both substrates constitute as interstitial space. Photos of each plot and subplot were also
collected. To account for edge effect on the mats and plug plots, a buffer of two meters
surrounding the plots was established; mats and plugs were planted two meters apart from
each other. Subplot data was gathered from the northwestern 1 x 1-meter areas surrounding
the plots, to understand the response of the existing plant community to the treatment types
and its potential effect on the experiment’s variables without planting. In June plant height
was measured using a bisecting ruler that fit over the quadrate (see Figure 34). Microsoft
Excel was used for data management, QA/QC and ArcGIS Insights was used for initial
analysis, data visualization and mapping. October data was analyzed in R Studio with a
Two-way MANOVA.
53
Figure 33: Coastal Prairie Monitoring application at Nestucca National Wildlife Refuge.
Figure 34: Monitoring quadrate and height ruler.
54
RESULTS
Following the treatments and plantings in December and January the first monitoring event
began on June 6, 2020 and ended on October 25, 2020. Sites were monitored from south
to north beginning at Rock Creek traveling northward to Nestucca and Willapa. Monitoring
occurred over the course of two weeks. During this initial visit all the project variables
were collected. Other research evidence collected included site, plot and subplot photo
points (see Figures 40 - 43). Additional photo point data can be accessed upon request.
Based on vegetative mat planting and scraping treatments, from June to October native
plant aerial cover was dominated by 15 species observed across all, some or one of the sites
(see Figure 36 and Appendix for plant codes). Based on vegetative mat planting and
mowing treatments, June native plant aerial cover was dominated by 13 species observed
across all, some or one of the sites (see Figure 37 and Appendix for plant codes).
Comparison bar charts display June’s percent native and non-native aerial cover based on
treatment and planting types (see Figures 44 & 45). These comparison bar charts are
repeated to show how June and Octobers’ percent native and non-native aerial cover based
on treatment and planting types changed throughout the projects’ growing season (see
Figures 52 & 53). Data collected during June provided the first opportunity to collect
information on the interaction effect of these variables which can be seen in the link charts
that reflect the top down hierarchical relationships of the treatments to the planting types
on total native aerial plant cover for June and October (see Figures 47 & 48). The link chart
can be understood as a data visualization tool to display data associations, with thicker
chart lines displaying more association between variable points. With points such as
planting or treatments type at the top of the chart having a greater association with a
55
variable, in this case native aerial cover. Across all planting and treatment types Willapa
saw the greatest increase in the sum of Viola adunca aerial cover over the project’s duration
from (68) in June to (128) in October. Between June and October, Nestucca saw a decrease
in the sum of Viola adunca aerial cover from (57) to (51) while Rock Creek’s cover
changed from (38) to (31) by October (see Figures 49 & 50). Scraped areas at Nestucca
with in subplots yielded Viola adunca germination and establishment (see Figure 35). As
built photo points were established during the initial planting and visited in June and
October to document the plant growth, phenology, and succession over the entire growing
season (see Figures 56 - 73). These photo points included captures of the mowed treatment,
scraped treatment and the full project area. Plot and subplot photos collected help to
visually illustrate the plant communities’ competition with the planting and treatment types
(see Figures 40 - 43).
Figure 35: Field monitoring observations. Viola adunca in scraped area. Solidago canadensis and Achillea
millefolium growing up through a mat at Nestucca. Mat Violas at Willapa.
56
Nestucca
Rock Creek
Willapa
Figure 36: June 2020 native plant aerial cover by site and specie. Filtered by scraping and vegmat.
Nestucca
Rock Creek
Willapa
Figure 37: June 2020 native plant aerial cover by site and specie. Filtered by mowing and vegmat.
57
Nestucca
Rock Creek
Willapa
Figure 38: October 2020 native plant aerial cover by site and specie. Filtered by scraping and vegmat.
Nestucca
Rock Creek
Willapa
Figure 39: October 2020 native plant aerial cover by site and specie. Filtered by mowing and vegmat.
58
Figure 40: January 2020 plot 5 planted in scraped area with vegetative mat.
Figure 41: June 2020 plot 5 planted in scraped area with vegetative mat.
59
Figure 42: January 2020 plot 5 scraped area planted with plant plugs.
Figure 43: June 2020 plot 5 scraped area planted with plant plugs.
60
Figure 44: June 2020 percent native aerial cover based on treatment and planting type.
Figure 45: June 2020 percent non-native aerial cover based on treatment and planting type.
61
Figure 46: June 2020 non-native height based on treatment and planting type.
Figure 47: June 2020 relation of treatment and planting type on native plant aerial cover.
62
Figure 48: October 2020 relation of treatment and planting type on native plant aerial cover.
Willapa
Rock
Creek
Nestucca
Figure 49: June 2020 total Viola adunca aerial cover by site.
63
Willapa
Nestucca
Rock
Creek
Figure 50: October 2020 total Viola adunca aerial cover by site.
Figure 51: October Viola adunca with vegetative mat on scraped areas. Willapa, Nestucca and Rock Creek.
64
Figure 52: October 2020 percent native aerial cover based on treatment and planting type.
Figure 53: October 2020 percent non-native aerial cover based on treatment and planting type.
65
Figure 54: June 2020 relation of treatment and planting type on target native plant aerial cover.
Figure 55: October 2020 relation of treatment and planting type on target native plant aerial cover.
66
Willapa National Wildlife Refuge
Figure 56: December 2019 vegetative mats with scraping at Willapa.
Figure 57: June 2020 vegetative mats with scraping at Willapa.
67
Figure 58: October 2020 vegetative mats with scraping at Willapa.
Figure 59: December 2019 vegetative mats with mowing at Willapa.
68
Figure 60: June 2020 vegetative mats with mowing at Willapa.
Figure 61: October 2020 vegetative mats with mowing at Willapa.
69
Nestucca Bay National Wildlife Refuge
Figure 62: January 2020 vegetative mats with scraping at Nestucca.
Figure 63: June 2020 vegetative mats with scraping at Nestucca.
70
Figure 64: October 2020 vegetative mats with scraping at Nestucca.
Figure 65: January 2020 vegetative mats with mowing at Nestucca.
71
Figure 66: June 2020 vegetative mats with mowing at Nestucca.
Figure 67: October 2020 vegetative mats with mowing at Nestucca.
72
Rock Creek Siuslaw National Forest
Figure 68: January 2020 vegetative mats with scraping at Rock Creek.
Figure 69: June 2020 vegetative mats with scraping at Rock Creek.
73
Figure 70: October 2020 vegetative mats with scraping at Rock Creek.
Figure 71: January 2020 vegetative mats with mowing at Rock Creek.
74
Figure 72: June 2020 vegetative mats with mowing at Rock Creek.
Figure 73: June 2020 vegetative mats with mowing at Rock Creek.
75
DISCUSSION & CONCLUSION
The future potential for this research to explore and grow is immense. There are many other
plants that host endangered insects in peril, this restoration technology holds opportunities
to explore and improve the functions of these relationships (Bierzychudek & Warner,
2015; Dunn, 2005; Hill et al., 2018; Shuey et al., 2016; Stiling & Moon, 2005). Following
a Two-way MANOVA test of the results for October’s monitoring with planting type as
the factor, indicated a statistically significant difference in native to non-native aerial plant
cover between planting types (plugs and the vegetative mats) for October F(3, 416) =
203.39, p ≤ (0.001). This statistically significant difference can be seen in the stack bar
charts of the two planting types for October (see Figure 75). Additionally, the study was
able to analyze the interaction effect between treatment types (scraping and mowing) on
native and non-native aerial plant cover for October plots. An additional Two-way
MANOVA test with treatment as the factor indicated a statistically significant difference
in native aerial plant cover between the restoration treatment types (mowing and scraping)
for October F(3, 416) = 15.58, p ≤ (0.001). The vegetative mats grew Viola adunca, other
native plant species and maintained interstitial space more effectively than the plugs
planted in scraped or mowed areas (see Figures 44, 52, 74 & 75). Moreover, scraped area
treatments planted with plugs and mats grew Viola aduna and the other native target
species more effectively than the mowed area treatments (see Figures 54 & 55). While
plugs in the scraped areas grew native species like Viola adunca and other target species
in amounts comparable to the mats, the amount of open and available bare ground (baregr)
for future plant recruitment in the scraped plug planted areas was less in comparison to the
space still available for plant recruitment in the vegetative mats (vegmat) (see Figure 75).
76
Figure 74: June target native aerial cover by planting type.
Figure 75: October target native aerial cover by planting type.
77
The vegetative mats were also associated with less non-native vegetative height across the
sites (see Figure 46). This is another key finding as plant height influences Viola adunca's
ability to grow as well as Speyeria z. hippolyta’s ability to seek out the plant to lay its eggs
(Schaeffer, 1992). Prior research points to habitats that used to support the butterfly as
having violet densities ranging from (20 to 100 per m2) (Kiser, 1993; Schaeffer, 1992).
While both the plug and mat plantings in scraped areas were able to yield Viola cover, site
variability, soil moisture, avian disturbances to the mats, and other variables did not allow
for Viola adunca coverage to be greater than (16%) at sites like Willapa at any one plot
(see Figures 76-78). With next year marking the first year for the release of captively reared
Speyeria z. hippolyta at Willapa this is encouraging, as the site is one of the more degraded
and undergoing active restoration. However, it should also be noted that Willapa was the
first site planted on 12/16 and 12/17/2019, which could have played a factor in Viola
growth and recorded cover as Nestucca and Rock Creek were planted a month later. Future
research into the vegetative mats ability over time to achieve these Viola densities as well
as further research to investigate if adult butterflies are attracted to the mat structure and
plant community for ovipositioning and other behavior, should be conducted (Kiser, 1993).
Research into Speyeria z. hippolyta’s larval survival within the mats themselves would
prove to be invaluable information in the quest to save a highly localized insect, while
moving towards a global consciousness of insect conservation (Bierzychudek & Warner,
2015; Bierzychudek et al., 2009; Crone et al., 2007; Samways, 2020; Thorpe & Stanley,
2011). Other interesting field observations of the vegetative mats included qualitative data
recording on the phenological suppression of the vegetative mats to the surrounding plant
78
community, which showed some association between the plant phenology to the planting
and treatment types. In addition to suppressing non-native aerial plant cover and height,
the mats also appeared to slow the flowering and fruiting of both native and non-native
species. Plugs of Viola adunca planted appeared to flower in June whereas Viola adunca
growing within the mats were flowering and fruiting during monitoring events in October
(see Figure 35). Plant associations to each other in the mat and scraped treatment areas
warrant additional research and a management experimentation at sites such as Rock Creek
and Nestucca, where scraped areas with a seed bank of Viola adunca yielded germination
and establishment of the plant (see Figure 35). The role of soil disturbance in Viola
adunca’s germination along with other species could provide an alternative to fire or
herbicide (Almasi & Kollmann, 2007; Dunwiddie & Martin, 2016; Jutila & Grace, 2002).
Mats could be planet on hillsides or hilltops to provide a Viola colony in which seed banks
could be built, promoted and stimulated around the mat by scraping the surrounding soil
biannually. Scraping also played an instrumental role in the mats ability to establish soil
contact and the planted plugs ability to yield greater Viola growth by reducing the overall
existing plant biomass (see Figures 76 – 78). Other link charts created for June and October
datasets show the top down hierarchical relationships of the planting types on native aerial
plant cover by species (see Figures 79 & 80). Here the association of the planting type to
the sum of native plant species aerial cover, is reflected in the thickness of the lines that
point to the plant code, with thicker lines reflecting greater associations with the planting
type (see Appendix for native plant codes). Data exploration of these link charts and others
along with an ArcGIS Story Map of the MES Research project can be found online at
(https://arcg.is/998KK2).
79
Scraped
Mowed
Figure 76: Willapa October 2020 planted Viola adunca aerial cover.
Scraped
Mowed
Figure 77: Nestucca October 2020 planted Viola adunca aerial cover.
80
Scraped
Mowed
Figure 78: Rock Creek October 2020 planted Viola adunca aerial cover.
Figure 79: Link chart of June native plant species aerial cover and association with planting type.
81
Figure 80: Link chart of October native plant species aerial cover and association with planting type.
The association of the native plant community to planting and treatment types could help
to reveal a greater understanding of the correct combinations of prairie restoration
prescriptions. Developing a clearer understanding of these best management practices over
time will inform a place based approach towards future restoration and habitat
enhancement in the coastal prairie ecosystem (Petix et al., 2018). These associations can
help to create a high resolution understanding of how localized adaptive management
practices effect plant structure, and how the structure of the plant community itself might
influence the survival and quality of host plants and other restoration target species
(Awmack & Leather, 2002; Bauerfeind & Fischer, 2013; Menéndez et al., 2007; Sivakoff
et al., 2016). An ordination test on June and October datasets would be of value to
developing a sense of the role ocean proximity and salt spray might play in maintaining
coastal prairie plant community height and structure across all sites. Soil and surface level
salinity experimentation should be conducted on the vegetative mats to observe the
response of non-native and native plants such as Viola adunca to examine the herbicidal
effects of salt water (Kiser, 1993; Schaeffer, 1992).
82
Figure 81: Viola adunca ecosystem. The role of structure and plant associations for insects.
Based on visual observations of some of the most successful mats harboring the project’s
three target species, it could be hypothesized that the three may help to co-facilitate Viola
adunca’s biology and protect the life cycle of the caterpillar of Speyeria z. hippolyta as it
moves through its six instar phases and the violet propagates (see Figure 81 - 83). Eggs
laid on Viola adunca hatch, the larva consumes its egg casing and begins its new diet of
Viola adunca leaves. As the caterpillar moves closer to diapause it moves into the Festuca
romeri for a protected place to go through diapause. Festuca romeri may help provide
Viola adunca with the soil structure and moisture it needs for germination and
establishment of new seed with micro shade from the bunch grass. Following diapause, the
83
larva wakes up and walks out to eat on Viola adunca leaves and is protected visually from
predators by Fragaria chiloensis as it develops and seeks new violets. The evergreen leaves
of Fragaria chiloensis could also help to shade and shed water towards Viola aduca,
promoting further seed and leaf production by the plant. These leaves may also help to
preclude light from non-natives while also keeping vegetative growth lower and lateral
benefiting Viola adunca. Moreover, Fragaria chiloensis could attract pollinators of the
violet and another insect in need of conservation action––solitary bees (Freitas & Sazima,
2003; Tonietto & Larkin, 2018). Finally, the caterpillar climbs up Festuca romeri to pupate
and metamorphizes into the form of the Oregon silverspot butterfly (Speyeria zerene
hippolyta). Insects reveal to us the value of life’s relationship to itself. A way to see one’s
self as part of a whole ecological value; protecting, restoring and inspiring our ecosystem.
Figure 82: Viola adunca mat ecosystem. The role of structure and plant associations for insects.
84
Figure 83: Speyeria zerene hippolyta ecosystem. Hard and soft ground etching, 2014.
85
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Appendices
Composite Native Plant List of All Sites
Native Plants
Plant Code
Plant Specie
native
achmil
Achillea millefolium
native
acmame
Acmispon americanus
native
acmpar
Acmispon parviflorus
native
agraeq
Agrostis aequivalvis
native
agrexa
Agrostis exarata
native
agrhal
Agrostis hallii
native
agrpal
Agrostis pallens
native
allcer
Allium cernuum
native
alnrub
Alnus rubra
native
anamar
Anaphalis margaritacea
native
anghen
Angelica hendersonii
native
angluc
Angelica lucida
native
aphocc
Aphanes occidentalis
native
aqufor
Aquilegia formosa
native
artsuk
Artemisia suksdorfii
native
astsub
Aster subspicatus
native
blespi
Blechnum spicant
native
botmul
Botrychium multifidum
native
brocar
Bromus carinatus
native
calcan
Calamagrostis canadensis
96
native
calnut
Calamagrostis nutkaensis
native
calcil
Calandrinia ciliata
native
camqua
Camassia quamash var. maxima
native
caroli
Cardamine oligosperma
native
carobn
Carex obnupta
native
carpan
Carex pansa
native
carros
Carex rossii
native
carsco
Carex scoparia
native
cartum
Carex tumulicola
native
caslit
Castilleja litoralis
native
chaang
Chamerion angustifolium
native
ciredu
Cirsium edule
native
claamo
Clarkia amoena var. caurina
native
claper
Claytonia perfoliata
native
clasib
Claytonia sibirica
native
dancal
Danthonia californica
native
daupus
Daucus pusillus
native
desces
Deschampsia cespitosa
native
elygla
Elymus glaucus ssp. glaucus
native
epicil
Epilobium ciliatum
native
equar
Equisetum arvense
native
erigla
Erigeron glaucus
native
erilan
Eriophyllum lanatum
native
erygut
Erythranthe guttata
97
native
fesam
Festuca ammobia
native
fesrom
Festuca roemeri
native
fesrub
Festuca rubra ssp. juncea
native
frachi
Fragaria chiloensis
native
galap
Galium aparine
native
gampur
Gamochaeta purpurea
native
glyspp
Glyceria spp.
native
habgre
Habenaria greenei
native
herlan
Heracleum lanatum
native
holdis
Holodiscus discolor
native
hosgra
Hosackia gracilis
native
iriten
Iris tenax
native
junbuf
Juncus bufonius var. bufonius
native
juneff
Juncus effusus
native
junens
Juncus ensifolius
native
junpat
Juncus patens
native
luplit
Lupinus littoralis
native
lupriv
Lupinus rivularis
native
luzcom
Luzula comosa
native
maidil
Maianthemum dilatatum
native
marore
Marah oregana
native
monfon
Montia fontana
native
navsqu
Navarretia squarrosa
native
philew
Philadelphus lewisii
98
native
phycap
Physocarpus capitatus
native
picsit
Picea sitchensis
native
plamar
Plantago maritima
native
plebra
Plectritis brachystemon
native
poamac
Poa macrantha
native
poapal
Poa palustris
native
polmun
Polystichum munitum
native
potpac
Potentilla pacifica
native
pruvul
Prunella vulgaris var. vulgaris
native
pteaqu
Pteridium aquilinum
native
pucnut
Puccinellia nutkaensis
native
ranocc
Ranunculus occidentalis
native
rosgym
Rosa gymnocarpa
native
rubpar
Rubus parviflorus
native
rubspe
Rubus spectabilis
native
ruburs
Rubus ursinus
native
rumocc
Rumex occidentalis
native
salsco
Salix scouleriana
native
salsit
Salix sitchensis
native
samarb
Sambucus racemosa var. arborescens
native
sancra
Sanicula crassicaulis
native
scigla
Scirpus glaucus
native
scrcal
Scrophularia californica ssp. californica
native
sidhir
Sidalcea hirtipes
99
native
sisbel
Sisyrinchium bellum
native
siscal
Sisyrinchium californicum
native
sisida
Sisyrinchium idahoense var. idahoense
native
solelo
Solidago elongata
native
solgil
Solidago simplex v. gillmanii
native
solspa
Solidago simplex var. spathulata
native
spirom
Spiranthes romanzoffiana
native
stamex
Stachys mexicana
native
symchi
Symphyotrichum chilense
native
triwor
Trifolium wormskioldii
native
vicgig
Vicia gigantea
native
vioadu
Viola adunca
Composite Non-native Plant List of All Sites
Non - Native
Plants
Plant Code
Plant Specie
non_native
agrrep
Agropyron repens
non_native
agrcap
Agrostis capillaris
non_native
agrgig
Agrostis gigantea
non_native
agrsto
Agrostis stolonifera
non_native
airpra
Aira praecox
non_native
alopra
Alopecurus pratensis
non_native
ammare
Ammophila arenaria
100
non_native
antodo
Anthoxanthum odoratum
non_native
arrela
Arrhenatherum elatius ssp. elatius
non_native
belper
Bellis perennis
non_native
cerfon
Cerastium fontanum ssp. vulgare
non_native
cerglo
Cerastium glomeratum
non_native
cirvul
Cirsium vulgare
non_native
crecap
Crepis capillaris
non_native
cynspp
Cynosurus spp.
non_native
dacglo
Dactylis glomerata
non_native
dandec
Danthonia decumbens
non_native
daucar
Daucus carrota
non_native
digpur
Digitalis purpurea
non_native
erocic
Erodium cicutarium
non_native
fesaru
Festuca arundinacea
non_native
fesrub
Festuca rubra
non_native
gerdis
Geranium dissectum
non_native
germol
Geranium molle
non_native
gnauli
Gnaphalium uliginosum
non_native
hollan
Holcus lanatus
non_native
hypper
Hypericum perforatum
non_native
hyprad
Hypochaeris radicata
non_native
ilaaqu
Ilex aquifolium
non_native
lampur
Lamium purpureum
non_native
lapcom
Lapsana communis
101
non_native
leuvul
Leucanthemum vulgare
non_native
lolper
Lolium perenne
non_native
lolspp
Lolium spp.
non_native
lotcor
Lotus corniculatus
non_native
lotuli
Lotus uliginosus
non_native
malneg
Malva neglecta
non_native
matdis
Matricaria discoidea
non_native
medlup
Medicago lupulina
non_native
myodis
Myosotis discolor
non_native
parvis
Parentucellia viscosa
non_native
permac
Persicaria maculosa
non_native
phaaru
Phalaris arundinacea
non_native
plalan
Plantago lanceolata
non_native
plamaj
Plantago major
non_native
poaann
Poa annua
non_native
poapra
Poa pratensis
non_native
poatri
Poa trivialis
non_native
pruvul
Prunella vulgaris var. vulgaris
non_native
ranpar
Ranunculus parviflorus
non_native
ranrep
Ranunculus repens
non_native
rubarm
Rubus armeniacus
non_native
rublac
Rubus laciniatus
non_native
rumace
Rumex acetosella
non_native
rumcri
Rumex crispus
102
non_native
rumspp
Rumex spp.
non_native
sagpro
Sagina procumbens
non_native
scharu
Schedonorus arundinaceus
non_native
senjac
Senecio jacobaea
non_native
senmin
Senecio minimus
non_native
sensyl
Senecio sylvaticus
non_native
senvul
Senecio vulgaris
non_native
sisoff
Sisymbrium officinale
non_native
solspp
Solanum spp.
non_native
sonasp
Sonchus asper
non_native
taroff
Taraxacum officinale
non_native
tridub
Trifolium dubium
non_native
tripra
Trifolium pratense
non_native
trirep
Trifolium repens
non_native
vulmyu
Vulpia myuros
103
Vegetative Mat Viola adunca Hoop House Survival Table
Vegetative
Mat ID
Total Viola
Plant count
12.10.2019
(Directly
sown
12.13.18)
Survival Total Viola
rate out Plant count
3.22.2019
of
(Directly
~ 324
sown
seeds
12.13.18)
Survival Total
Total
rate out Viola Plant Survival
count
rate out
of
6.14.2019 of
~ 324
(Directly
~ 324
seeds
sown
seeds
12.13.18)
WA SOP 1
94
29%
69
21 %
48
15 %
WA SOP 2
85
26%
14
4%
51
16 %
WA SOP 3
144
44 %
28
9%
47
15%
WA SOP 4
123
38 %
4
1%
69
21 %
OR CLO 1
201
62 %
31
10 %
68
21 %
OR CLO 2
259
80 %
32
10 %
91
28 %
OR CLO 3
270
83 %
41
13 %
85
26 %
OR CLO 4
256
79 %
13
4%
60
19 %
OR CLO 5
252
78 %
30
9%
77
24 %
OR CLO 6
267
82 %
11
3%
50
15 %
OR CLO 7
280
86 %
35
11 %
101
31 %
OR CLO 8
255
79 %
34
11 %
67
21 %
Average
207
64%
29
9%
68
21 %
Notes: Each initial vegetative mat grown in a hoop house 3x1 meters in size sown with ~ 324 Viola adunca
seeds for each sowing. SOP = Salish prairie seed source, CLO = Cascade lowlands seed source.
104
Cost Benefit Table of Treatment and Planting Types
Site
Scraping
Time
Mowing
Time
Plug
Time
Mat
Time
Tools
Willapa
10 min
5 min
5 hrs 30
min
35 min
Tractor Flail mower
Nestucca
10 min
5 min
5 hrs 30
min
35 min
Tractor Flail mower
Rock Creek
7 hrs
3 hrs
5 hrs
35 min
Weed Whacker
15 min
50 min
30 min
Rake
Rototiller
Total
7 hrs
4 hrs
35 min
Total Cost @
$35 per hour
$270
$140
16 hrs
30 min
1 hrs
$ 578
$62
Humans
45 min
Total
$1050
105
