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OVIPOSITION PREFERENCE IN TAYLOR’S CHECKERSPOT
BUTTERFLIES (EUPHYDRYAS EDITHA TAYLORI): COLLABORATIVE
RESEARCH AND CONSERVATION WITH INCARCERATED WOMEN
by
Dennis Aubrey
A Thesis
Submitted in partial fulfillment
of the requirements for the degree
Master of Environmental Studies
The Evergreen State College
June 2013
©2013 by Dennis Aubrey. All rights reserved.
This Thesis for the Master of Environmental Studies Degree
by
Dennis Aubrey
has been approved for
The Evergreen State College
By
________________________
Carri J. LeRoy
Member of the Faculty
________________________
Date
ABSTRACT
Oviposition preference in Taylor’s checkerspot butterflies (Euphydryas editha
taylori): Collaborative research and conservation with incarcerated women
Dennis Aubrey
Taylor’s checkerspot butterfly (Euphydryas editha taylori) is a federally
threatened pollinator of increasingly rare prairies in the Willamette Valley-Puget
Tough-Georgia Basin ecoregion. Since the arrival of European settlers, land use
changes, habitat fragmentation, and invasive species have contributed to a decline
in available native host plants for E. e. taylori larvae. The most commonly utilized
host is now lance-leaf plantain (Plantago lanceolata), an exotic species long
prevalent in the area. None of the known native hosts are ideal for supporting E. e.
taylori recovery efforts, so P. lanceolata is currently planted at butterfly
reintroduction sites. Golden paintbrush (Castilleja levisecta), a federally
threatened perennial, does not now co-occur with E. e. taylori but may have been
an important host historically and could be more suitable than the known native
hosts. Previous work has shown that oviposition preference is: 1) heritable and
may provide clues as to which hosts were historically important, and 2) is
correlated with larval success so might indicate which native hosts would be most
effective at restoration sites. I undertook a manipulative oviposition preference
experiment to determine which potential hosts were preferred by E. e. taylori
among P. lanceolata, C. levisecta, and harsh paintbrush (Castilleja hispida), a
known native host. The two Castilleja spp. were preferred equally, but both were
preferred over P. lanceolata. If further research confirms the suitability of C.
levisecta as a host for E. e. taylori, restoration efforts for the two species could be
united, and the effectiveness of both might be synergistically increased. This
project was undertaken collaboratively with inmates at Mission Creek Corrections
Center for Women with support from the Sustainability in Prisons Project and the
Washington Department of Fish and Wildlife, and seeks to benefit multiple
stakeholders through an interdisciplinary intersection of conservation biology and
social sustainability.
TABLE OF CONTENTS
LIST OF FIGURES................................................................................................vi
ACKNOWLEDGMENTS.....................................................................................vii
INTRODUCTION & LITERATURE REVIEW.....................................................1
Taylor’s checkerspot....................................................................................4
Species status, life history, and restoration challenges....................5
Anticipated effects of regional climate change................................8
Habitat restoration..........................................................................10
Translocation and captive breeding...............................................13
Oviposition host plants..................................................................14
Golden paintbrush......................................................................................15
ARTICLE MANUSCRIPT....................................................................................18
Introduction................................................................................................18
Methods......................................................................................................22
Site description...............................................................................22
Data collection...............................................................................23
Statistical analysis..........................................................................26
Results........................................................................................................26
Discussion..................................................................................................28
Acknowledgements....................................................................................30
References..................................................................................................30
DISCUSSION & BROADER IMPACTS.............................................................35
Ecological Implications.............................................................................35
Taylor’s checkerspot......................................................................35
iv
Golden paintbrush..........................................................................38
Collaboration..............................................................................................39
The Sustainability in Prisons Project.............................................39
Value of collaborative conservation..............................................45
Social Implications.....................................................................................46
Transforming prisons.....................................................................46
Community benefits.......................................................................52
Involving underserved audiences...................................................53
An interdisciplinary thesis.........................................................................53
Conclusions................................................................................................54
References..................................................................................................57
iv
LIST OF FIGURES
INTRODUCTION & LITERATURE REVIEW
Figure 1 – Historic prairies..........................................................................2
Figure 2 – Current prairies...........................................................................2
ARTICLE MANUSCRIPT
Figure 1 – Details of oviposition preference testing methods...................25
Figure 2 – Mean oviposition preference score..........................................27
Figure 3 – Trial results per pairing............................................................28
DISCUSSION & BROADER IMPACTS
Plate 1 – Butterfly rearing facility at MCCCW.........................................42
Plate 2 – Inmate technicians working in greenhouse interior....................42
Plate 3 – Taylor’s checkerspot ovipositing on golden paintbrush.............50
vi
ACKNOWLEDGEMENTS
The butterfly facility at Mission Creek Corrections Center for Women is managed
by the Sustainability in Prisons Project and Washington Department of
Corrections staff, overseen by the Washington Department of Fish and Wildlife,
assisted by the endangered butterfly lab at the Oregon Zoo (Portland, Oregon),
and funded by a US Fish and Wildlife Grant through the US Army Compatible
Use Buffer program. I thank all of these partners, The Evergreen State College,
and WDOC administration for their support of both this research and the
overarching vision of bringing science and nature into prisons. I also thank the
individuals that made this research a reality: biologist Mary Linders from
WDFW; Kelli Bush, Joslyn Trivett, and Carl Elliott from SPP; undergraduate
interns Caitlin Fate and Chelsea Oldenburg from TESC; Superintendent Wanda
McRae, Anne Shoemaker, Leo Gleason, Sherri Albrecht, and Dan Davis from
MCCCW; Ted Thomas of the U.S. Fish and Wildlife Service; Mary Jo Andersen
and Karen Lewis from the Oregon Zoo; and five inmates, who will remain
unnamed, that supported and participated in gathering these data. Lastly, my
reader Carri LeRoy provided tireless guidance on everything from methods design
to statistical analysis and preparation for peer-reviewed publication.
vii
INTRODUCTION & LITERATURE REVIEW
The prairies and oak savannas of the Willamette Valley-Puget Sound-Georgia
Basin (WPG) ecoregion, in the western United States, are increasingly rare and
are recognized as one of the most endangered ecosystem types in the region (Noss
et al. 1995, Floberg et al. 2004, Stanley et al. 2008, Dunwiddie and Bakker 2011).
Early descriptions of the vegetation and habitat types within the WPG come from
David Douglas who explored the area in the early 1800’s. Then Lang (1961)
briefly described the prairies of the south Puget lowlands, a subregion of the
WPG, as being a mosaic of grasslands, oak and conifer savannas, and wetlands.
The first overview of WPG prairies was provided by Franklin and Dyrness
(1973), but more detailed surveys were still needed. Giles (1970) and del Moral
and Deardoff (1976) surveyed small subsets of regional prairies but not until
Chappel and Crawford (1997) was the vegetation exhaustively catalogued. These
1997 surveys provide a valuable baseline for current and future analyses of
changing plant ranges and assemblages.
Today, one of the defining characteristics of WPG prairies is their
scarcity. In the south Puget lowlands, high quality prairie cover has declined to
3% of its historic extent based on soil surveys (Crawford and Hall 1997);
however, if semi-native and non-native grasslands are included, 24.4% remain
(Figures 1 & 2). The first to note the spatial decline of the grassland/woodlands in
the region was Giles (1970), who examined Douglas-fir (Pseudotsuga menzesii
(Mirb.) Franco) encroachment on south Puget lowland prairies. Further studies
1
Figure 1. Historic prairies (based on soil surveys), in the south Puget
lowland ecoregion: 173,261 acres; largest patch: 63,641 acres; mean
patch size: 262 acres; adapted from Crawford et al. 1994
Figure 2. Remaining grasslands and prairies (2005): 42,353 acres
(24.4%); largest patch: 3,778 acres (5.9%); mean patch size: 18 acres
(6.9%); adapted from Crawford and Hall 1997
2
surveyed the flora of these ecosystems in natural (del Moral and Deardorff 1976)
and disturbed (Jackson 1982) settings, but it was not until Clampitt (1993) that the
differences between natural and disturbed Puget lowland prairies were quantified.
Clampitt (1993) concluded that no native prairies remain in western Washington,
and that at least one native species (Aster curtis) was unable to persist in disturbed
habitats. More projects followed these, and it is now clear that land use changes,
habitat fragmentation and invasion by exotics have all contributed to the
continuing decline in both the extent and functionality of these prairie ecosystems
(Fimbel 2004, Grosboll 2004, Stanley et al. 2008), and many obligate species
have become imperiled as a result (Dennehy et al. 2011, Hamman et al. 2011,
Schultz et al. 2011, Wold et al. 2011).
One important concern for situations involving multiple declining species
involves the potential of lost interactions. For example, Kearns et al. (1998)
explored “endangered mutualisms” with respect to loss of pollination services.
Particularly, they discuss the effects of population decline and habitat
fragmentation in decreasing the pollinator/host interactions among declining
species, and argue that such impacts could range in severity according to the
dependence of the interaction. This was one of the earliest times this concept had
been discussed in a community ecology context, and it opened the door to a
growing body of work on the topic of co-extinction (Koh et al. 2004, 2004,
Rezende et al. 2007, Dunn et al. 2009). Further work modeling the effects of lost
mutualisms across phylogenetic trees was done by Rezende (2007), showing that
co-extinction leads to “non-random pruning” of phylogenetic tree branches. Pin
3
Koh et al. (2004) modeled other species relationships at risk of facilitating coextinction, including parasites and their hosts such as butterflies and their larval
host plants, and concluded that 6300 known species were “co-endangered”
because of an interaction with another listed species. Dunn et al. (2009)
summarized that the two broad interactions most likely to facilitate co-extinction
are mutualism and parasitism, due to the highly specific dependence often
associated with these relationships.
Two threatened species found on WPG prairies, the ranges of which were
broadly overlapping historically but do not now co-occur, are Taylor’s
checkerspot butterfly (Euphydryas editha taylori), and golden paintbrush
(Castilleja levisecta). Possible historic interactions between these two species
would certainly have included mutualistic pollination, but if it could be shown
that E. e. taylori utilizes C. levisecta as a larval host, it could also have included
parasitism. If we accept the premise that mutualism and parasitism are separately
the two most dangerous interactions for declining species in terms of coextinction risk (Dunn et al. 2009), we can conclude that the continued isolation of
E. e. taylori and C. levisecta is a particularly urgent conservation concern.
TAYLOR’S CHECKERSPOT BUTTERFLY
Euphydryas editha taylori is a non-migratory butterfly species, federally listed as
potentially endangered (2012), which once flourished on glacial outwash prairies,
low elevation grassy balds and coastal grassland sites from southern British
Columbia to central Oregon (Grosboll 2004, Schultz et al. 2011, Severns and
4
Grosboll 2011). The species was first named by W. H. Edwards in 1888 after
Reverend George W. Taylor, one of the first lepidopterists to work in British
Columbia (Shepard and Guppy 2011). Gunder (1929) thereafter described two
other subspecies (E. e. barnesi, and E. e. victoriae) that are now considered
synonymous with E. e. taylori. Phenotypically, E. e. taylori is the darkest editha
subspecies. It has black wings brightly checkered with orange and white spots,
and an average wing span of ca. 4 cm, making E. e. taylori one of the smallest
editha subspecies.
Species status, life history, and restoration challenges
Taylor’s checkerspots were relatively abundant until fairly recently, according to
Pyle (1974) and Dornfield (1980) who said that in western Oregon before 1970
they were known to “swarm by the thousands.” Since 1970, factors such as land
use change, habitat fragmentation, and invasion of remnant prairies by exotic
shrubs and grasses all combined to reduce populations of E. e. taylori to the point
that they were thought to be extinct (Pyle 2002, Severns and Warren 2008). It was
not until they were rediscovered by a junior author during the preparation of The
Butterflies of Cascadia (Pyle 2002) that conservation efforts began. Since then,
efforts to quantify the status of the species (Shepard 2000, Ross 2003, Black and
Vaughan 2005, Stinson 2005) have led to the conclusion that eight known
populations of E. e. taylori continue to persist (Schultz et al. 2011).
Before its decline, E. e. taylori had not been intensively studied, so initial
hypotheses of E. e. taylori habitat needs and host plant interactions were
5
augmented by cautious inference from research done with other E. editha
subspecies. Fortunately, these are some of the most studied butterflies in North
America (Ehrlich and Hanski 2004). For example, work by Weiss, Murphy, and
White (1988) had shown that topographic diversity, and its associated
microclimatic heterogeneity, was an important determinant of habitat quality for
California populations of bay checkerspots (E. e. bayensis). This conclusion was
drawn from several observations. First, on warmer slopes, post-diapause larvae
pupated earlier and pupa developed to eclosion more rapidly than on
progressively cooler slopes. Furthermore, females which eclosed earlier were
more reproductively successful because egg clutches laid earlier tended to be
more successful on a wider variety of slopes than those laid comparatively later.
This was because larval host plants on the sunnier slopes began to senesce in the
latter part of the season, which caused significant mortality in pre-diapause larvae.
Therefore, warmer slopes were advantageous for post-diapause larvae and adults,
but eggs and pre-diapause larvae showed better survivorship on cooler slopes
(Weiss et al. 1988).
Other insights gleaned from previous E. editha research related to
dispersal traits and metapopulation dynamics. Early work by Ehrlich (1961) had
shown that E. editha was similar to other butterflies in that, despite the high
potential vagility (ability to disperse across barriers) associated with flight, their
populations tended to remain fairly sedentary. This understanding about the
difference between potential and actual vagility in E. editha, and the implications
of low actual vagility on gene flow within metapopulations and the species’
6
ability to colonize new sites (or recolonize old ones), led to a four year study of a
single metapopulation at Jasper Ridge, California (Ehrlich 1965). This study
showed that very little gene flow occurred between populations even when there
was no discernible habitat discontinuity separating them. Furthermore,
populations tended to shift spatially very little and did not expand to take
advantage of unutilized resources at the fringe. Later work showed that
populations were subject to relatively frequent extirpations (Ehrlich et al. 1980)
but that the few colonizations of new sites which did occur (Singer and Ehrlich
1979) served to offset this, resulting in a metapopulation which persisted as a
shifting mosaic of relatively isolated subpopulations (Singer and Ehrlich 1979,
Harrison et al. 1988). Singer and Ehrlich (1979) warned of the potential impact of
any factor which decreased the rate at which new sites were successfully
colonized. In light of this previous research on related species, the need for
relatively large contiguous habitat patches could be especially problematic for E.
e. taylori which exists in a highly fragmented landscape (Char and Boersma 1995,
Dunwiddie and Bakker 2011, Schultz et al. 2011).
Another threat to the continued persistence of E. e. taylori is posed by a
shift in plant assemblage on remaining prairies, from native forbs and bunch
grasses to invasive shrubs, forbs, and tall grasses (Stanley et al. 2008, Dunwiddie
and Bakker 2011). This shift may inhibit the ability of E. e. taylori to colonize
new sites and so reduce the functionality of metapopulations. Tall grasses in
particular may be harmful to E. e. taylori persistence (Weiss 1999, Severns and
Warren 2008). Weiss (1999) showed that California populations of E. e. bayensis
7
tended to persist in areas dominated by native grasses, but often crashed shortly
after invasion by taller exotic species. Severns and Warren (2008) showed that
gravid E. e. taylori females chose sites for oviposition that were surrounded by a
higher abundance of native plants and short grasses, as opposed to taller exotics.
In many cases invasion of prairies by exotic plants causes the remaining
natives (if any remain) to persist only on less suitable, more xeric soils (Fimbel
2004), a fact which may exacerbate the effects of invasive plants on E. e. taylori
populations. Even a small shift towards earlier senescence by E. e. taylori’s native
host plants could reduce larval survival, since a major source of mortality for E.
editha larvae is premature host plant senescence (Ehrlich 1961, Mackay 1985,
Grosboll 2011). Even a shift of a single week could mean the difference between
larvae successfully entering diapause or dying in the sun on a desiccated host
plant.
Anticipated effects of regional climate change
Another potential influence on E. e. taylori populations, which may be
increasingly severe in the future, is posed by regional climate change. In a study
of two extirpations of E. e. bayensis in California, McLaughlin et al. (2002) found
that their population declines were more precipitous as a result of increased
variability in precipitation. Global and regional climate models predict that
precipitation variability in the Pacific Northwest will increase (Solomon et al.
2007). McLaughlin et al. (2002) also modeled extant populations of E. e. bayensis
to examine the influences of such a trend, and found that increased precipitation
8
variability caused populations to fluctuate dramatically leading to swift
extinctions, especially when coupled with increased fragmentation of habitat.
Current regional climate models predict a general warming throughout the
WPG of 1.1 °C by the 2020’s and 3.0 °C by the 2080’s (Mote and Salathé 2010).
Precipitation variability is expected to increase, with more winter rainfall but
prolonged summer droughts (Mote and Salathé 2010, Bachelet et al. 2011). The
frequency of extreme weather events is also expected to increase. Potential effects
of these changes to E. e. taylori have not been specifically studied, but could
include increased population variability (McLaughlin et al. 2002), temporal shifts
in life stages or changes in diapause length, or increased mortality from
anomalous weather events.
In addition to direct effects of climate change on E. e. taylori, indirect
effects may also exist from changes to habitat characteristics and host/nectar plant
availability. The effects of regional climate change on WPG prairies have been
predicted by effect simulations and warming experiments. Effect simulations have
been focused primarily on trees, and predict a range shift by P. menzesii
northward (Hamann and Wang 2006) and upward (Rehfeldt et al. 2006, Coops
and Waring 2011). Because of this, it is anticipated that forest encroachment on
lowland WPG prairies will be reduced (Bachelet et al. 2001, Shafer et al. 2001,
Rehfeldt et al. 2006, Littell et al. 2010, Coops and Waring 2011) except in the
northernmost parts of the ecoregion (Hamann and Wang 2006). In grassland sites
across the globe, several warming experiments have shown an overall decline in
plant biodiversity (Zavaleta et al. 2003, Klein et al. 2004, Walker et al. 2006),
9
which corroborates the prediction of decreased competition from P. menzesii, and
also goes a step further and predicts species loss across a wider range of taxa.
Still, plants native to WPG prairies are well adapted to summer droughts and
nutrient-poor soils. Fimbel (2004) found that natives were often able to persist in
marginal conditions unsuitable for exotics, and Pfeifer-Meister et al. (2008) found
that native vs. exotic success was controlled by moisture and nutrient availability.
Therefore, there is the potential that the loss of biodiversity associated with a
warming climate might favor native plants. Complicating the picture, however,
are other experiments that found the negative effects associated with warming
were offset by increases in nutrient availability which might favor invasives
(Shaver et al. 2000, Rustad et al. 2001, de Valpine and Harte 2001, An et al. 2005,
Suttle et al. 2007). Because of these contrasting factors, a summative prediction of
climate change impacts to WPG prairies is difficult to make (Bachelet et al.
2011), but even small changes to host plant availability could have dramatic
effects on E. e. taylori populations.
Habitat restoration
Facing the multitude of challenges listed above, conservation efforts for E. e.
taylori have been underway for over a decade. These efforts have focused on
conservation of existing populations (primarily through invasive plant removal),
restoration of habitat for reintroduction, translocation, and captive breeding.
The restoration of habitat for E. e. taylori has been informed by several
studies. Hays et al. (2000) conducted an extensive survey of two south Puget
10
lowland prairies (Scatter Creek Wildlife Area and Johnson Prairie on Joint-Base
Lewis McChord) to assess habitat characteristics and plant usage by E. e. taylori.
This study provided an early baseline for restoration targets at sites being
prepared for translocation of the butterflies. Previous studies with related species
also help inform E. e. taylori restoration decisions, such as Ehrlich and Murphy
(1987) who reviewed conservation lessons learned from several long-term studies
with E. editha spp. and provided insight on supplying resources for all life stages
in the design of restoration projects. Another study, which informs our
understanding of E. e. taylori’s ideal habitat characteristics, was provided by
Singer (1972) who showed that gopher mounds provided enough microclimatic
variation to increase larval survival in E. e. bayensis. Larval host plants growing
on the mounds resisted summer drought longer than those growing in the
intermound space, and provided a mechanism for larval survival in dry years.
These findings may be relevant for E. e. taylori because Mazama pocket gophers
(Thomomys mazama) historically occupied a similar range of Puget lowland
prairies, and currently co-exist with E. e. taylori in the location of its largest
extant population. Mazama pocket gophers are also candidates for listing under
the Endangered Species Act.
Another important habitat characteristic which E. e. taylori has adapted to
is the presence of fire. Western Washington prairies were burned by Native
Americans nearly every year, for about 15,000 years, prior to the arrival of
European settlers in the middle of the 19th century (Morris 1934, Lang 1961,
Norton 1979, Leopold and Boyd 1999, Fimbel 2004, Storm and Shebitz 2006).
11
Humans burned prairies annually in many locations to increase the availability of
edible forbs, and every few years in other places to increase the abundance of
berries (Norton 1979, Fimbel 2004, Storm and Shebitz 2006). This adaptation of
E. e. taylori to fire-altered habitats may be partly responsible for the unusual
location of its largest extant population: the Artillery Impact Area (AIA) at Joint
Base Lewis-McChord (Linders 2012). Fires still burn the prairie nearly every
summer on the AIA, set by practice shelling with explosive ordinance (Tveten
1997). Other factors which may also contribute to E. e. taylori persistence at this
site include the sheer size (> 3000 ha) of the habitat fragment (MacArthur 1967,
Quammen 2012), as well as the presence of Mazama pocket gophers (Stinson
2005). Also, the restriction against development and recreational use of military
lands may reduce other negative human influences on E. e. taylori populations.
Endangered butterflies living on a valuable army training asset is a
strangely beneficial relationship for E. e. taylori recovery efforts. Department of
Defense biologists are tasked with species conservation and restoration on federal
lands. A dedicated staff of these biologists and ecologists is employed at JBLM
and is actively engaged in on-site restoration on a year-round basis. Furthermore,
a grant program, called the Army Compatible Use Buffer (ACUB) program,
focuses on purchasing and restoring non-military land in the vicinity of military
bases. One of the stated purposes of the ACUB program is reducing the negative
influences of training activities on imperiled species, and it has been instrumental
in supporting E. e. taylori captive rearing and translocation efforts in the south
Puget lowlands (Linders 2012).
12
Translocation and captive breeding
When recovery efforts began for E. e. taylori, there were two populations that
were considered robust enough to serve as sources of individuals for captive
breeding and translocation. These were the AIA at JBLM as mentioned above and
another site in western Washington, a series of grassy knolls called the Bald Hills
(Linders 2007). The Bald Hills population is now considered extirpated (Grosboll
2011) leaving only one source population range-wide.
Nine unoccupied sites were initially considered as potential restoration
areas for experimentally reintroducing E. e. taylori (Linders 2007). These
included sites both on and off the JBLM military base. To date, E. e. taylori
releases have occurred at four sites (Linders 2012): Scatter Creek Wildlife Area
(since 2007), Range 50 on the AIA at JBLM (2009-2011), Pacemaker on 13th
Division Prairie at JBLM (2012), and Glacial Heritage Preserve (since 2012).
Because of the need for an increasing number of animals for release, and
the uncertainty of the source population, effort has also been focused on the
development of captive breeding methods (Grosboll 2004, Linders 2007, 2012,
Barclay et al. 2009). Captive breeding began with a pilot project by Grosboll
(2004) who attempted to rear 126 eggs collected from a wild female using two
different host plants, lance-leaf plantain (Plantago lanceolata L.) and harsh
paintbrush (Castilleja hispida Benth.). His results showed no difference from
hatching to diapause, but better survival from diapause to eclosion for the C.
hispida group. Efforts were moved to the Oregon Zoo in 2004, where a successful
standardized protocol has been developed (Barclay et al. 2009). A second captive
13
breeding institution was added to the project in 2012, in association with the
Sustainability in Prisons Project, at Mission Creek Corrections Center for
Women. This facility was able to build on the work accomplished by the Oregon
Zoo in achieving a 96.6% egg to diapause survivorship in their first year of
operation. With both facilities operating successfully, the largest release to date
was held in 2013, when over 6000 animals were released at Scatter Creek
Wildlife Area and Glacial Heritage Preserve.
Oviposition host plants
Ecological restoration to prepare sites for reintroduction of E. e. taylori is
underway, but conservation planners are uncertain which plants were historically
the butterfly’s most important larval hosts (Severns and Warren 2008). At present,
E. e. taylori primarily utilizes the introduced exotic, P. lanceolata for oviposition.
One reason for this is that E. e. taylori utilizes an unpalatability defense, by
ovipositing selectively on plants which contain iridoid glycosides (Grosboll 2004,
Schultz et al. 2011). Upon hatching, the larvae consume these monoterpenes,
sequestering them in their tissues and ultimately discouraging predation
throughout their life cycle (Bowers 1981). Plantago lanceolata contains iridoid
glycosides, as do the other native plants E. e. taylori is known to oviposit on, such
as C. hispida, shortspur seablush (Plectritis congesta (Lindl.) D.C.), and maiden
blue-eyed Mary (Collinsia parviflora Lindl.). Despite having suitable native
hosts, however, E. e. taylori has come to be almost completely dependent on P.
14
lanceolata due its spatial density and abundance, and the decreasing abundance of
the native species (Severns and Warren 2008).
The fact that E. e. taylori has switched hosts to a potentially invasive
exotic plant presents a problem for restoration management. The idea of
introducing a noxious weed to otherwise high quality native prairie is unpopular,
but other options are few. The native plants known to be occasional larval hosts to
E. e. taylori typically senesce on WPG prairies before the larvae are able to
successfully enter diapause (Mary Linders, personal communication). In contrast
to the natives, P. lanceolata tolerates drought quite well and often persists
throughout the summer.
GOLDEN PAINTBRUSH
One native plant which shows promise as a larval host is another federallythreatened species, golden paintbrush (Castilleja levisecta Greenm.). This iridoid
glycoside-producing perennial shares with E. e. taylori approximate historic range
(Wentworth 2001, Lawrence and Kaye 2008, 2011), preferred habitat type, and
reasons for population decline.
Castilleja levisecta was first collected in 1875 in Victoria, B.C., and was
first described by J. N. Greenman in 1898. Historically, it has been collected from
over 30 sites, but by 1981 it had declined in abundance and was listed on the first
publication of the Washington Natural Heritage Program’s list of endangered
species. Field surveys by Sheehan and Sprague (1984) and Evans, Schuller, and
Augenstein (1984) quantified how rare it had become, which led to its 1997
15
federal threatened-species listing. Following this, Wentworth (1994) further
studied the phenology and life history traits of C. levisecta. Unfortunately, the
species was already extirpated from most of its range before Wentworth did his
research, so the true variability of its phenology and preferred habitat
characteristics have been difficult to estimate (Gammon 1995, Lawrence and
Kaye 2006).
Currently, there are 11 isolated populations of C. levisecta, ten of which
are in the San Juan Islands and British Columbia (Lawrence and Kaye 2008,
2011). Additionally, there are no co-occurring populations of C. levisecta and E.
e. taylori. Because of this, it is unknown if E. e. taylori will oviposit on C.
levisecta, but it is recognized that in several ways it might be highly suitable. For
example, C. levisecta occupies slightly more hydric microsites than the known
native hosts and it often persists well into the summer months (Wentworth 2001).
Additionally, its growth form may provide more available biomass for larval
consumption than C. hispida, so it might be able to host larger populations per
plant. These lines of evidence, combined with the fact that the timing of the two
species’ decline has been relatively coincident, have led to speculation that C.
levisecta could be an ancestral host for E. e. taylori (Stinson 2005).
Despite the apparent suitability C. levisecta as a larval host, it has not been
experimentally reintroduced to E. e. taylori habitat sites because congeneric C.
hispida is actively planted and the two could hybridize if grown together
(Lawrence and Kaye 2008). Research is currently ongoing to address the question
of Castilleja spp. hybridization rates, but until this is known the only option
16
would be to replace C. hispida with C. levisecta at restoration sites, and that
represents too much of a risk without verification of the suitability of C. levisecta
as a host plant. However, C. levisecta is currently reintroduced at separate
restoration sites, and if it could be shown that E. e. taylori will select it for
oviposition, the two restoration efforts might be joined. This has the potential to
increase the effectiveness of both C. levisecta and E. e. taylori recovery efforts,
and is the subject of this thesis.
17
ARTICLE MANUSCRIPT
Formatted for submission to Northwest Science
Oviposition preference by Taylor’s checkerspot butterfly (Euphydryas editha
taylori) among lance-leaf plantain (Plantago lanceolata), harsh paintbrush
(Castilleja hispida), and golden paintbrush (Castilleja levisecta)
ABSTRACT
Taylor’s checkerspot butterfly (Euphydryas editha taylori) is a federally
threatened pollinator of increasingly rare prairies in the Willamette Valley-Puget
Trough-Georgia Basin ecoregion. Since the arrival of European settlers, several
factors have helped reduce available native host plants for E. e. taylori larvae. The
most common host is now Plantago lanceolata, an exotic species long prevalent
in the area. None of the known native hosts are ideal for supporting E. e. taylori
restoration. Federally threatened Castilleja levisecta may have been important
historically but does not now co-occur with E. e. taylori. Previous work has
shown that oviposition preference is: 1) heritable and may provide clues to which
hosts were historically important, and 2) correlated with larval success so might
indicate which hosts would be most effective for restoration. We undertook an
oviposition preference experiment to determine which potential hosts were
preferred by E. e. taylori among P. lanceolata, C. levisecta, and C. hispida. The
two Castilleja spp. were preferred equally and both were preferred over P.
lanceolata. If further research confirms the suitability of C. levisecta as a host for
E. e. taylori, restoration efforts for the two species could be united, and the
effectiveness of both might be synergistically increased.
INTRODUCTION
The prairies and oak savannas of the Willamette Valley-Puget Trough-Georgia
Basin (WPG) ecoregion are increasingly rare and are recognized as one of the
most endangered ecosystem types in North America (Noss et al. 1995, Floberg et
al. 2004, Stanley et al. 2008, Dunwiddie and Bakker 2011). Land use changes,
habitat fragmentation and invasion by exotics have all contributed to the decline
18
of both the extent and functionality of these habitats (Fimbel 2004, Grosboll 2004,
Stanley et al. 2008), and many obligate species have become imperiled as a result
(Dennehy et al. 2011, Hamman et al. 2011, Schultz et al. 2011, Wold et al. 2011).
Among these are the Taylor’s checkerspot butterfly (Euphydryas editha taylori),
and golden paintbrush (Castilleja levisecta).
Euphydryas editha taylori is a non-migratory butterfly that has been
federally listed as a potentially endangered species (2012). It once flourished on
glacial outwash prairies, low elevation grassy balds and coastal grassland sites
from southern British Columbia to central Oregon (Grosboll 2004, Schultz et al.
2011, Severns and Grosboll 2011). However, in recent decades habitat loss and
degradation have reduced it to only eight isolated populations (Stinson 2005,
Schultz et al. 2011). Exacerbating the threat to E. e. taylori populations is the
declining presence of native forbs (historically the most important food and nectar
plants for the species) on remnant prairies (Stanley et al. 2008, Dunwiddie and
Bakker 2011), which inhibits the ability of E. e. taylori to recolonize the parts of
its range from which it has been extirpated. Also, many native prairie plants are
now restricted to more xeric sites due to competition with invasive exotics
(Fimbel 2004), which may be problematic because early host plant senescence has
been shown as a primary source of mortality for pre-diapause Euphydryas editha
larva (Mackay 1985).
At present, E. e. taylori primarily utilizes the introduced exotic, lance-leaf
plantain (Plantago lanceolata L.) for oviposition. One reason for this is that E. e.
taylori utilizes an unpalatability defense, by ovipositing selectively on plants
19
which contain iridoid glycosides (Grosboll 2004, Schultz et al. 2011). Upon
hatching, the larvae consume these monoterpenes, sequestering them in their
tissues and ultimately discouraging predation throughout their life cycle (Bowers
1981). Plantago lanceolata contains iridoid glycosides, as do the three native
plants that E. e. taylori is known to oviposit on harsh paintbrush (Castilleja
hispida Benth.), shortspur seablush (Plectritis congesta (Lindl.) D.C.), and
maiden blue-eyed Mary (Collinsia parviflora Lindl.), yet due to the decreasing
abundance of the native species and the spatial density of P. lanceolata patches,
E. e. taylori has come to be almost completely dependent on this exotic plant
(Severns and Warren 2008).
The fact that E. e. taylori has switched hosts to a potentially invasive
exotic plant presents a problem for restoration management. The introduction of
potentially invasive P. lanceolata to otherwise high quality native prairie is
problematic for land managers, but few other options exist at present. The native
plants known to be occasional larval hosts to E. e. taylori typically senesce on
WPG prairies before E. e. taylori larvae are able to successfully enter diapause
(Mary Linders, personal communication). In contrast to the natives, P. lanceolata
often persists throughout the summer.
An ideal host plant for E. e. taylori would, 1) contain an suitable array of
iridoid glycosides to be chosen for oviposition, 2) provide enough biomass to
support prediapause larva into the third instar when they begin to disperse, and 3)
persist long enough to allow larvae to successfully enter diapause. The only plant
20
currently available to E. e. taylori on WPG prairies, which possesses all three of
these qualities, is P. lanceolata.
One native plant which shows promise as a larval host is another
federally-threatened species, golden paintbrush (Castilleja levisecta). It is an
iridoid glycoside-producing perennial which shares with E. e. taylori approximate
historic range (Wentworth 2001, Lawrence and Kaye 2008, 2011), preferred
habitat types, and reasons for population decline. Currently there are no cooccurring populations of C. levisecta and E. e. taylori. Because of this, it is
unknown if E. e. taylori will oviposit on C. levisecta, but it is recognized that in
several ways it might be highly suitable. For example, C. levisecta occupies
slightly more mesic microsites than the known native hosts and it often persists
well into the summer months (Wentworth 2001). Additionally, the growth form of
C. levisecta may provide more available biomass for larval consumption than C.
hispida, so it might be able to host larger populations per plant. These
characteristics of C. levisecta, in addition to the relatively coincident decline of
both species, has led to speculation that C. levisecta could be an ancestral host for
E. e. taylori (Stinson 2005).
Despite its potential suitability, C. levisecta has not been experimentally
reintroduced to E. e. taylori habitat sites because congeneric C. hispida is actively
planted at these locations and the two species could hybridize if grown together
(Lawrence and Kaye 2008). Research is currently ongoing to address the question
of hybridization rates, but until these are known, the reintroduction of C. levisecta
at E. e. taylori restoration sites would require replacing C. hispida entirely, and
21
that may present too much of a risk prior to verification of its suitability as a host
plant. In the interim, C. levisecta is reintroduced at separate restoration sites;
however, if it could be shown that E. e. taylori will select C. levisecta for
oviposition, the two recovery efforts might be joined, potentially increasing the
effectiveness of both efforts.
To assess the viability of C. levisecta as a host plant, we undertook a
manipulative oviposition preference study to compare the likelihood of it being
selected by E. e. taylori from among C. hispida and P. lanceolata. Our hypotheses
were that 1) the two native Castilleja spp. would be preferred over P. lanceolata
due to their historical coexistence with E. e. taylori, and 2) C. levisecta would be
the most preferred overall due to its potential suitability as a host plant and the
possibility of an unknown historical interaction.
METHODS
Site description
Research was conducted in a purpose-built butterfly breeding facility at Mission
Creek Corrections Center for Women (MCCCW) in Belfair, Washington, USA,
under the guidance of the Sustainability in Prisons Project (SPP). The SPP is a
collaboration between The Evergreen State College (TESC) and the Washington
Department of Corrections (WDOC) which seeks to engage incarcerated men and
women in science, conservation, and sustainability, in order to reduce the
environmental, social, and human costs of prisons (LeRoy et al. 2012).
22
The butterfly facility is a 3 x 7.3 m partitioned greenhouse with UVtransmitting glass panels. Research was conducted in the smaller of two rooms (3
x 2.4 m) while normal rearing and breeding activities were confined to the larger.
Trained inmate butterfly technicians performed all research activities, with daily
oversight by a graduate and two undergraduate students. In keeping with the
SPP’s mission, inmate technicians were engaged as collaborators and were
involved in many phases of the work, including: planning, methods refining, data
collection, and manuscript review.
Data Collection
Methods for testing oviposition preference were developed with California
populations of E. editha (Singer 1982, Singer et al. 1991, 1992), and with
Melitaea cinxia (Singer and Lee 2000). These methods were designed to compare
preference between just two potential hosts. We made pairwise comparisons
among C. levisecta, C. hispida, and P. lanceolata for a total of three complete
comparison sets. Each comparison involved 10 individual trials, each with a
different butterfly selected randomly from five different captively-reared lineages.
Butterflies were F1 descendants of individual wild-caught females,
collected from Range 76 on Joint Base Lewis-McChord in 2011. This site hosts
the species’ largest extant population, and currently supports the collection of all
captive colony founders.
In general, oviposition preference testing is possible because the
butterflies display behaviors that indicate selection prior to oviposition (Singer
23
1982). Upon alighting on a potential host, the butterflies taste for the presence of
iridoid glycosides with specially adapted fortarsi. If conditions are not adequate
they will not oviposit and move to another location. Butterflies often decline or
accept different individuals of the same species, apparently selecting for an
unknown but specific chemical signature. If the butterfly approves of the potential
host’s alkaloid signature, she may tap her antennae or wave her wings in further
investigation before finally curling her abdomen and touching her ovipositor to
the plant, indicating final oviposition selection.
Following Singer (1982, Singer et al. 1991, 1992, Singer and Lee 2000),
oviposition preference was determined by sequentially offering individual gravid
females (n=30), test plant individuals (n=3) of two species per comparison
(Figure 1). Each plant was contained within a small screen enclosure. The
butterflies, which were not initially motivated to oviposit, were placed into an
enclosure with a randomly chosen plant and observed for five minutes for
oviposition behavior, before being offered the next plant. After a positive
attempted oviposition, which was denoted by the female touching her ovipositor
to a leaf surface for two seconds, she was placed in an empty enclosure for five
minutes before being offered the next plant. Testing sessions continued until
females chose all six plants. They were then allowed to oviposit completely in a
separate enclosure, on the plant of highest preference.
Three plants of each species were used in each trial to account for withinspecies variation in alkaloid signatures. Plants were randomly selected for each
trial from pools of 80 individuals. The plants were all in reasonably good health,
24
Figure 1 – Details of oviposition preference testing methods carried out at the
endangered butterfly rearing facility at Mission Creek Corrections Center for
Women, Belfair, Washington, USA
and each selected individual was used only in a single trial. Plants were all from
south Puget lowland genetic stock, and were grown under similar conditions.
These plants were propagated from the same populations used for Taylor’s
checkerspot restoration efforts. The butterflies themselves are also from the same
lineages that are reared for release on south Puget lowland prairie restoration
sites.
25
Throughout the trials, temperature was maintained within a range of 24-32
°C. Other environmental variables were assumed to be relatively standardized due
to the randomized pairings. Over 1200 individual five-minute trials were
performed during the course of the study.
Statistical Analysis
Preference between potential hosts for each butterfly was assessed by averaging
the rank of acceptance. For example if, during a trial between C. levisecta and C.
hispida, C. levisecta was selected first, second, and fourth, it would be given a
score of 2.33. We call this statistic the mean selection rank, and lower selection
ranks indicate higher preference. Additionally, accepted plants were only
considered preferred if the next plant offered was declined, so plants accepted
sequentially were given the same averaged rank, creating the possibility of a
given trial resulting in no preference shown for either species. Overall preference
between plant species was determined by comparing the overall means of all three
comparisons using a one-way ANOVA and Tukey’s HSD test.
RESULTS
Comparison of mean selection rank (Figure 2) showed that E. e. taylori did not
equally prefer the three plants (F(2,27) = 18.02, p<0.0001). Post-hoc tests revealed
that the two Castilleja spp. were preferred equally, but each was preferred over P.
lanceolata.
26
Figure 2 – Mean oviposition preference among potential host plants. Each
trial resulted in a mean selection rank based on selection order of the six
plants offered, resulting in a score between two and five with low scores
indicating higher preference.
The record of individual trial results (Figure 3) shows that for the 10
butterflies offered the C. levisecta and P. lanceolata pairing, nine preferred C.
levisecta and one showed equal preference for both. Between C. hispida and P.
lanceolata, seven preferred C. hispida, one preferred P. lanceolata, and two
showed equal preference for both. Between the two Castilleja spp., four preferred
C. levisecta, three preferred C. hispida, and three showed equal preference for
both.
27
# of Preference Selections
10
8
tie
6
PLLA
4
CAHI
CALE
2
0
CALE vs. PLLA
CAHI vs. PLLA
CALE vs. CAHI
Comparison Pairing
Figure 3 – Results of the 10 trials in each oviposition preference
comparison pairing.
DISCUSSION
Our results not only show that E. e. taylori will select C. levisecta for oviposition,
but that its preference for C. levisecta is equal to its preference for C. hispida, the
most suitable of the currently known native host plants. Adding another suitable
native host to the E. e. taylori reintroduction site planting mix may increase the
effectiveness of recovery efforts.
Both Castilleja spp. were preferred over P. lanceolata despite the fact that
the butterflies had consumed P. lanceolata exclusively as larvae. This can be
explained by research with the conspecific bay checkerspot (Euphydryas editha
bayensis), which found that oviposition preference was heritable (Singer 1988).
Singer’s research with E. e. bayensis (1988) also showed that oviposition
preference was correlated with offspring growth rate, indicating that a mother’s
use of her preferred oviposition host conferred an advantage to her offspring.
28
Since the two Castilleja spp. in our study were preferred over P. lanceolata, it is
possible that E. e. taylori is at a disadvantage when utilizing P. lanceolata, its
most common larval host.
The observed trend in oviposition preference of C. levisecta over C.
hispida, as indicated by mean selection rank, was non-significant. However, the
mean score for C. levisecta was higher than for C. hispida, a trend corroborated
by the record of individual trial results. Because of this, we suspect that a larger
sample size might have shown C. levisecta to be the most preferred host overall;
further inquiry will be required to investigate this.
More work also needs to be done in the field to determine the efficacy of
C. levisecta as a larval host, but if it could be utilized in E. e. taylori
reintroduction site plantings, either alongside or in place of C. hispida, the
effectiveness of two threatened species recovery efforts might be synergistically
increased. The potential exists to provide a valuable native host to E. e. taylori
while also creating more planting sites and a more robust metapopulation for C.
levisecta. In an age of ambitious conservation goals and limited resources,
efficiency is paramount to overall success.
In a novel collaboration, we have found that the effective employment of
non-traditional partners, in this case incarcerated women, can also help increase
the available resources for conservation and improve overall efficiency. We hope
that our project opens doors to more collaborative conservation science in
correctional facilities and other non-traditional environments in the future.
29
Acknowledgments
The butterfly facility at Mission Creek Corrections Center for Women is managed
by the Sustainability in Prisons Project and Washington Department of
Corrections staff, overseen by the Washington Department of Fish and Wildlife,
assisted by the endangered butterfly lab at Oregon Zoo (Portland, Oregon), and
funded by a US Fish and Wildlife Grant through the US Army Compatible Use
Buffer program. We thank these partners, The Evergreen State College, and
WDOC administration for their support of both this research and the overarching
vision of science in prisons. We also thank the individuals that helped make this
research a reality: Kelli Bush, Joslyn Trivett and Carl Elliott from SPP,
undergraduate interns Caitlin Fate and Chelsea Oldenburg from TESC,
Superintendent Wanda McRae, Anne Shoemaker, Leo Gleason, Sherri Albrecht,
and Dan Davis from MCCCW, Ted Thomas of the U.S. Fish and Wildlife
Service, Mary Jo Andersen and Karen Lewis of the Oregon Zoo, and and five
inmates, who will remain unnamed, that supported and participated in gathering
these data..
REFERENCES
Bowers, M. D. 1981. Unpalatability as a defense strategy of western checkerspot
butterflies (Euphydryas scudder, Nymphalidae). Evolution 35:367–375.
Dennehy, C., E. R. Alverson, H. E. Anderson, D. R. Clements, R. Gilbert, and T.
N. Kaye. 2011. Management strategies for invasive plants in pacific
30
northwest prairies, savannas, and oak woodlands. Northwest Science
85:329–351.
Dunwiddie, P. W., and J. D. Bakker. 2011. The future of restoration and
management of prairie-oak ecosystems in the Pacific Northwest.
Northwest Science 85:83–92.
Fimbel, C. 2004. Habitat enhancement for rare butterflies on Fort Lewis prairies.
Report to The Nature Conservancy, South Sound Office, Olympia
Washington.
Floberg, J., M. Goering, G. Wilhere, C. MacDonald, C. Chappell, C. Rumsey, Z.
Ferdana, A. Holt, P. Skidmore, T. Horsman, E. Alverson, C. Tanner, M.
Bryer, P. Lachetti, A. Harcombe, B. McDonald, T. Cook, M. Summers,
and D. Rolph. 2004. Willamette Valley-PugetTrough-Georgia Basin
ecoregional assessment. The Nature Conservancy with support from the
Nature Conservancy of Canada, Washington Department of Fish and
Wildlife, Washington Department of Natural Resources (Natural Heritage
and Nearshore Habitat programs), Oregon State Natural Heritage
Information Center and the British Columbia Conservation Data Centre.
Grosboll, D. N. 2004. Captive rearing the endangered mardon skipper (Polites
mardon) and Taylor’s checkerspot (Euphydryas editha taylori) butterflies:
Initial results (Lepidoptera, Nymphalidae). Proceedings of the Species at
Risk Pathways to Recovery Conference, Victoria, B. C.
Hamman, S. T., P. W. Dunwiddie, J. L. Nuckols, and M. McKinley. 2011. Fire as
a restoration tool in Pacific Northwest prairies and oak woodlands:
31
Challenges, successes, and future directions. Northwest Science 85:317–
328.
Lawrence, B. A., and T. N. Kaye. 2008. Direct and indirect effects of host plants:
Implications for reintroduction of an endangered hemiparasitic plant
(Castilleja levisecta). Madroño 55:151–158.
Lawrence, B. A., and T. N. Kaye. 2011. Reintroduction of Castilleja levisecta:
Effects of ecological similarity, source population genetics, and habitat
quality. Restoration Ecology 19:166–176.
LeRoy, C. J., K. Bush, J. Trivett, and B. Gallagher. 2012. The Sustainability in
Prisons Project: An overview (2004-12). Gorham Printing, Olympia, Wa.
Mackay, D. A. 1985. Conspecific host discrimination by ovipositing Euphydryas
editha butterflies: Its nature and its consequences for offspring
survivorship. Researches on Population Ecology 27:87–98.
Noss, R. F., E. T. LaRoe III, and J. M. Scott. 1995. Endangered ecosystems of the
United States: A preliminary assessment of loss and degradation. Report
to the US Department of the Interior, National Biological Service,
Washington D. C.
Schultz, C. B., E. Henry, A. Carleton, T. Hicks, R. Thomas, A. Potter, M. Collins,
M. Linders, C. Fimbel, S. Black, H. E. Anderson, G. Diehl, S. Hamman,
R. Gilbert, J. Foster, D. Hays, D. Wilderman, R. Davenport, E. Steel, N.
Page, P. L. Lilley, J. Heron, N. Kroeker, C. Webb, and B. Reader. 2011.
Conservation of prairie-oak butterflies in Oregon, Washington, and British
Columbia. Northwest Science 85:361–388.
32
Severns, P. M., and D. Grosboll. 2011. Patterns of reproduction in four
Washington State populations of Taylor’s checkerspot (Euphydryas editha
taylori) during the spring of 2010. Report to The Nature Conservancy,
Olympia, Washington.
Severns, P. M., and A. D. Warren. 2008. Selectively eliminating and conserving
exotic plants to save an endangered butterfly from local extinction.
Animal Conservation 11:476–483.
Singer, M. C. 1982. Quantification of host preference by manipulation of
oviposition behavior in the butterfly Euphydryas editha. Oecologia
52:224–229.
Singer, M. C., and J. R. Lee. 2000. Discrimination within and between host
species by a butterfly: Implications for design of preference experiments.
Ecology Letters 3:101–105.
Singer, M. C., D. Ng, and R. A. Moore. 1991. Genetic variation in oviposition
preference between butterfly populations. Journal of Insect Behavior
4:531–535.
Singer, M. C., D. Ng, and C. D. Thomas. 1988. Heritability of oviposition
preference and its relationship to offspring performance within a single
insect population. Evolution 42:977-985.
Singer, M. C., D. Vasco, C. Parmesan, C. D. Thomas, and D. Ng. 1992.
Distinguishing between “preference” and “motivation” in food choice: An
example from insect oviposition. Animal Behaviour 44, Part 3:463–471.
33
Stanley, A. G., T. N. Kaye, and P. W. Dunwiddie. 2008. Regional strategies for
restoring invaded prairies. Native Plants Journal 9:247 –254.
Stinson, D. W. 2005. Draft status report for the Mazama pocket gopher, streaked
horned lark, and Taylor’s checkerspot. Report to the Washington
Department of Fish and Wildlife, Olympia, Washington.
Wentworth, J. B. 2001. The demography and population dynamics of Castilleja
levisecta, a federally threatened perennial of Puget Sound grasslands. In R.
S. Reichard, P. Dunwiddie, J. Gamon, A. Kruckeberg, and D. Salstrom,
editors. Conservation of Washington’s Native Plants and Ecosystems.
Washington Native Plant Society, Seattle, Washington.
Wold, E. N., J. E. Jancaitis, T. H. Taylor, and D. M. Steeck. 2011. Restoration of
agricultural fields to diverse wet prairie plant communities in the
Willamette Valley, Oregon. Northwest Science 85:269–287.
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DISCUSSION & BROADER IMPACTS
ECOLOGICAL IMPLICATIONS
Taylor’s checkerspot
One of the problems facing E. e. taylori conservation is a lack of firm
understanding surrounding which plants are the species’ most important native
oviposition hosts. Currently the butterfly almost exclusively utilizes P. lanceolata,
which is a potentially invasive exotic species, for oviposition. Several native
plants are known to be occasional larval food sources, but oviposition hosts are
more strategically valuable than other food plants. By laying their eggs in
clutches, E. e. taylori females are essentially choosing individual plants which
will support her young through their first few instars. The literature is unclear
about which plant species were historically the most important for E. e. taylori, in
part because P. lanceolata has been abundant on WPG prairies for so long. It was
reportedly common around Fort Vancouver by 1825 (Nisbet 2009). The historical
and current abundance of P. lanceolata, and the degree to which it is utilized,
make it difficult to describe E. e. taylori’s most important native plant
interactions.
Another reason it is difficult to understand the historical web of native
species interactions is the lack of some plant species, like Castilleja levisecta, that
once occupied similar historic ranges with E. e. taylori, but now do not. Many
species have been impacted by the loss, fragmentation, and degradation of WPG
prairie habitat.
35
It is likely we will never know for sure what the most important native
plants were for E. e. taylori prior to the arrival of P. lanceolata, but the results of
this research may provide clues to the mystery. Both Castilleja spp. were
preferred for oviposition over P. lanceolata. It may be that these were important
ancestral host plants for E. e. taylori.
Because the prairies with the richest soils were typically the first to be
converted to agriculture, many native plants have been pushed to the edges of
their preferred range. It may be that the Castilleja spp. are less suitable as larval
hosts for E. e. taylori when growing on more xeric soils. Early host plant
senescence has been shown to be a primary cause of mortality for pre-diapause E.
e. bayensis larvae (Ehrlich 1961, Mackay 1985, Grosboll 2011), and in some
cases small variations in soil moisture have been shown to have large impacts on
larval survival (Singer 1972, Weiss et al. 1988). Native host plants are more likely
to senesce before the caterpillars are able to safely enter diapause if they are
growing in marginal habitat. Because of this, it could be argued that P. lanceolata
may have been both bane and boon to E. e. taylori’s persistence. It is a bane in
that exotic species are influential in pushing natives to the fringe in the first place,
but a boon in being an acceptable surrogate for E. e. taylori larvae in the absence
of suitable native host populations.
The suitability of P. lanceolata as a larval host stems from three factors.
First, it is much less prone to desiccation than any of the known native hosts. It
often persists well into autumn, so it is almost guaranteed not to senesce before E.
e. taylori larvae enter diapause in early July. Second, it contains iridoid
36
glycosides, similar to those found in the known native hosts, which E. e. taylori
sequester in their tissues to maintain their unpalatability defense. Third, P.
lanceolata individuals are well distributed and tend to exist in dense enough
populations that they both: 1) are relatively likely to be found by gravid E. e.
taylori females, and 2) can support lots of hungry dispersing caterpillars.
Despite the fact that P. lanceolata provides key resources for E. e. taylori,
there are problems associated with planting it at prairie restoration sites. Plots
designated for reintroduction of the butterfly are typically stocked with known
larval hosts and nectar plants, but the idea of planting a potentially invasive exotic
species on high quality native prairie is problematic. Since invasive behavior is
often triggered by the crossing of an unknown population threshold (Crooks and
Soule 2001, Sakai et al. 2001), land managers must be cautious in deciding how
many P. lanceolata plants represent the ideal balance between function and risk.
Our results show that C. levisecta is preferred by E. e. taylori for
oviposition over P. lanceolata. If future research continues to suggest that C.
levisecta would be a valuable addition to the suite of native plants used in E. e.
taylori recovery efforts, it might be possible to reduce the number of P.
lanceolata individuals needed at reintroduction sites. This, in turn, could reduce
the likelihood of P. lanceolata crossing a hidden population threshold at any
given site and initiating outbreak conditions. Although P. lanceolata is known to
be invasive in a wide variety of habitat types, including south Puget lowland
prairies, it might be possible to slow its spread if planting densities are kept low.
37
Golden paintbrush
Like E. e. taylori, C. levisecta is a federally threatened species. Also like E. e.
taylori, it has a recovery plan, restoration sites, and facility-based cultivation
projects for generating reintroduction stock.
Being a known and potentially valuable larval host for E. e. taylori may
benefit C. levisecta. If C. levisecta is utilized in restoration plantings for the
butterfly, it could increase the total number of C. levisecta plugs planted every
year, and increase the total number of its recovery sites. Furthermore, the species
interaction between E. e. taylori and C. levisecta may increase public awareness
of both species, as people interested in one will be more likely to learn about the
other and the synergy the two share. Public awareness, in turn, is critical in
determining funding priorities and community support.
Synergy in conservation may also have value of its own accord. As with
most activities that do not generate profit, ecological restoration and threatened
species conservation are limited by funding and volunteer support. If two
threatened species can be conserved synergistically, such limiting resources can
be used more efficiently, and the net effort can be made more effective. These
outcomes could lead to more total effort for the two species in question, or it
could preserve resources for use by other projects.
38
COLLABORATION
Sustainability in Prisons Project
The Sustainability in Prisons Project (SPP) is a partnership between The
Evergreen State College (TESC) and the Washington Department of Corrections
(WDOC). It strives to involve incarcerated persons in science, sustainability, and
conservation, while engaging them as colleagues and stakeholders, for the sake of
ecological and social restoration.
The SPP began in 2004 with a science and sustainability lecture series that
spawned several sustainability projects at Cedar Creek Corrections Center
(CCCC), a minimum-security men’s prison near Littlerock, Washington. Waste
sorting, composting, recycling, gardening, rainwater recapture, and even a green
roof project, all began from the inspiration of the SPP lecture series. Many of
these ideas then quickly caught on at other Washington prisons, first when local
media started covering the efforts, and then even more so when it became known
how much money CCCC was saving.
The sustainability initiatives inspired by the SPP lecture series are now for
the most part carried forward under the broader umbrella of WDOC Sustainable
Operations (in partnership with SPP). Direction for these measures happens at
both the statewide and the individual facility level. Sustainability in Washington
prisons has taken a life of its own, and is one reason why the state is now
recognized as a world leader in the greening of corrections (LeRoy et al. 2012).
Between 2005 and 2010, WDOC reduced solid waste to landfills by 35%,
increased diversion to recycling by 89%, increased composting operations by
39
90%, decreased potable water use by over 100 million gallons annually, reduced
transportation fuel consumption by 25%, and reduced total carbon emissions by
approximately 40%. In addition, in 2010 prison gardens and farms yielded over
123,000 kg of produce for consumption by inmates and donations to food banks
(LeRoy et al. 2012).
Meanwhile, the science and sustainability lecture series continues and has
expanded to five prisons. Bringing informal science and environmental education
into prisons remains a priority for the SPP. More than 100 lectures and 26
workshops have been held at five prisons, involving 2400 inmates and 280
WDOC staff attendees (LeRoy et al. 2012).
In 2008, another SPP program was added. A partnership with the
Washington Department of Fish and Wildlife (WDFW) allowed CCCC to become
a rearing institution for Washington State endangered Oregon spotted frogs (Rana
pretiosa). These were reared for release onto wetland sites at Joint Base LewisMcChord, with professional biologists from other rearing institutions such as the
Woodland Park Zoo and the Oregon Zoo working collaboratively with inmates
from CCCC. When the project began, there was some debate about whether the
inmates would be able to match the success of professional rearing institutions,
but those doubts were soon erased. The CCCC frog program had the highest
survivorship and most developed frogs of any institution, and was named “best
rearing facility” in 2009, 2010, and 2011. Additionally, inmates participated in
conducting relevant research including a growth comparison between two distinct
40
frog populations with WDFW, and a predator evasion response experiment with
the Oregon Zoo.
The following year in 2009, the SPP began a rare and endangered prairie
plant propagation program at Stafford Creek Corrections Center (SCCC), a
medium-security men’s facility near Aberdeen. It employs up to 10 inmates and
has produced over 600,000 plants for south Puget Sound restoration sites both on
and off JBLM. The SPP recently doubled the capacity of this program by adding a
similar program at the Washington Corrections Center for Women (WCCW), a
medium-security facility near Gig Harbor.
In an added layer of synergy, many of the plants raised at SCCC and
WCCW are planted on E. e. taylori restoration sites. These sites now host
butterflies raised by inmates at the Mission Creek Corrections Center for Women
(MCCCW), a minimum security prison near Belfair
Like the other SPP conservation projects, the butterfly program at
MCCCW is made possible by a diverse assemblage of collaborating partners. The
facility, a purpose-built 3 x 7.3 m partitioned greenhouse with UV-transmitting
glass panels (Plates 1 & 2), was built with a US Fish and Wildlife Service
(USFWS) grant, using funds from the Department of Defense’s Army Compatible
Use Buffer (ACUB) program. Captive rearing activities are supported by WDFW
and the Native Butterfly Conservation Lab at the Oregon Zoo. Funds for the
overall E. e. taylori recovery effort also come from ACUB and USFWS, and are
overseen by WDFW.
41
Plate 1 – Butterfly rearing facility at MCCCW, with flying shade cloth, winter
diapause shed, and raised garden beds for larval food plants
Plate 2 – Mary Jo Andersen of the Oregon Zoo working with inmates inside
the greenhouse at the beginning of the 2012 rearing season
42
Inmates at MCCCW helped build the butterfly greenhouse in 2011, and
then were trained to care for butterflies using painted ladies (Vanessa cardui) as a
training surrogate. These butterflies have short life cycles and are very forgiving
in terms of the conditions that they require for survival. Therefore, inmates were
able to use the E. e. taylori rearing and breeding protocols developed by the
Oregon Zoo, practicing through five complete life cycles before ever seeing a
Taylor’s checkerspot. When the E. e. taylori rearing season began, the inmates
quickly proved that the trust placed in their abilities by all of the funding and
conservation partners was well warranted. In 2012, 701 checkerspots were
released into the wild, 92 successful breeding introductions were made, resulting
in 3,624 pre-diapause larvae (180% of the pre-season target), and an egg-todiapause survivorship of 96.6%. The end of the rearing season is in early July
when the animals go into diapause, whereupon the bulk of them were taken to the
established diapause area at the Oregon Zoo. The other 500 were kept at MCCCW
over the winter as a trial. Of these, 100% survived. The entire cohort was returned
to MCCCW in late February for wake-up, after which 3400 were released,
bringing the total number of E. e. taylori individuals restored to the prairie so far
by MCCCW to over 4,000.
The success of the project can be attributed to at least three things: the
inmates, the facility, and the collaboration. The inmate butterfly technicians at
MCCCW have been meticulous, careful, thorough, and dedicated. They have
taken ownership of the project and its goals, they go out of their way to do things
better, and they keep records over and above what they have been asked to keep.
43
In some cases they have developed new methods to do things that have been
incorporated into protocols at the Oregon Zoo. Other times they have learned
things such as, if you rub the feet of a butterfly from the outside of its screen
enclosure it will bask its wings contentedly. At the end of the flight season in
2012, they took it on themselves to give hospital-like care to the older butterflies,
hand-feeding them honey water from tiny spoons. The point is, they have time on
their hands to do this work as thoroughly as can be imagined.
The second reason for the success of the project has been the facility. One
of the limiting variables at the Oregon Zoo facility is natural light. The E. e.
taylori rearing facility at the Oregon Zoo is housed in what was formerly a giant
air-conditioning unit for polar bears. They have two small windows to let in
natural light, which is important for butterfly life-stage cues and development.
Most of the time the butterflies have to make due with supplemental lighting on
timers. For breeding, staff have to hang them in small enclosures in front of the
windows, and then hope that clouds do not block the sun. At MCCCW, there is so
much light in the facility that breeding can be accomplished on the most
miserably drizzly overcast Washington day. The enclosures are hung near the
roof, the heat is cranked up, and the butterflies waste no time copulating as if it
were a hot sunny day on the prairie.
The third reason for the success of the MCCCW butterfly rearing facility
is the collaboration among numerous partners. The work would not be possible
without all the groups working together. The Department of Defense paid for the
facility, USFWS oversaw the funds, WDFW provides overarching project
44
leadership, training and rearing support comes from the Oregon Zoo, there is a
graduate student coordinator as well as faculty and staff support from SPP/TESC,
funding and staff support from WDOC, and the inmates themselves; all of these
partners play crucial roles in this truly synergistic conservation effort.
Value of collaborative conservation
Collaborative conservation has been identified as a rising phenomenon (Brick et
al. 2001, Lauber et al. 2011) which often provides significant benefits to multiple
partners. It can help involve community partners who might not otherwise
contribute, bridge the information divide between scientists and citizens, and
foster sharing of resources such as funding and labor. Additionally, multiple
partners can increase overall effectiveness by allowing partner groups to work
within their strengths. Furthermore, partners can reap the benefits of other groups’
strengths and the services they provide.
The butterfly program at MCCCW is an excellent example of
collaborative conservation. Every partner benefits and the net result is a more
effective effort than any group could accomplish alone. It has been described as a
5-way win-win situation: WDFW gets a second rearing facility where inmates do
professional work at a fraction of the cost, the Oregon Zoo enjoys greater
resilience in its captive colony and increased breeding capacity, WDOC receives
valuable programming for its inmates and receives positive media attention,
TESC is able to provide project management experience to a graduate student and
gains a new opportunity for research, and the DOD and JBLM get help restoring a
45
species that threatens to curtail training activities on an important practice range.
Other winners are the inmates themselves, who get the opportunity to contribute a
valuable service to society, are provided an environment where they can do
rehabilitative self-work while nurturing living beings, enjoy a collegial
relationship with academics and conservation professionals, and are exposed to
science and laboratory techniques for possible future study or employment.
Every partner wins. Every group is enjoying a success they could never
achieve alone. No matter how success is measured, the collaboration takes the
efforts of each partner and returns an emergent triumph. A prison raises
endangered butterflies, an army base gets to keep training with live artillery, a
state conservation agency doubles its output with negligible increase in cost, and a
college known for interdisciplinary learning puts a student right in the middle of
all of it; none of these benefits would be possible without the collaboration.
SOCIAL IMPLICATIONS
Transforming prisons
A retributive criminal justice paradigm has been historically prevalent in the
United States and around the world, and has largely continued to be so into the
present. Beginning in the 1990’s, however, increasing investments were made in
skill-building and re-entry programs, reflecting a shift toward a more restorative
criminal justice system (Phelps 2011).
The retributive and restorative justice paradigms are philosophically very
different in several important ways, as described in the criminal justice literature
46
(Wenzel et al. 2008, Dancig-Rosenberg and Gal 2013). Retributive justice
attempts to solve rule-breaking through punishment and the threat of punishment.
Interaction is one-directional, a power and status disparity is implied, and an
individualistic mentality is encouraged. Conversely, restorative justice attempts to
solve rule-breaking through rehabilitative programming and reinforcing the
offender’s role in society. Interaction is bi-directional, the value of every member
of society is implied, and a focus on community contribution is encouraged.
Retributive and restorative criminal justice paradigms both attempt to
solve the same societal problem, but they do so in nearly opposite ways, leading
some to claim that they may work against each other (Bazemore 1998). This idea
holds that rehabilitative programming may soften the effects of punishment, and
punishment may lessen the effects of rehabilitation. This concept of polarity is
one of the standard arguments for those who consider punishment-only
incarceration the best model. Since the retributive justice paradigm continues to
be the undeniable foundation of incarceration, this sentiment is not uncommon
within the prison system itself.
Organizations like SPP, however, are proud to champion the growth of
restorative justice, and believe that rehabilitative programming does the opposite
of limiting the effectiveness of punishment. It works with rather than against
incarceration and adds to the functionality of the justice system as a whole, while
also tending to increase prison safety at the same time (LeRoy et al. 2012). This
way of thinking is borne out by more recent criminal justice literature. An
upcoming article in the Carrozo Law Review (Dancig-Rosenberg and Gal 2013)
47
redefines the role of retributive justice by placing punishment as a useable tool in
a larger restorative framework.
Under the restorative justice paradigm, the long-term goal of corrections is
founded in the assumption that inmates serve their time and then return to our
communities to become our neighbors again. Reports have shown that purely
punitive penal systems do not do an efficient job rehabilitating people, and are in
fact often more likely to “lead to more crime following release” (Chen and
Shapiro 2007). Harsher sentences have been correlated with poorer post-release
employment rates (Western et al. 2001), while prison stays have been shown to
increase introversion and violent tendencies (Bolton et al. 1976), and peer-to-peer
social interaction in prison has been linked to an evolution of criminal tendencies,
or a crime learning effect (Glaeser et al. 1996, Bayer et al. 2004). If the goal of
corrections is to get people back on track so that they can become functional
members of our society, then a different approach is needed.
Rehabilitative programming can take many forms. If the measurement of
rehabilitative success is getting people back on their feet and functioning in
society, then the most direct methods are programming elements that have been
shown empirically to reduce recidivism such as formal education and re-entry job
skills training. Other types of programming which also can be considered
rehabilitative include informal education, opportunities for offenders to contribute
to the outside community, and activities that decrease negative emotions during
incarceration (such as gardening or access to a library).
48
If there is a weakness in the effectiveness of the SPP’s conservation
programs in terms of rehabilitation, it is that emphasis on developing marketable
skills for the post-release job market is not the primary focus. That said, every
other category of rehabilitative programming is covered. At MCCCW, a wide
range of environmental and scientific topics are discussed by inmate butterfly
technicians and the TESC graduate student overseeing the project, during extra
time budgeted every week for that purpose. In addition, several lectures have been
brought to MCCCW for the general population to increase education about
butterflies.
Opportunities to contribute meaningfully to the larger community are also
considered important for rehabilitation. The butterfly technicians at MCCCW are
able to play an integral part in efforts to restore a federally threatened pollinator to
south Puget lowland prairies. Furthermore, they are helping conduct relevant
research and contributing to the body of scientific literature about the species they
work with. In fact, in the case of the oviposition preference study, two of the
technicians were so involved and took such ownership that they earned spots as
co-authors on the scientific manuscript being prepared for peer-reviewed
publication. Also, before the study there had been no documentation of E. e.
taylori using C. levisecta as an oviposition host at all, so after it was clear that this
was indeed happening a camera and macro-lens were brought to the greenhouse.
One of the inmates was an avid amateur photographer before her incarceration,
and she got to be the first person to document the relationship (Plate 3).
49
Plate 3 – A Taylor’s checkerspot female ovipositing on golden paintbrush; this
photo was taken by an inmate butterfly technician at MCCCW and is the first
documentation of this interaction between the two threatened species.
Another opportunity to contribute is that, in a larger sense, the inmates’
success at MCCCW is paving the way for this model to spread. By showing that
butterflies and prisons are a good match, and doing so with high success rates, the
inmate butterfly technicians are making their program an example to the world.
When other states or nations go to sell the idea of endangered butterflies in prison
to policy makers and land managers, the MCCCW butterfly program provides
evidence that the model works. By working hard, taking ownership, and caring
for their charges with delicate patience, they are potentially paving the way for
other women to have similar opportunities, in other states, or possibly even
around the world. Since 2012, another butterfly program has been started at the
50
Washington State Penitentiary, and plans are being made for a butterfly rearing
program in Oregon.
There are several types of prison programming that are considered
rehabilitative, but one of the common themes is the reduction of negative
emotions. Typical examples of programming thought to be effective at this are
jobs, recreation, gardening, access to books or art supplies, and religious services.
I argue that the butterfly program at MCCCW has the potential to reduce negative
emotions in several ways. First, working with and nurturing living organisms,
then watching them develop and metamorphose into butterflies may be
therapeutic, and provides an example of transformation and change for people
undergoing changes within themselves. Second, the greenhouse itself is a peaceful
environment, outside the fenced yard and all its rush and intensity. The butterfly
technicians at MCCCW have reported listening to the caterpillars chew leaves on
a quiet sunny afternoon, and have often commented on the meditative quality of
the working environment. Next, the SPP is committed to interacting with inmate
technicians as colleagues rather than employees. Their ideas are welcomed and
often implemented by the facility at the Oregon Zoo. They have the opportunity to
sit with a graduate student every week, and discussion is welcomed on any
scientific topic they are interested in. Furthermore, the inmates are able to interact
with college professors and agency biologists within the framework of a
professional relationship. They are even allowed to attend (with custody staff
escort) annual working group meetings for the range-wide E. e. taylori recovery
effort, where they are able to hear presentations and discussions on all the latest
51
ideas, from genetics research to site suitability reports and wild population
updates. I argue that the validation and respect inherent in being treated like a
partner can also serve to reduce negative emotions and perhaps be rehabilitative.
Community benefits
The reality of reentry is that inmates return home. As discussed above, they finish
their sentences and return to our communities, rejoining family and becoming our
neighbors. The aspects of prison aimed at rehabilitation are the ones that serve a
functional purpose for the future good of society.
The SPP is helping prisons by providing a new avenue for functionality in
an increasingly connected and aware society. The benefits of rehabilitative
programming were not newly invented by SPP, however. The elegance of the SPP
idea is that it brings another social goal, ecological restoration, into the picture. In
fact, SPP brings several societal needs to the table at the same time and uses
collaboration to address them all. If all aspects of this multi-dimensional
community benefit were realized, prisons would be more functional in the
community, the military would be able to keep us all safer, conservation efforts
would be more effective, students would become better graduates for the working
world, and scientific knowledge would increase.
52
Involving underserved audiences
There is growing emphasis in the scientific community on reaching out from the
ivory tower to involve a wider audience in science education (Nadkarni 2004,
2006, 2007, McCallie et al. 2009, Bonney et al. 2009). Benefits of outreach and
informal science education may include broader discussions, new ideas, greater
community participation, more new student enrollments, and increased interest
through media exposure.
Involving inmates in relevant scientific research brings a novel audience
into the discussion. Educational programming in prisons has traditionally been
almost entirely through high school equivalency, associate’s degree programs,
and religious learning, but involving inmates in research brings science into the
prison community in a new way (Weber 2012). Inmates who participate learn
through experience, and may extend the effects of science education by talking to
fellow inmates about their jobs.
Other underserved audiences may also benefit from involvement in
relevant scientific research, such as people in restrictive institutions like jails,
mental institutions, retirement homes, and public schools. The work of forming
these partnerships is beyond the scope of SPP, but the model that SPP has created
may inspire other groups to seek out such novel collaborations in the future.
AN INTERDISCIPLINARY THESIS
In working for the SPP, coordinating the butterfly program at MCCCW and
carrying out this oviposition preference research, many lines of interdisciplinarity
53
were explored, joining conservation biology with environmental education and
social justice. I worked within the criminal justice system and helped provide
corrections programming while bringing collaborative conservation to an
underserved scientific audience. I learned about restoration ecology on butterfly
release sites, and endangered species protection working with state agencies. I
learned how to staff and manage a captive breeding facility, and played a role in a
multi-partner collaboration. I participated in informal science education and
public outreach, speaking at conferences, community events, and a public
elementary school. I would argue that my work with incarcerated women and
endangered butterflies, and the oviposition preference research that we did
together, reflects a truly interdisciplinary MES thesis.
CONCLUSIONS
In my opinion, the most important implication of the oviposition preference study
and its results was that there may now be the opportunity to bring two threatened
species together for the mutual benefit of both. If further research continues to
indicate that C. levisecta would be a suitable oviposition host for E. e. taylori, it
could be planted at the butterfly reintroduction sites. This unified restoration
approach could provide a more diverse assemblage of resources for the animals
while reducing the need to plant exotic P. lanceolata on high quality native
prairie, while at the same time adding planting sites to the C. levisecta recovery
effort.
54
The most important aspect of doing this research at MCCCW has been in
establishing a successful model for conservation work in prisons, which is being
expanded to other situations. By showing that a program like the one at MCCCW
can come on-line quickly, and function both effectively and inexpensively, we are
paving the way for other similar programs to follow. Our success makes the idea
of prisons as conservation partners attractive and easier to sell to policy makers in
other states. In the future, I would like to see more states emulating this model,
with a variety of other species and other partners.
I think that an ideal system moving forward would be to have zoos and
other professional rearing institutions shift from long-term rearing to protocol
development. They could spend several years developing a successful protocol for
a particular species, then move the operation into a prison where labor is cheaper
and serves the dual function as rehabilitative inmate programming. The zoo could
then shift its focus to developing a protocol for a new species. This system would
help prison-based facilities be successful, while also solving a funding dilemma
for zoos and conservation agencies. One of the problems with long-term rearing
operations is that they become increasingly difficult to fund as years go by. Space
and money are always at a premium in zoos and it is often easier to fund exciting
new projects than continue projects that have been around for a decade or more.
A partnership between zoos and prisons is a natural fit for the rearing of
butterflies, and if I could choose a trajectory for my career as an MES graduate, I
would hope to facilitate that partnership in many different states. Each new
partnership would provide a springboard for new programs in more prisons,
55
helping restore more ecosystems while at the same time providing rehabilitative
programming opportunities to more inmates. If I could, I would make helping the
synergistic metamorphosis of conservation and incarceration my life’s work.
56
References
An, Y., S. Wan, X. Zhou, A. A. Subedar, L. L. Wallace, and Y. Luo. 2005. Plant
nitrogen concentration, use efficiency, and contents in a tallgrass prairie
ecosystem under experimental warming. Global Change Biology 11:1733–
1744.
Bachelet, D., B. R. Johnson, S. D. Bridgham, P. V. Dunn, H. E. Anderson, and B.
M. Rogers. 2011. Climate change impacts on western Pacific Northwest
prairies and savannas. Northwest Science 85:411–429.
Bachelet, D., R. P. Neilson, J. M. Lenihan, and R. J. Drapek. 2001. Climate
change effects on vegetation distribution and carbon budget in the United
States. Ecosystems 4:164–185.
Barclay, E., M. J. Anderson, H. Wendt, and M. Arnold. 2009. Husbandry manual,
Taylor’s checkerspot Euphydryas editha taylori. Report to the Oregon
Zoo, Portland, Oregon.
Bayer, P., R. Pintoff, and D. E. Pozen. 2004. Building criminal capital behind
bars: Social learning in juvenile corrections. Discussion paper No. 864,
Economic Growth Center, Yale University.
Bazemore, G. 1998. Restorative justice and earned redemption communities,
victims, and offender reintegration. American Behavioral Scientist
41:768–813.
Black, S. H., and D. M. Vaughan. 2005. Species profile: Euphydryas editha
taylori. in M. D. Shepherd and D. M. Vaughan, editors. Red List of
57
Pollinator Insects of North America. The Xerces Society for Invertebrate
Conservation, Portland, OR.
Bolton, N., F. V. Smith, K. J. Heskin, and P. A. Banister. 1976. Psychological
correlates of long-term imprisonment. British Journal of Criminology
16:38.
Bonney, R., H. Ballard, R. Jordan, E. McCallie, T. Phillips, J. Shirk, and C. C.
Wilderman. 2009. Public participation in scientific research: Defining the
field and assessing its potential for informal science education. CAISE
Inquiry Group Report, Center for Advancement of Informal Science
Education, Washington, D.C.
Bowers, M. D. 1981. Unpalatability as a defense strategy of western checkerspot
butterflies (Euphydryas scudder, Nymphalidae). Evolution 35:367–375.
Brick, P., D. Snow, and S. V. D. Wetering. 2001. Across the great divide:
Explorations in collaborative conservation and the American West. Island
Press, Washington D. C.
Chappell, C., and R. Crawford. 1997. Native vegetation of the south Puget Sound
landscape. in P. Dunn and K. Ewing, editors. Ecology and Conservation of
the South Puget Sound Prairie Landscape. The Nature Conservancy,
Seattle, WA.
Char, P., and P. D. Boersma. 1995. The effects of prairie fragmentation on
butterfly species in western Washington. Report to The Nature
Conservancy and the U. S. Army, Fort Lewis, University of Washington,
Seattle, Washington.
58
Chen, M. K., and J. M. Shapiro. 2007. Do harsher prison conditions reduce
recidivism? A discontinuity-based approach. American Law and
Economics Review 9:1–29.
Clampitt, C. A. 1993. Effects of human disturbances on prairies and the regional
endemic Aster curtis in western Washington. Northwest Science 67:163–
169.
Coops, N. C., and R. H. Waring. 2011. Estimating the vulnerability of fifteen tree
species under changing climate in Northwest North America. Ecological
Modeling 222:2119–2129.
Crawford, R. C., C. Chappell, B. Stephens, C. Soper, and D. Rolph. 1995.
Inventory and mapping of endangered native ecosystems of Ft. Lewis.
Report to the Environmental and Natural Resources Division, Fort Lewis,
Washington. Washington Department of Natural Resources, Natural
Hertiage Program and The Nature Conservancy, Olympia, Washington.
Crawford, R., and H. Hall. 1997. Changes in the south Puget prairie landscape. in
P. V. Dunn and K. Ewing, editors. Ecology and Conservation of the South
Puget Sound Prairie Landscape. The Nature Conservancy of Washington,
Seattle, Washington.
Crooks, J. A., and M. E. Soule. 2001. Lag times in population explosions of
invasive species: Casues and implications. in O. T. Sandlund, P. J. Schei,
and Å. Viken, editors. Invasive Species and Biodiversity Management.
Kluwer Academic Publishers, Dordrecht, The Netherlands.
59
Dancig-Rosenberg, H., and T. Gal. 2013. Restorative criminal justice. Cardozo
Law Review, forthcoming.
Dennehy, C., E. R. Alverson, H. E. Anderson, D. R. Clements, R. Gilbert, and T.
N. Kaye. 2011. Management strategies for invasive plants in Pacific
Northwest prairies, savannas, and oak woodlands. Northwest Science
85:329–351.
Dornfeld, E. J. 1980. Butterflies of Oregon. Timber Press, Forest Grove, OR.
Dunn, R. R., N. C. Harris, R. K. Colwell, L. P. Koh, and N. S. Sodhi. 2009. The
sixth mass coextinction: Are most endangered species parasites and
mutualists? Proceedings of the Royal Society B, Biological Sciences
276:3037–3045.
Dunwiddie, P. W., and J. D. Bakker. 2011. The future of restoration and
management of prairie-oak ecosystems in the Pacific Northwest.
Northwest Science 85:83–92.
Ehrlich, P. R. 1961. Intrinsic barriers to dispersal in checkerspot butterfly. Science
134:108–109.
Ehrlich, P. R. 1965. The population biology of the butterfly, Euphydryas editha.
II. The Structure of the Jasper Ridge Colony. Evolution 19:327–336.
Ehrlich, P. R., and I. Hanski. 2004. On the wings of checkerspots: A model
system for population biology. Oxford University Press, New York, NY.
Ehrlich, P. R., D. D. Murphy, M. C. Singer, C. B. Sherwood, R. R. White, and I.
L. Brown. 1980. Extinction, reduction, stability and increase: The
60
responses of checkerspot butterfly (Euphydryas) populations to the
California drought. Oecologia 46:101–105.
Evans, S., R. Schuller, and E. Augenstein. 1984. A report on Castilleja levisecta
Greenm. at Rocky Prairie, Thurston County, Washington. Report to The
Nature Conservancy, Washington Field Office, Seattle, Washington.
Fimbel, C. 2004. Habitat enhancement for rare butterflies on Fort Lewis prairies.
Report to The Nature Conservancy, South Sound Office, Olympia
Washington.
Floberg, J., M. Goering, G. Wilhere, C. MacDonald, C. Chappell, C. Rumsey, Z.
Ferdana, A. Holt, P. Skidmore, T. Horsman, E. Alverson, C. Tanner, M.
Bryer, P. Lachetti, A. Harcombe, B. McDonald, T. Cook, M. Summers,
and D. Rolph. 2004. Willamette Valley-PugetTrough-Georgia Basin
ecoregional assessment. The Nature Conservancy with support from the
Nature Conservancy of Canada, Washington Department of Fish and
Wildlife, Washington Department of Natural Resources (Natural Heritage
and Nearshore Habitat programs), Oregon State Natural Heritage
Information Center and the British Columbia Conservation Data Centre.
Franklin, J. F., and C. T. Dyrness. 1973. Natural Vegetation of Oregon and
Washington. Oregon State University Press, Corvallis, OR.
Gammon, J. 1995. Report on the status of Castilleja levisecta. Report to the
Washington Natural Heritage Program, Department of Natural Resources,
Olympia, Washington
61
Giles, L. J. 1970. The ecology of the mounds on Mima Prairie with special
reference to Douglas-fir invasion. M.S. thesis, University of Washington.
Glaeser, E. L., B. Sacerdote, and J. A. Scheinkman. 1996. Crime and social
interactions. The Quarterly Journal of Economics 111:507–548.
Grosboll, D. N. 2004. Captive rearing the endangered mardon skipper (Polites
mardon) and Taylor’s checkerspot (Euphydryas editha taylori) butterflies:
Initial results (Lepidoptera, Nymphalidae). Proceedings of the Species at
Risk Pathways to Recovery Conference, Victoria, B. C.
Grosboll, D. N. 2011. Taylor’s checkerspot (Euphydryas editha taylori)
oviposition habitat selection and larval hostplant use in Washington State.
MES thesis. The Evergreen State College.
Hamann, A., and T. Wang. 2006. Potential effects of climate change on
ecosystem and tree species distribution in British Columbia. Ecology
87:2773–2786.
Hamman, S. T., P. W. Dunwiddie, J. L. Nuckols, and M. McKinley. 2011. Fire as
a restoration tool in Pacific Northwest prairies and oak woodlands:
Challenges, successes, and future directions. Northwest Science 85:317–
328.
Harrison, S., D. D. Murphy, and P. R. Ehrlich. 1988. Distribution of the bay
checkerspot butterfly, Euphydryas editha bayensis: Evidence for a
metapopulation model. The American Naturalist 132:360–382.
Jackson, L. E. 1982. Comparison of phenological patterns in prairie and subalpine
meadow communities. Northwest Science 56:316–328.
62
Karl, T. R., and R. W. Knight. 1998. Secular trends of precipitation amount,
frequency, and intensity in the United States. Bulletin of the American
Meteorological Society 79:231–241.
Kearns, C. A., D. W. Inouye, and N. M. Waser. 1998. Endangered mutualisms:
The conservation of plant-pollinator interactions. Annual Review of
Ecology and Systematics 29:83–112.
Klein, J. A., J. Harte, and X.-Q. Zhao. 2004. Experimental warming causes large
and rapid species loss, dampened by simulated grazing, on the Tibetan
Plateau. Ecology Letters 7:1170–1179.
Koh, L. P., R. R. Dunn, N. S. Sodhi, R. K. Colwell, H. C. Proctor, and V. S.
Smith. 2004. Species coextinctions and the biodiversity crisis. Science
305:1632–1634.
Lang, F. A. 1961. A study of vegetation change on the gravelly prairies of Pierce
and Thurston Counties, Western Washington. M.S. thesis, University of
Washington.
Lauber, T. B., R. C. Stedman, D. J. Decker, and B. A. Knuth. 2011. Linking
knowledge to action in collaborative conservation. Conservation Biology
25:1186–1194.
Lawrence, B. A., and T. N. Kaye. 2006. Habitat variation throughout the historic
range of golden paintbrush, a Pacific Northwest endemic: Implications for
restoration. Northwest Science 80:140–152.
63
Lawrence, B. A., and T. N. Kaye. 2008. Direct and indirect effects of host plants:
Implications for reintroduction of an endangered hemiparasitic plant
(Castilleja levisecta). Madroño 55:151–158.
Lawrence, B. A., and T. N. Kaye. 2011. Reintroduction of Castilleja levisecta:
Effects of ecological similarity, source population genetics, and habitat
quality. Restoration Ecology 19:166–176.
Leopold, E. B., and R. Boyd. 1999. An ecological history of old prairie areas in
southwestern Washington. in R. Boyd, editor. Indians, Fire and Land in
the Pacific Northwest. Oregon State University Press, Corvalis, OR.
LeRoy, C. J., K. Bush, J. Trivett, and B. Gallagher. 2012. The Sustainability in
Prisons Project: An overview (2004-12). Gorham Printing, Olympia, WA.
Linders, M. J. 2007. Development of captive rearing and translocation methods
for Taylor’s checkerspot (Euphydryas editha taylori) in south Puget
Sound, Washington: 2006-2007 Annual Report. Report to the Washington
Department of Fish and Wildlife, Olympia, Washington.
Linders, M. J. 2012. Captive rearing and translocation of Taylor’s checkerspot in
south Puget Sound: 2011-2012 ACUB Annual Progress Report. Report to
the Washington Department of Fish and Wildlife, Olympia, Washington.
Littell, J. S., E. E. Oneil, D. McKenzie, J. A. Hicke, J. A. Lutz, R. A. Norheim,
and M. M. Elsner. 2010. Forest ecosystems, disturbance, and climatic
change in Washington State, USA. Climatic Change 102:129–158.
MacArthur, R. H. 1967. The Theory of Island Biogeography. Princeton
University Press, Princeton, NJ.
64
Mackay, D. A. 1985. Conspecific host discrimination by ovipositing Euphydryas
editha butterflies: Its nature and its consequences for offspring
survivorship. Researches on Population Ecology 27:87–98.
McCallie, E., L. Bell, T. Lohwater, J. H. Falk, J. L. Lehr, B. V. Lewenstein, C.
Heedham, and B. Wiehe. 2009. Many experts, many audiences: Public
engagement with science and informal science education. A CAISE
Inquiry Group Report, Center for Advancement of Informal Science
Education, Washington, D.C.
McLaughlin, J. F., J. J. Hellmann, C. L. Boggs, and P. R. Ehrlich. 2002. Climate
change hastens population extinctions. Proceedings of the National
Academy of Sciences 99:6070–6074.
del Moral, R., and D. C. Deardorff. 1976. Vegetation of the Mima Mounds,
Washington State. Ecology 57:520–530.
Morris, W. G. 1934. Forest fires in western Washington and western Oregon.
Oregon Historical Quarterly 35:313–339.
Mote, P. W., and E. P. Salathé. 2010. Future climate in the Pacific Northwest.
Climatic Change 102:29–50.
Nadkarni, N. M. 2004. Not preaching to the choir: Communicating the
importance of forest conservation to nontraditional audiences.
Conservation Biology 18:602–606.
Nadkarni, N. M. 2006. The moss-in-prison project: Disseminating science beyond
academia. Frontiers in Ecology and the Environment 4:442–443.
65
Nadkarni, N. M. 2007. Ecological outreach to faith- based communities. Frontiers
in Ecology and the Environment 5:332.
Nisbet, J. 2009. The Collector: David Douglas and the Natural History of the
Northwest. Sasquatch Books, Seattle, WA.
Norton, H. H. 1979. The association between anthropogenic prairies and
important food plants in western Washington. Northwest Anthropological
Research Notes 13:175–200.
Noss, R. F., E. T. LaRoe III, and J. M. Scott. 1995. Endangered ecosystems of the
United States: A preliminary assessment of loss and degradation. Report
to the U. S. Department of the Interior, National Biological Service,
Washington D. C.
Phelps, M. S. 2011. Rehabilitation in the punitive era: The gap between rhetoric
and reality in U.S. prison programs. Law & Society Review 45:33–68.
Pyle, R. M. 2002. The Butterflies of Cascadia: A Field Guide to All the Species of
Washington, Oregon, and Surrounding Territories. Seattle Audubon
Society, Seattle, WA.
Pyle, R. M., and S. A. Society. 1974. Watching Washington Butterflies: An
Interpretive Guide to the State’s 134 species, Including Most of the
Butterflies of Oregon, Idaho, and British Columbia. Seattle Audubon
Society, Seattle, WA
Quammen, D. 2012. The Song Of The Dodo: Island Biogeography in an Age of
Extinctions. Scribner Island Press, New York, NY
66
Rehfeldt, G. E., N. L. Crookston, M. V. Warwell, and J. S. Evans. 2006.
Empirical analyses of plant‐climate relationships for the western United
States. International Journal of Plant Sciences 167:1123–1150.
Rezende, E. L., J. E. Lavabre, P. R. Guimarães, P. Jordano, and J. Bascompte.
2007. Non-random coextinctions in phylogenetically structured
mutualistic networks. Nature 448:925–928.
Ross, D. 2003. First year (2003) cencus of the Carwell Hill population of Taylor’s
checkerspot (Euphydryas editha taylori) in Benton County, Oregon.
Report to the Xerces Society, Corvallis, Oregon.
Rustad, L., J. Campbell, G. Marion, R. Norby, M. Mitchell, A. Hartley, J.
Cornelissen, J. Gurevitch, and GCTE-NEWS. 2001. A meta-analysis of
the response of soil respiration, net nitrogen mineralization, and
aboveground plant growth to experimental ecosystem warming. Oecologia
126:543–562.
Sakai, A. K., F. W. Allendorf, J. S. Holt, D. M. Lodge, J. Molofsky, K. A., S.
Baughman, R. J. Cabin, J. E. Cohen, N. C. Ellstrand, D. E. Mccauley, P.
O’neil, I. M. Parker, J. N. Thompson, and S. G. Weller. 2001. The
population biology of invasive species. Annual Review of Ecology and
Systematics 32:305–332.
Schultz, C. B., E. Henry, A. Carleton, T. Hicks, R. Thomas, A. Potter, M. Collins,
M. Linders, C. Fimbel, S. Black, H. E. Anderson, G. Diehl, S. Hamman,
R. Gilbert, J. Foster, D. Hays, D. Wilderman, R. Davenport, E. Steel, N.
Page, P. L. Lilley, J. Heron, N. Kroeker, C. Webb, and B. Reader. 2011.
67
Conservation of prairie-oak butterflies in Oregon, Washington, and British
Columbia. Northwest Science 85:361–388.
Severns, P. M., and D. Grosboll. 2011. Patterns of reproduction in four
Washington State populations of Taylor’s checkerspot (Euphydryas editha
taylori) during the spring of 2010. Report to The Nature Conservancy,
Olympia, Washington.
Severns, P. M., and A. D. Warren. 2008. Selectively eliminating and conserving
exotic plants to save an endangered butterfly from local extinction.
Animal Conservation 11:476–483.
Shafer, S. L., P. J. Bartlein, and R. S. Thompson. 2001. Potential changes in the
distributions of western North America tree and shrub taxa under future
climate scenarios. Ecosystems 4:200–215.
Shaver, G. R., J. Canadell, F. S. Chapin, J. Gurevitch, J. Harte, G. Henry, P.
Ineson, S. Jonasson, J. Melilo, L. Pitelka, and L. Rustad. 2000. Global
warming and terrestrial ecosystems: A conceptual framework for analysis.
BioScience 50:871–882.
Sheehan, M., and N. Sprague. 1984. Report on the status of Castilleja levisecta
Greenm. Report to The Nature Conservancy, Seattle, Washington.
Shepard, J., and C. Guppy. 2011. Butterflies of British Columbia: Including
Western Alberta, Southern Yukon, the Alaska Panhandle, Washington,
Northern Oregon, Northern Idaho, and Northwestern Montana. UBC
Press, Vancouver B. C.
68
Shepard, J. H. 2000. Status report on Taylor’s checkerspot Euphydryas editha
taylori (Edwards) (Lepidoptera: Nymphalidae) in British Columbia.
Report to the Committee on the Status of Endangered Wildlife in Canada,
Ottawa, Ontario.
Singer, M. C. 1972. Complex components of habitat suitability within a butterfly
colony. Science 176:75–77.
Singer, M. C. 1982. Quantification of host preference by manipulation of
oviposition behavior in the butterfly Euphydryas editha. Oecologia
52:224–229.
Singer, M. C., and P. R. Ehrlich. 1979. Population dynamics of the checkerspot
butterfly Euphydryas editha. Fortschritte der Zoologie 25:53–60.
Singer, M. C., and J. R. Lee. 2000. Discrimination within and between host
species by a butterfly: Implications for design of preference experiments.
Ecology Letters 3:101–105.
Singer, M. C., D. Ng, and R. A. Moore. 1991. Genetic variation in oviposition
preference between butterfly populations. Journal of Insect Behavior
4:531–535.
Singer, M. C., D. Ng, and C. D. Thomas. 1988. Heritability of oviposition
preference and its relationship to offspring performance within a single
insect population. Evolution 42:977-985.
Singer, M. C., D. Vasco, C. Parmesan, C. D. Thomas, and D. Ng. 1992.
Distinguishing between “preference” and “motivation” in food choice: An
example from insect oviposition. Animal Behaviour 44, Part 3:463–471.
69
Solomon, S., D. Qin, M. Manning, Z. Chen, M. Marquis, K. B. Averyt, M.
Tignor, and H. L. Miller. 2007. Contribution of working group I to the
fourth assessment report of the Intergovernmental Panel on Climate
Change, 2007. Cambridge University Press, New York, NY.
Stanley, A. G., T. N. Kaye, and P. W. Dunwiddie. 2008. Regional strategies for
restoring invaded prairies. Native Plants Journal 9:247 –254.
Stinson, D. W. 2005. Draft status report for the Mazama pocket gopher, streaked
horned lark, and Taylor’s checkerspot. Report to the Washington
Department of Fish and Wildlife, Olympia, Washington.
Storm, L., and D. Shebitz. 2006. Evaluating the purpose, extent, and ecological
restoration applications of indigenous burning practices in southwestern
Washington. Ecological Restoration 24:256–268.
Suttle, K. B., M. A. Thomsen, and M. E. Power. 2007. Species interactions
reverse grassland responses to changing climate. Science 315:640–642.
Tveten, R. 1997. Fire effects on prairie vegetation, Fort Lewis, Washington. in P.
V. Dunn and K. Ewing, editors. The Ecology and Conservation of the
South Puget Sound Landscape. The Nature Conservancy of Washington,
Seattle, WA.
de Valpine, P., and J. Harte. 2001. Plant responses to experimental warming in a
montane meadow. Ecology 82:637–648.
Walker, M. D., C. H. Wahren, R. D. Hollister, G. H. R. Henry, L. E. Ahlquist, J.
M. Alatalo, M. S. Bret-Harte, M. P. Calef, T. V. Callaghan, A. B. Carroll,
H. E. Epstein, I. S. Jónsdóttir, J. A. Klein, B. Magnússon, U. Molau, S. F.
70
Oberbauer, S. P. Rewa, C. H. Robinson, G. R. Shaver, K. N. Suding, C. C.
Thompson, A. Tolvanen, Ø. Totland, P. L. Turner, C. E. Tweedie, P. J.
Webber, and P. A. Wookey. 2006. Plant community responses to
experimental warming across the tundra biome. Proceedings of the
National Academy of Sciences 103:1342–1346.
Weber, S. R., 2012. Environmental education in prison: A comparison of teaching
methods and their influence in inmate attitudes and knowledge of
environmental topics. MES thesis. The Evergreen State College.
Weiss, S. B. 1999. Cars, cows, and checkerspot butterflies: Nitrogen deposition
and management of nutrient-poor grasslands for a threatened species.
Conservation Biology 13:1476–1486.
Weiss, S. B., D. D. Murphy, and R. R. White. 1988. Sun, slope, and butterflies:
Topographic determinants of habitat quality for Euphydryas Editha.
Ecology 69:1486–1496.
Wentworth, J. B. 2001. The demography and population dynamics of Castilleja
levisecta, a federally threatened perennial of Puget Sound grasslands. in R.
S. Reichard, P. Dunwiddie, J. Gamon, A. Kruckeberg, and D. Salstrom,
editors. Conservation of Washington’s Native Plants and Ecosystems.
Washington Native Plant Society, Seattle, Washington
Wenzel, M., T. G. Okimoto, N. T. Feather, and M. J. Platow. 2008. Retributive
and restorative justice. Law and Human Behavior 32:375–389.
Western, B., J. R. Kling, and D. F. Weiman. 2001. The labor market
consequences of incarceration. Crime & Delinquency 47:410–427.
71
Wold, E. N., J. E. Jancaitis, T. H. Taylor, and D. M. Steeck. 2011. Restoration of
agricultural fields to diverse wet prairie plant communities in the
Willamette Valley, Oregon. Northwest Science 85:269–287.
Zavaleta, E. S., M. R. Shaw, N. R. Chiariello, H. A. Mooney, and C. B. Field.
2003. Additive effects of simulated climate changes, elevated CO2, and
nitrogen deposition on grassland diversity. Proceedings of the National
Academy of Sciences 100:7650–765
72