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Western Monarch Population Plummets: Status, Probable Causes, and Recommended Conservation Actions

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Frontiers in Ecology and Evolution
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  • The Xerces Society
  • Xerces Society for Invertebrate Conservation

Abstract and Figures

Western monarch butterflies dropped by ~97% of their average historic abundance between the 1980s and mid-2010s. In winter 2018–2019, the population plummeted even farther, to fewer than 30,000 monarchs, which represents a single year drop of 86% and a drop of >99% since the 1980s. The population may now be hovering at its quasi-extinction threshold. In this Perspectives piece, we: (1) Place the current status in context, (2) Highlight the most likely window during the annual life cycle when the population declined, (3) Review probable causes of long-term declines, and (4) Recommend steps that the public, policy makers, and land managers can take to recover western monarchs. The available studies reinforce the hypotheses that overwintering habitat loss and loss of central California breeding habitat, as well as pesticide use, are likely important contributors to the western monarch's long-term decline. The most limiting part of the migratory cycle appears to be concentrated during the overwintering stage and/or in early spring. If western monarchs are in fact entering an extinction vortex, they need extraordinary efforts—focused on the most vulnerable periods of the annual cycle— to save the migration. Critical short-term conservation priorities are to (1) Protect, manage and restore overwintering habitat, (2) Protect monarchs and their habitat from pesticides, (3) Restore breeding and migratory habitat in California, (4) Protect, manage, and restore summer breeding and fall migration monarch habitat throughout the western monarch's range, and (5) Fill research gaps to inform western monarch recovery strategies.
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PERSPECTIVE
published: 03 July 2019
doi: 10.3389/fevo.2019.00258
Frontiers in Ecology and Evolution | www.frontiersin.org 1July 2019 | Volume 7 | Article 258
Edited by:
Jay E. Diffendorfer,
United States Geological Survey,
United States
Reviewed by:
Tyler Flockhart,
University of Maryland Center for
Environmental Science (UMCES),
United States
Ralph Grundel,
U.S. Geological Survey, Great Lakes
Science Center, Chesterton, IN,
United States
*Correspondence:
Elizabeth E. Crone
elizabeth.crone@tufts.edu
Specialty section:
This article was submitted to
Conservation,
a section of the journal
Frontiers in Ecology and Evolution
Received: 05 March 2019
Accepted: 18 June 2019
Published: 03 July 2019
Citation:
Pelton EM, Schultz CB, Jepsen SJ,
Black SH and Crone EE (2019)
Western Monarch Population
Plummets: Status, Probable Causes,
and Recommended Conservation
Actions. Front. Ecol. Evol. 7:258.
doi: 10.3389/fevo.2019.00258
Western Monarch Population
Plummets: Status, Probable Causes,
and Recommended Conservation
Actions
Emma M. Pelton 1, Cheryl B. Schultz 2, Sarina J. Jepsen 1, Scott Hoffman Black 1and
Elizabeth E. Crone 3
*
1The Xerces Society for Invertebrate Conservation, Portland, OR, United States, 2Department of Biological Sciences,
Washington State University, Vancouver, WA, United States, 3Department of Biology, Tufts University, Medford,
MA, United States
Western monarch butterflies dropped by 97% of their average historic abundance
between the 1980s and mid-2010s. In winter 2018–2019, the population plummeted
even farther, to fewer than 30,000 monarchs, which represents a single year drop of
86% and a drop of >99% since the 1980s. The population may now be hovering
at its quasi-extinction threshold. In this Perspectives piece, we: (1) Place the current
status in context, (2) Highlight the most likely window during the annual life cycle when
the population declined, (3) Review probable causes of long-term declines, and (4)
Recommend steps that the public, policy makers, and land managers can take to recover
western monarchs. The available studies reinforce the hypotheses that overwintering
habitat loss and loss of central California breeding habitat, as well as pesticide use,
are likely important contributors to the western monarch’s long-term decline. The most
limiting part of the migratory cycle appears to be concentrated during the overwintering
stage and/or in early spring. If western monarchs are in fact entering an extinction
vortex, they need extraordinary efforts—focused on the most vulnerable periods of the
annual cycle— to save the migration. Critical short-term conservation priorities are to (1)
Protect, manage and restore overwintering habitat, (2) Protect monarchs and their habitat
from pesticides, (3) Restore breeding and migratory habitat in California, (4) Protect,
manage, and restore summer breeding and fall migration monarch habitat throughout
the western monarch’s range, and (5) Fill research gaps to inform western monarch
recovery strategies.
Keywords: Danaus plexippus plexippus, western monarchs, quasi-extinction, conservation, population trends
INTRODUCTION
Monarch butterflies (Danaus plexippus plexippus) across North America have been undergoing
a multi-decade decline (Semmens et al., 2016; Schultz et al., 2017). Nonetheless, the crash
of the western population (Figure 1) in winter 2018–2019 was particularly stunning. In
2017, we estimated that the overwintering population had dropped by 97% of its average
historic abundance, from 3 to 10 million to 200–300 thousand butterflies (Schultz et al.,
2017). In winter 2018-2019, the population plummeted to fewer than 30,000 monarchs,
Pelton et al. Western Monarch Population Plummets
FIGURE 1 | Western monarchs breed west of the Rocky mountains
and primarily overwinter at over 200 sites (black points) along the Pacific coast
in California. During the spring, monarchs leave the overwintering habitat
(colored blue) to disperse (orange arrows) across the West. The butterflies
breed continuously across the West during the summer (colored white); in the
fall, they return (blue arrows) to the overwintering grounds. [Tag recoveries in
Mexico show that at least some western monarchs migrate to central Mexico,
mixing with the eastern monarch overwintering population; whether or not
monarchs from Mexico return to the West in the spring has not been
documented, but is suspected (dashed orange arrow)]. The authors have
monitored monarch breeding phenology and milkweed at 12 sites throughout
the West (orange points) as part of a multi-year study.
which represents a single year drop of 86%, and a >99% drop
since the 1980s (Figure 2A).
In this Perspective, we: (1) Place the current status in
context, both how trends compare to the eastern population
and potential implications of dropping to unprecedentedly low
abundance in the West, (2) Highlight the most likely window
during the annual life cycle when the population declined,
(3) Review probable causes of long-term declines, and (4)
Use our understanding of drivers of declines to recommend
steps that the public, policy makers, and land managers can
take including identifying knowledge gaps for which focused
mechanistic studies could contribute to developing more effective
and efficient conservation actions.
STATUS OF WESTERN MONARCHS IN
WINTER 2018–2019
Since 1997, volunteers have estimated the overwintering
population in California each fall at coastal groves (Xerces Society
Western Monarch Thanksgiving Count, 2019). The 2018 Xerces
Thanksgiving Count revealed a new low—only 28,429 monarchs
were tallied—<1% of the historic population (Figure 2A). The
current trend in western monarchs is in contrast to eastern
monarchs, which hit the highest estimated population size in
the last decade in winter 2018–2019 with 6.05 hectares occupied
(Rendón-Salinas et al., 2019).
We know from our past analyses that a western population
of <30,000 butterflies is unprecedented. The 2018 Thanksgiving
count mirrors a textbook extinction vortex (Gilpin and Soule,
1986), in the sense that fluctuations in abundance—which have
been happening throughout the past 30 years—become riskier as
the population becomes smaller. As populations become smaller,
“ordinary” environmental variation can cause a population to
drop below a point from which extinction is inevitable, unless
extraordinary measures are taken. We call this point the quasi-
extinction threshold. In 2016, a group of experts proposed
30,000 butterflies as the quasi-extinction threshold for western
monarchs (Schultz et al., 2017). Now, it is suddenly imperative to
know if the experts were correct, and, if so, what extraordinary
measures need to be taken to preserve the population.
In general, we know very little about what happens when
formerly large populations become small. Individuals in small
populations may have reduced mating success, suffer increased
predation, and lose other benefits of schooling or flocking
(Courchamp et al., 1999). These effects due to small population
size are known as “Allee effects” and are difficult to estimate
in wild populations because they are only expressed after a
population has begun to decline to extinction (Liermann and
Hilborn, 1997). Therefore, setting quasi-extinction thresholds
is one of the most subjective steps of population viability
analysis (e.g., Frick et al., 2010; McGowan et al., 2017). If the
published quasi-extinction threshold is correct, then positive
density-dependent processes associated with Allee effects could
lead to further rapid decline. If the quasi-extinction threshold is
incorrect, we will see the western monarch recover to a larger
population size. Regardless, this serves as a call to intensify efforts
to boost abundance to healthy enough numbers in the wild for
the population to be able to sustain itself through normal ups and
downs in the population size.
ENVIRONMENTAL DRIVERS
Causes of Rapid Decline From 2017 to 2018
Given the large drop in western monarchs from 2017 to 2018,
some are tempted to blame the weather for the low numbers.
Late rainy season storms swept across California in March. There
was a severe and extended wildfire season in the West and smoke
was widespread at times. California is still recovering from a
historic drought. Large amplitude inter-annual fluctuations are
an intrinsic aspect of butterfly population dynamics, and causes
of year-to-year variation are not necessarily the same as the
causes of long-term declines. Nonetheless, it is important to try
to understand western monarch abundance throughout the year
from winter 2017–2018 through winter 2018–2019, when the
decline occurred.
Starting in winter 2016–2017, the Xerces Society and
volunteers began a second count at overwintering sites, the New
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Pelton et al. Western Monarch Population Plummets
FIGURE 2 | Western monarch abundance at (A) overwintering and (B) breeding sites. In both panels, shaded areas show 95% confidence limits. (A) Western
monarch butterfly 1981–2018 estimates for overwintering abundance during the Thanksgiving Count time period in coastal California. Estimates for 1981–2017 were
calculated with state space models (Schultz et al., 2017), scaled to be comparable to the raw count from 2018 (shown). (B) Monarch egg and larva counts per stem
at all 12 monitoring sites (shown in Figure 1) throughout the season in 2017 and 2018. Curves were fitted with generalized additive models (Wood, 2011) to show
general trends in abundance. The fact that the two curves are parallel suggests that densities were lower by the time monarchs arrived in 2018; the decline does not
appear to be due to different dynamics during breeding. Note the log scale and 10-fold difference among years.
Year’s count (centered around New Year’s Day, to complement
the Thanksgiving Count 6 weeks earlier). Monarch abundance at
the New Year’s Count had declined by 43% on average in 2017
(n=44 sites), 49% on average in 2018, (n=115 sites) and 36%
in 2019 (n=130 sites), when compared to monarch abundance
at those same sites during the Thanksgiving Count. These data
suggest that monarch butterflies did not have exceptionally low
survival between November 2017 and January 2018, compared to
the previous year.
In addition to counts at overwintering sites, we started
monitoring summer breeding of western monarchs in 2017 at
12 sites throughout the West (Figure 1). Across these 2 years,
the density of monarch eggs and larvae was consistently lower in
2018 than 2017 (Figure 2B), with about a 10-fold decline between
the 2 years (average immature monarchs/stem =0.0273 [95% CI
=0.0025, 0.2953] in 2017 and 0.0022 [95% CI =0.0001, 0.0429]
in 2018; paired t-test of site averages between years: t= 2.53, df
=10, P=0.030). We therefore suggest that the drop measured
at Thanksgiving 2018 originated before the beginning of the 2018
breeding season, either late during the overwintering season or
very early in the breeding season.
This inference is consistent with Espeset et al. (2016) who
concluded that western monarch declines were concentrated in
early spring. Of the environmental events that seemed “unusual”
in 2017–2018, this pattern points to the possible negative effects
of unusually heavy rains in March 2018 with the caveat that many
other factors may have caused the population drop, including the
interaction of weather with habitat quality at overwintering sites,
and habitat inland from the coast in California, where the first
generation breeds.
Causes of Long-Term Declines
In the larger eastern population, declines have largely been
attributed to overwintering habitat loss (Brower et al., 2012;
Vidal et al., 2013) and breeding habitat loss, especially through
the use of herbicides (e.g., Pleasants and Oberhauser, 2012;
Flockhart et al., 2014). We (Crone et al., in press) recently
evaluated climate and land use factors simultaneously as potential
drivers of western monarch abundance. Trends in abundance
were more strongly associated with land use variables including
coastal development in overwintering areas and pesticide use
(glyphosate and neonicotinoid insecticides) in breeding areas
than climate variables in both overwintering and breeding areas
(Crone et al., in press). These results are consistent with the
hypotheses that overwintering habitat loss and loss of central
California breeding habitat are important for western monarchs
(see Espeset et al., 2016) and that trends in pesticide use likely
contribute to declining monarch populations as well as declines
in other butterfly taxa (see also Forister et al., 2016).
In addition to this broad scale analysis, we estimated
daily survival using data from Tuskes and Brower (1978),
for comparison with population declines estimated from
Thanksgiving and New Year’s counts. Daily survival at Natural
Bridges near Santa Cruz was 0.995 (95% CI 0.988, 0.997) and
at Santa Barbara was 0.991 (0.989, 0.993). Over 6 weeks (the
approximate time between Thanksgiving to New Year’s counts),
this historical estimate translates into a 29% drop (95% CI
12–40%) using estimates from Santa Cruz and a 32% drop
(95% CI 26–37%) using estimates from Santa Barbara. Hence,
based on the best available evidence, apparent survival during
winter in recent years (36–49% drop) has been lower than
it was in the past. This change reinforces the importance of
overwintering habitat quality on the long-term decline of the
western monarch population. At the present time, we have not
found comparable data to evaluate whether breeding season
survival or reproduction have changed in western monarchs.
URGENT STEPS FOR CONSERVATION
To date, western monarchs have received far less conservation
attention and financial resources than the larger eastern
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Pelton et al. Western Monarch Population Plummets
population. Nonetheless, the western monarch breeds
across most of the US west of the Rocky Mountains,
a significant portion of the monarchs overall North
American range. It makes an important contribution to the
resilience, redundancy, and representation of the species as a
whole (see definition in Shaffer and Stein, 2000).
While the precise causes of the recent dramatic drop in
the western population, as well as the longer term decline,
remain unknown, this knowledge gap should not prevent
conservation action. We suggest that a precautionary approach
be taken to remediate potential causes of decline. Specifically
we recommend efforts (1) to protect, enhance, and actively
manage overwintering sites; (2) to protect monarch habitat
from pesticides, particularly systemic insecticides (including
neonicotinoids); (3) to supplement larval and adult resources-
especially in the early spring-in California; (4) to identify, protect,
and enhance monarch habitats throughout the West, and (5)
to prioritize research efforts to answer questions critical to
developing an effective and efficient recovery strategy. Here, we
briefly explain our recommendations, and their relationship to
the causes of western monarch declines, described above. These
recommendations and relevant resources are expanded in in our
“Western Monarch Call to Action.”1
Protect, Manage, and Restore
Overwintering Habitat
Our analyses (“Environmental drivers” above) point to the
importance of monarch habitat in winter and early spring,
prior to the breeding season. Conservation biologists have long
known that efforts focused only on one stage of a species’ life
cycle (e.g., breeding) may not be sufficient if populations are
limited by another life stage [e.g., overwintering (Brown et al.,
2017)]. Despite the importance of monarchs to Californians
and the state’s tourism economy, few overwintering sites
are meaningfully protected (International Environmental Law
Project and the Xerces Society, 2012) and sites continue to be
destroyed—indeed, from 2017 to present, over one dozen sites
have either been newly destroyed or are reported to be threatened
by inappropriate tree trimming, removal, and/or development
(Xerces Society Western Monarch Overwintering Sites Database
2019, unpublished). To protect remaining habitat, overwintering
sites could be designated as Environmentally Sensitive Habitat
Areas (ESHAs) by the California Coastal Commission, protected
as Critical Habitat if monarchs were listed under the federal
Endangered Species Act, protected by California Department of
Fish and Wildlife if monarchs were listed as endangered under
the California Endangered Species Act, or a new law could be
created by the California state legislature to protect overwintering
sites from destruction.
To address the need for active management of overwintering
sites, the majority of which occur on publicly owned land, a
greater financial investment is needed. The Monarch Butterfly
and Pollinator Rescue Program (California Assembly Bill 2421),
was signed into law in 2018, and $3 million was allocated to this
program. An additional $3.9 million was recently allocated for
1www.savewesternmonarchs.org
restoration of overwintering sites owned by the City of Goleta.
While these represent important steps forward, more resources
are needed to restore and manage the over 200 actively used
overwintering sites. While there are no published estimates,
restoring a significant number of overwintering sites would easily
require tens of millions of dollars and, more importantly, would
benefit from sustained funding to continue to manage the groves
for monarchs in the long-term. Of the Top 50 priority sites
identified by Pelton et al. (2016) many of the most important
sites are owned by the California Department of Parks and
Recreation, followed by cities, U. S. Department of Defense,
East Bay Regional Parks District, and county, university, and
other state and federal agencies as well as private entities. Some
of these owners do not encourage or permit the planting of
eucalyptus (the dominant tree used by monarchs in California
during overwintering), nor are these land managers necessarily
focused on managing for monarch overwintering habitat—and,
in some cases, may be unaware of the full extent of overwintering
habitat within their jurisdiction.
Protect Monarchs and Their Habitat From
Pesticides
In our analyses of long-term trends, insecticide and herbicide
use were almost as tightly associated with monarch declines as
overwinter habitat loss. Restricting insecticide and herbicide use
increases adult Lepidoptera abundance (Frampton and Dorne,
2007). Broadcast herbicide use can kill host and nectar plants
and have non-target effects on butterflies (Stark et al., 2012).
We advise protecting the most important monarch breeding
and overwintering habitats from insecticide and herbicide use.
Specifically, we recommend avoiding herbicide applications
that damage monarch breeding and migratory habitat such as
milkweed and wildflowers. These recommendations apply to
home gardens and lawns, as well as lands used for agriculture and
other purposes. If herbicides are used, we advise using targeted
application methods, avoiding large-scale broadcast applications
of herbicides, and taking precautions to limit off-site movement
of herbicides. Neonicotinoid insecticides, in particular, should be
avoided at all times in monarch habitat due to their persistence,
systemic nature, and toxicity. When purchasing milkweeds
or wildflowers from nurseries, we recommend ensuring that
they have not been treated with neonicotinoids or other
systemic insecticides.
Restore Breeding and Migratory Habitat in
California
Enhancing monarch breeding habitat may be able to partly
mitigate reductions in overwintering habitat quality because
larger populations at the end of the summer can potentially
withstand higher mortality. Numerous studies have quantified
the importance of host and nectar plants for butterfly populations
(Dennis et al., 2006; Dennis, 2010), and restoration efforts
which enhance host and nectar have been effective approaches
for the conservation of rare butterflies (Carleton and Schultz,
2013). We recommend planting native milkweeds in areas
where they historically grew in California, and, in particular,
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Pelton et al. Western Monarch Population Plummets
in the Coast Range, Central Valley, and the foothills of the
Sierra Nevada, areas where the first generation of monarchs are
produced each spring. Early emerging native species that may
be particularly important in spring include woollypod (Asclepias
eriocarpa), California (A. californica), and heartleaf milkweed
(A. cordifolia). However, commercial availability of these species
is limited. Later-emerging native California milkweed species
that are more readily available, and may also help, include
narrowleaf (A. fascicularis) and showy milkweed (A. speciosa).
In the desert southwest of California, we recommend rush
(A. subulata) and desert milkweed (A. erosa). We recommend
only planting milkweed >5 miles inland from overwintering
sites, as milkweed does not naturally grow close to the
coast north of Santa Barbara and milkweed at overwintering
sites can interrupt natural overwintering behavior. Tropical
milkweed (A. curassavica) is exotic to California, disrupts the
monarchs migratory cycle, and serves as a reservoir for monarch
pathogens (Satterfield et al., 2016). As such we recommend
against planting tropical milkweed. In places where tropical
milkweed already exists, we recommend cutting it back to
the ground in the fall (October/November) and repeatedly
throughout the winter to mimic native milkweed phenology
and break the disease cycle; ideally, it should be replaced by
native milkweed.
In addition, we recommend planting nectar-rich wildflowers,
especially those that bloom early in the spring (February–April)
and fall (September-October). If located close to the coast,
plants which bloom in the winter (November-January) may also
be useful.
Protect, Manage, and Restore Summer
Breeding and Fall Migration Monarch
Habitat Throughout the Western Monarch’s
Range
Identifying key areas of breeding and migrating habitat for
monarchs in the West remains a knowledge gap. Some
geographic regions contribute disproportionately to the eastern
monarch overwintering population in Mexico (e.g., Flockhart
et al., 2017), and it is important to know whether the same
is true for western monarchs. No data exist from which we
could meaningfully evaluate their importance for short- or
long-term population declines. Thus, while some of the most
important monarch habitat within its western breeding (Yang
et al., 2016; Dilts et al., 2019) and overwintering (Pelton
et al., 2016) range has already been identified, additional work
is needed to identify and rank these areas. We recommend
identifying existing monarch habitat, ensuring that it is managed
to protect monarchs (Xerces Society, 2018) and in some regions
and landscape types, we recommend habitat enhancement
or restoration. Habitat restoration in regions where monarch
habitat historically occurred, but have likely been lost (such
as the Columbia Plateau and Snake River Plain), as well as
riparian areas, are high priority areas outside of California.
Such restoration would likely benefit from habitat elements
beyond milkweed and nectar, such as shrubs or trees for roosting
and shade.
Fill Research Gaps to Inform Western
Monarch Recovery Strategies
Breeding and migrating habitat are only a few of the gaps
in our knowledge of western monarchs. We especially need
observations of monarch biology in places where human
populations are low (e.g., the Great Basin desert) and at times
of year when monarch butterflies are sparse (e.g., early spring
in western California, just as they leave the overwintering
grounds). We urge volunteers across the West to collect
observations of monarchs and milkweeds, especially in the early
spring (February–April), the period in which monarchs typically
leave the overwintering sites. Together these observations will
help answer questions about monarch breeding phenology.
In this year, when numbers are low in the West and high
in the East, targeted observations of monarch adults and
larvae may also tell us whether the West sees an influx of
monarchs arriving from Mexico (see Pyle, 2015). Monarch
adult, larva, egg, nectaring, and milkweed sightings can be
reported to the Western Monarch Milkweed Mapper2and
first adults observed can be reported to Journey North3
as well. More robust monitoring may be achieved through
increased western participation in the Integrated Monarch
Monitoring Program4.
We urge academic ecologists to conduct targeted
experimental and observational studies to complement
large-scale observations like the ones described above. In
both the eastern and western monarch populations, filling
knowledge gaps about demography throughout the life cycle
would allow us to design quantitative thresholds for conservation
and restoration. For example, it may be possible for targeted
actions at one point in the life cycle to make up for stresses
at other points. If climate change is making the landscape less
favorable, can we make up for this with improved breeding
or overwintering habitat quality and/or area? Can more
breeding habitat in the outer parts of the breeding distribution
make up for habitat loss at breeding or overwintering sites
in California? Intuitively, the answer is probably “yes, but
only partly.” To answer this in a more quantitative way,
we need a better understanding of how the life cycle pieces
fit together.
CONCLUSION
In closing, western monarchs are currently in peril. Their status
reflects a long-term decline due to some combination of habitat
loss and degradation in their overwintering and breeding range,
increased pesticide use, and possibly climate change. The recent
dramatic drop reflects conditions when the least is known about
western monarchs—where they are, what habitat they are using,
and what they need to survive, migrate and reproduce. In spite of
their current status, monarchs are resilient; we believe that rapid
conservation actions can recover the population. This recovery
will require the protection of monarchs and their habitat, as well
as targeted research to understand the unique life cycle of western
2www.monarchmilkweedmapper.org
3https://journeynorth.org/monarchs
4https://monarchjointventure.org/immp
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Pelton et al. Western Monarch Population Plummets
monarch butterflies. If we are going to take these actions, the time
is now.
DATA AVAILABILITY
The datasets for this study will not be made publicly
available because restrictions apply to some of the
datasets. Some of the datasets are in a publicly
accessible repository:
The Xerces Society Western Monarch Thanksgiving and New
Year’s Counts analyzed in this study can be found at www.
westernmonarchcount.org/data.
Restrictions apply to some of the datasets:
The Xerces Society Western Monarch Overwintering
Sites Database 2019 is not publicly available because
of privacy concerns with a subset of the information.
Requests to access the database should be directed to Emma
Pelton, monarchs@xerces.org.
The western monarch and milkweed phenology dataset
summarized in this manuscript are not publicly available because
it is part of a study currently in-progress. Requests to access the
datasets should be directed to Cheryl Schultz, schultzc@wsu.edu.
AUTHOR CONTRIBUTIONS
EP, SJ, and SB (along with others—see Acknowledgments)
oversee Thanksgiving and New Year’s Counts and maintain the
overwintering sites database. All authors contributed to funding
and implementing the 2017–2018 surveys in the breeding range.
EC conceived and ran all analyses with input from CS and EP. All
authors wrote and revised the manuscript.
FUNDING
Funds for the 2017–2018 breeding and phenology surveys
and analysis were provided by Department of Defense Legacy
Natural Resources Program (NR 16 Western Monarch) and the
U.S. Fish & Wildlife Service Coastal Program. Authors were
supported by their institutions (WSU, Tufts and Xerces Society)
and EC, SJ, EP, and CS were partly supported by the National
Science Foundation (NSF DEB 1920834).
ACKNOWLEDGMENTS
Thank you to the Western Monarch Thanksgiving Count
volunteers, particularly our regional coordinators, Mia Monroe,
and Katie Hietala-Henschell of the Xerces Society; Stephanie
McKnight of the Xerces Society and Cameron Thomas of
Washington State University for conducting the fieldwork
for the breeding and milkweed phenology project; fellow
western monarch researchers and conservation practitioners
for conversations that led to the development to the Western
Monarch Call to Action; US Fish and Wildlife Service Coastal
Program, Department of Defense, National Science Foundation,
and Xerces Society members and other funders for supporting the
work presented in this Perspective.
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Conflict of Interest Statement: The authors declare that the research was
conducted in the absence of any commercial or financial relationships that could
be construed as a potential conflict of interest.
Copyright © 2019 Pelton, Schultz, Jepsen, Black and Crone. This is an open-access
article distributed under the terms of the Creative Commons Attribution License (CC
BY). The use, distribution or reproduction in other forums is permitted, provided
the original author(s) and the copyright owner(s) are credited and that the original
publication in this journal is cited, in accordance with accepted academic practice.
No use, distribution or reproduction is permitted which does not comply with these
terms.
Frontiers in Ecology and Evolution | www.frontiersin.org 7July 2019 | Volume 7 | Article 258
... Land use may reflect habitat loss, a potential driver in the Western Monarch decline [9,10,30]. Therefore, for the COR model, we predicted certain land cover classifications would have positive associations with overwintering monarchs, such as forest and shrublands [30], that provide winter roost sites and winter nectar [14,23,31,32]. Conversely, we predicted that some categories, like moderate/high levels of development, could be negatively associated with monarch presence, implying low overwintering habitat availability. ...
... Western Monarchs are under threat of a long-term decline [9,80]. Protecting overwintering habitat is a key step in conservation planning for the long-term success and recovery of the western migratory population [10,81]. The multi-scale species distribution model presented here took a large-scale approach to identifying the climatic and local landscapelevel predictors of quality overwintering habitat. ...
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Western Monarch butterflies (Danaus plexippus) migrate from inland breeding ranges to coastal overwintering grounds in California. Given that migratory individuals may make multi-scale habitat selection decisions, we considered a multi-scale species distribution model (SDM) using range-wide climatic and local landscape-level predictors of migratory and overwintering habitat and community-science presence data. The range-wide model output was included as a predictor in the local-scale model, generating multi-scale habitat suitability. The top range-wide predictor was the minimum temperature in December, contributing 83.7% to the model, and was positively associated with presence. At the local scale, the strongest predictors of presence were the range-wide output and percent coverage of low and medium levels of development, contributing > 95%, with 61–63% from the range-wide output, with local-scale suitability coinciding with the California coastal zones. Development’s positive association with overwintering monarch presence was counterintuitive. It is likely that our local-scale model is overfit to these development zones, but it is unclear whether this overfitting resulted from modeler choices, monarchs overwintering close to human development, biased detection near human development, or a combination of these factors. Therefore, alternative approaches to collecting local-scale attribute data are suggested while recognizing the primacy of climate in restricting overwinter sites.
... For the eastern monarch population, both the timing of the fall migration southward (Culbertson et al., 2022), and spring arrival northward have gotten later over recent decades (Howard & Davis, 2015), although some portions of the eastern population have shown no change in fall migration timing (Ethier & Mitchell, 2023). Declines in the eastern North American monarch population (Brower et al., 2012;Pelton et al., 2019;Thogmartin et al., 2017;Vidal & Rendón-Salinas, 2014) have led to conservation efforts from national (USFWS, 2024) and international (Normile, 2023) perspectives. ...
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Phenological changes have been widely documented in animal and plant responses to directional environmental change. However, predicting the consequences of these shifts for species interactions and population viability requires knowledge of vital rate responses to biotic and abiotic drivers. Here, we paired long‐term phenology data documenting monarch butterfly abundance and occurrence of their milkweed hostplant with outdoor experiments in the central United States to ask how changes in spring arrival times to monarch breeding sites affect their development, survival, and within‐season population growth. Monarch arrival times did not change across the 17 years of monitoring, but the peak abundance of monarchs, which occurred just prior to their fall migration, shifted 9 days later in 2019 as compared to 2003. Summer population growth declined from 2003 to 2019, significant in ~80% bootstrap calculations. Phenological changes in milkweed occurrence mirrored changes in monarch abundance, happening later through time. Our field experiment showed that early season larval survival was highest when the timing of hatching matched the average timing of the first natural monarch cohort; survival was lowest when egg hatching shifted 14 days earlier. The results of our study indicate that earlier arrival of adult monarchs to summer breeding habitat would be costly for monarchs—but field survey data show that arrival times have not changed to date. Instead, the local changes we observed in the timing of peak abundance occurred towards the end of the breeding season, not the onset. At present, we conclude that changes in early season phenology are not a threat to eastern North American monarchs living in the central United States, but drivers of breeding‐season growth rates and changes in late‐season phenology merit further study, both in the central United States and in other parts of the monarch's range.
... Over the last two decades, eastern and western United States monarch butterfly (Danaus plexippus plexippus; Figure 1) populations have experienced significant declines, and in response, concern and causal factors have stimulated considerable interest, resulting in various hypotheses and conservation efforts (Pelton et al., 2019;Pocius et al., 2017;Thogmartin et al., 2017;USFWS, 2020a;Voorhies et al., 2019;Zaya et al., 2017). In December 2020, the US Fish and Wildlife Service (USFWS) listed the monarch butterfly in the US Federal Register as a candidate species under the Endangered Species Act of 1973 (USFWS, 2020b). ...
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Population declines of the monarch butterfly (Danaus plexippus plexippus) in North America have largely been attributed to the distribution and condition of species‐specific preferred nectar sources. In 2020, the US Fish and Wildlife Service (USFWS) listed the monarch butterfly in the US Federal Register as a candidate species under the Endangered Species Act of 1973. The USFWS ranked the availability, quality, and spatial distribution of nectar plants during autumn migration as the fourth most contributing factor to US monarch population declines. During the autumn migration through the Great Plains, monarchs seek nectar plants to accumulate lipid reserves for further migration to and overwintering in Mexico. We applied vegetation and rangeland health data from the US Department of Agriculture, Natural Resources Conservation Service, National Resources Inventory (NRI) to quantify species density and richness of monarch‐preferred nectar plants, associated rangeland conditions, and diversity of nectar sources along this autumn migration pathway. We focused specifically on longitudinal gradients W‐095‐100 and W‐100‐105 and discrete 5° latitudinal–longitudinal cells within those gradients. The respective NRI dataset spans 8211 rangeland sites sampled between 2009 and 2018. Approximately 84.4% of sites in W‐095‐100 and 72.5% of sites in W‐100‐105 contained monarch‐preferred nectar plants. Preferred nectar plants made up 7.4% of 2438 identified plant species in W‐095‐100 and 6.1% of 2371 identified plant species in W‐100‐105. For W‐095‐100, preferred nectar plant densities were highest for the 5° cell covering portions of US states Oklahoma and Kansas and lowest for the 5° cell at the US–Mexico border. In W‐100‐105, preferred nectar plant densities decreased linearly from north to south. Preferred nectar plant densities were greater for 5° cells in W‐100‐105 (50.5 billion plants) as compared with W‐095‐100 (44.4 billion plants). Consistent with trends in preferred nectar source density, rangeland conditions assessed by similarity indices and rangeland health protocols were generally lowest for 5° cells spanning the US–Mexico border. The results provide the most comprehensive assessment to date for preferred nectar sources of the monarch butterfly along the Great Plains autumn migration to Mexico and document generally decreasing nectar sources and habitat conditions at southern latitudes in this ecologically important pathway.
... Monarch butterfly (Danaus plexippus) populations in North America have declined precipitously over the past several decades (Pelton et al. 2019;Monarch Watch 2022), prompting increased public messaging to plant milkweed wherever possible, particularly in agricultural areas which are seen by proponents of the milkweed limitation hypothesis to be essential for monarch recovery efforts (Thogmartin et al. 2017). However, agricultural landscapes throughout the United States have become increasingly toxic to insects in recent decades, driven largely by the rise in the use of highly potent neonicotinoid insecticides (Douglas et al. 2020). ...
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Milkweed plants in agricultural landscapes throughout the United States and southern Canada are believed to be vitally important for the imperiled monarch butterfly (Danaus plexippus) populations. However, studies have found that these plants often assimilate a slew of pesticides from the surrounding landscape, including highly potent and widely applied neonicotinoid insecticides. This has prompted concern over the potential impacts of these compounds on monarch populations and has created a growing interest in determining the direct lethal and sublethal consequences of exposure. Fewer studies have considered how neonicotinoids may interact with milkweed defensive chemistry to indirectly influence monarch performance. Here we addressed this question by investigating whether uptake of a widely used neonicotinoid insecticide, clothianidin, could alter milkweed (Asclepias syriaca) defense responses and subsequently impact monarch growth and feeding. We found that clothianidin-treated milkweed plants grew taller, and produced monarch larvae that weighed more and consumed more leaf tissue than larvae feeding on control plants. After five days of monarch herbivory, clothianidin-treated plants had higher levels of the phytohormone, jasmonic acid, but similar levels of salicylic acid relative to control plants. Neither latex nor cardenolide production was impacted by clothianidin assimilation. Overall, these findings indicate that clothianidin exposure can improve the vitality of common milkweed plants, and may subsequently impact monarch performance.
... The butterfly can be subdivided into two populations: a western population that overwinters on the Pacific coast of California and then migrates to breeding ground west of the Rocky Mountains, and an eastern population with two-way migration from an overwintering area in Mexico to breeding ground across eastern North America and back to Mexico in autumn (Gao et al. 2020). Studies have demonstrated a decline in monarchs' population at overwintering and breeding grounds in both western and eastern populations (Pelton et al. 2019;Wilcox et al. 2019;Grant et al. 2021;Kendrick and McCord 2023). ...
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Climate warming and landscape level changes are considered to be among the most serious of anthropogenic stresses to the environment. Consequences of land-use change include habitat loss and fragmentation, which are considered to be primary forces in the decline of species worldwide. Climate change not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human-mediated threats. Understanding species’ response to landscape structure and climate change can inform our understanding of their ecology and aid in the development of management and conservation strategies. Monarch butterflies (Danaus plexippus) are well-known long-distance migrants whose population has suffered a drastic decline in recent decades. Here I investigated response of monarchs’ breeding phenology to climate warming and also assessed landscape characteristics associated with monarchs’ breeding-site and habitat selection during different seasons. My result demonstrated that the breeding phenology of monarchs was more responsive to temperature in warmer regions than in colder ones. I also found that, although often neglected, landscape compositional and configurational heterogeneity can be of great importance for monarch conservation. I recommend that the landscapes surrounding monarch’s ideal habitat, like grasslands, be managed to maintain a diversity of land cover types. Implications for insect conservation: My finding demonstrates temporal shifts in the breeding phenology of monarchs in response to global warming, which can lead to a mismatch between a butterfly and its host plants and can further alter the species interactions. My results also showed the importance of landscape compositional and configurational heterogeneity along with grassland patches for the monarch population. Therefore, wildlife managers should focus on maintaining diverse land-cover types with different landscape structures for better management of monarch populations. Implications for conservation In today’s world, land use, land cover change, and climate change have led to a decline in biodiversity. Climate change is causing significant shifts in insect phenology, which can affect insect abundance, species interactions, and ecosystem services. My finding provides evidence of temporal shifts in the breeding phenology of monarchs in response to global warming, which can lead to a mismatch between a butterfly and its host plants and can further alter the species interactions. Even though we cannot control the climate, this knowledge may help to better prepare for monarch conservation. My results showed the importance of landscape compositional and configurational heterogeneity along with grassland patches for the monarch population. As conservation resources become increasingly limited, targeted management at the landscape level is necessary to get the most results for the conservation dollar. Based on my findings, instead of only focusing on maintaining grassland areas, managers should equally focus on maintaining diverse land-cover types with different landscape structures.
... First, predicting specialists allows for more targeted conservation because specialists are more strongly dependent on a limited suite of hosts to complete their life cycle. For example, planting milkweed, the larval host of the declining monarch butterfly, is seen as a critical part of this specialist's recovery strategy (Pelton et al. 2019). For bees, diet breath information could be used to inform seed mixes used to increase bee diversity in agriculture (Morandin and Kremen 2013;Seitz et al. 2020) and urban areas (Gerner and Sargent 2022). ...
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An animal’s diet breadth is a central aspect of its life history, yet the factors determining why some species have narrow dietary breadths (specialists) and others have broad dietary breadths (generalists) remain poorly understood. This challenge is pronounced in herbivorous insects due to incomplete host plant data across many taxa and regions. Here, we develop and validate machine learning models to predict pollen diet breadth in bees, using a bee phylogeny and occurrence data for 682 bee species native to the United States, aiming to better understand key drivers. We found that pollen specialist bees made an average of 72.9% of their visits to host plants and could be predicted with high accuracy (mean 94%). Our models predicted generalist bee species, which made up a minority of the species in our dataset, with lower accuracy (mean 70%). The models tested on spatially and phylogenetically blocked data revealed that the most informative predictors of diet breadth are plant phylogenetic diversity, bee species’ geographic range, and regional abundance. Our findings also confirm that range size is predictive of diet breadth and that both male and female specialist bees mostly visit their host plants. Overall, our results suggest we can use visitation data to predict specialist bee species in regions and for taxonomic groups where diet breadth is unknown, though predicting generalists may be more challenging. These methods can thus enhance our understanding of plant-pollinator interactions, leading to improved conservation outcomes and a better understanding of the pollination services bees provide.
... 5 The monarch butterflies west of the Rockies are at a high risk of local extirpation, likely due to environmental changes in that region. 62 However, the strong migratory connectivity of the species largely prevents the larger census size east of the Rockies from bolstering the western numbers (but see 63 ). Historically, the Afro-Palearctic population of painted ladies has demonstrated long-term demographic sta-bility, albeit with large short-term population fluctuations, partially due to outbreak dynamics. ...
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Some insects, such as the painted lady butterfly Vanessa cardui, exhibit complex annual migratory cycles spanning multiple generations. Traversing extensive seas or deserts is often a required segment of these migratory journeys. We develop a bioavailable strontium isoscape for Europe and Africa and then use isotope geolocation combining hydrogen and strontium isotopes to estimate the natal origins of painted ladies captured north and south of the Sahara during spring and autumn, respectively. Our findings reveal moderate migratory connectivity across the Sahara characterized by a broad-front, parallel migration. We also report evidence of a leapfrog migration, wherein early autumn migrants from higher latitudes cover greater distances southward than their late autumn counterparts. This work represents a major advancement in understanding insect migratory patterns and connectivity, particularly across extensive barriers, which is essential for understanding population dynamics and predicting the impacts of global change on insect-mediated ecosystem services.
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Introduction Identifying habitat of migratory species to effectively support conservation and management requires careful consideration of (1) the data used to inform habitat models, (2) the biology of the organism, (3) land tenure, and (4) the needs of the target audience. Methods To provide this information for western U.S. monarch butterflies, a population undergoing decline, we modeled habitat during the spring and fall migrations. Our approach controlled for biases in citizen science locality data, the principal source of monarch observation data, and incorporated needs for milkweed host plants in the spring and nectar plants in the fall. Results and Discussion The results showed the distribution of habitat for spring and fall migration, where the Coast Range and Central Valley in California and riparian areas throughout the range were particularly important. Just 29% of predicted habitat for spring and fall migrations, combined, overlapped between the two seasons. Although the U.S. federal government manages 53% of the land in the western U.S., government land makes up just 11.7% of the spring migration range and 23.5% of the fall migration range. State and local governments and non-governmental organizations (NGOs) manage an additional 4.2% of the spring and 4.0% of the fall migration habitat. Thus, like eastern monarchs, western monarchs rely heavily on land under private ownership for their migration and to be successful, monarch conservation efforts must embrace a public-private approach. Among federal agencies, the Bureau of Land Management (BLM; 5.9% of spring and 9.7% of fall habitat) and Forest Service (3.3% of spring and 9.2% of fall habitat) manage the greatest shares. Less than half of the government and NGO owned habitat for both migrations is managed for biodiversity conservation rather than multiple uses (spring, 46.0%; fall, 36.5%). We created custom model outputs for the BLM to highlight areas of both regional and local importance for migrating monarchs in each BLM administrative unit, enabling managers across the agency to contribute to recovery. The outcomes provide input at a relevant spatial scale to support actions such as habitat restoration, riparian zone protection, and pesticide use reduction to enhance conditions for migrating monarchs on both government and private lands.
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The monarch butterfly (Danaus plexippus) is a vagile species that undertakes an annual, multi-generational migration across North America. The abundance of this species at both eastern and western overwintering sites in Central Mexico and California indicates a population decline. Success of continental-scale conservation programs for a migratory species depends on providing, maintaining, and protecting habitats at appropriate temporal and spatial scales. Here, dynamics of monarch continental-scale migration and gene flow were obtained by combined stable isotope, morphological, and genetic analyses. These analyses were applied to temporal monarch samples collected from May to September during 2016–2021 at locations in Iowa, USA and spatial collections from Pennsylvania, Delaware, Iowa, Ohio, Nevada, Idaho, Hawaii, 3 Australian locations during July and August 2016, and Texas in April 2021. Evidence of seasonal multi-generational migration was obtained through δ2H analyses of spatial collections, which was corroborated by decreased wing hue (a morphological marker for non-migratory individuals). In Iowa, 10–15% of monarchs represented migrants from southern areas throughout the breeding season and 6% were migrants from the North in midsummer. Limited sequence variation detected across the mitochondrial genome impacted the capability to detect significant population genetic variation in our North American samples. However, 2 novel substitutions were identified and predicted to be fixed among Australia samples, contributing to intercontinental differentiation from counterparts in North America. Our assessment of temporal and spatial population dynamics across the North American monarch breeding range provides insight into continental-scale migration and previously undetected mitochondrial DNA variation among globally distributed populations.
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Monarch butterfly (Danaus plexippus L.) populations have declined in North America. The International Union for Conservation of Nature (IUCN) recently classified the species as endangered, sparking public concern and conservation efforts. Our approach to conservation is through cryopreservation of germinal cells and tissue. The goal of this study was to develop a cryopreservation protocol for monarch spermatozoa to ensure successful long-term storage. Cryopreserved sperm cells would provide a reserve of monarch germplasm, which could be utilized in the event of population loss. In this study, sperm cell bundles collected from male monarch butterflies were cryopreserved in a cryoprotective medium and stored in liquid nitrogen. To determine the post-cryopreservation sperm cell viability, a subsample of preserved sperm bundles were thawed rapidly, and their viability was qualified using a sperm live/dead stain. We are presenting a protocol to preserve and store genetic material and viable sperm bundles of the monarch butterfly. To date, this is the first report of successful cryopreservation of monarch germplasm which sets the foundation for cryostorage and could be extensible to other vulnerable lepidopterans.
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The monarch butterfly is one of the most easily recognized and frequently studied insects in the world, and has recently come into the spotlight of public attention and conservation concern because of declining numbers of individuals associated with both the eastern and western migrations. Historically, the larger eastern migration has received the most scientific attention, but this has been changing in recent years, and here we report the largest-ever attempt to map and characterize non-overwintering habitat for the western monarch butterfly. Across the environmentally and topographically complex western landscape, we include 8,427 observations of adults and juvenile monarchs, as well as 20,696 records from 13 milkweed host plant species. We find high heterogeneity of suitable habitats across the geographic range, with extensive concentrations in the California floristic province in particular. We also find habitat suitability for the butterfly to be structured primarily by host plant habitat associations, which are in turn structured by a diverse suite of climatic variables. These results add to our knowledge of range and occupancy determinants for migratory species and provide a tool that can be used by conservation biologists and researchers interested in interactions among climate, hosts and host-specific animals, and by managers for prioritizing future conservation actions at regional to watershed scales.
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Assessment of a species’ status is a key part of management decision making for endangered and threatened species under the U.S. Endangered Species Act. Predicting the future state of the species is an essential part of species status assessment, and projection models can play an important role in developing predictions. We built a stochastic simulation model that incorporated parametric and environmental uncertainty to predict the probable future status of the Sonoran desert tortoise in the southwestern United States and North Central Mexico. Sonoran desert tortoise was a Candidate species for listing under the Endangered Species Act, and decision makers wanted to use model predictions in their decision making process. The model accounted for future habitat loss and possible effects of climate change induced droughts to predict future population growth rates, abundances, and quasi-extinction probabilities. Our model predicts that the population will likely decline over the next few decades, but there is very low probability of quasi-extinction less than 75 years into the future. Increases in drought frequency and intensity may increase extinction risk for the species. Our model helped decision makers predict and characterize uncertainty about the future status of the species in their listing decision. We incorporated complex ecological processes (e.g., climate change effects on tortoises) in transparent and explicit ways tailored to support decision making processes related to endangered species.
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Introduced plants can positively affect population viability by augmenting the diet of native herbivores, but can negatively affect populations if they are subpar or toxic resources. In organisms with complex life histories, such as insects specializing on host plants, the impacts of a novel host may differ across life stages, with divergent effects on population persistence. Most research on effects of novel hosts has focused on adult oviposition preference and larval performance, but adult preference may not optimize offspring performance, nor be indicative of host quality from a demographic perspective. We compared population growth rates of the Baltimore checkerspot butterfly, Euphydryas phaeton, on an introduced host, Plantago lanceolata (English plantain), and the native host Chelone glabra (white turtlehead). Contrary to the previous findings suggesting that P. lanceolata could be a population sink, we found higher population growth rates (λ) on the introduced than the native host, even though some component parameters of λ were higher on the native host. Our findings illustrate the importance of moving beyond preference–performance studies to integrate vital rates across all life stages for evaluating herbivore–host plant relationships. Single measures of preference or performance are not sufficient proxies for overall host quality nor do they provide insights into longer term consequences of novel host plant use. In our system, in particular, P. lanceolata may buffer checkerspot populations when the native host is limiting, but high growth rates could lead to crashes over longer time scales.
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The butterfly fauna of lowland Northern California has exhibited a marked decline in recent years that previous studies have attributed in part to altered climatic conditions and changes in land use. Here, we ask if a shift in insecticide use towards neonicotinoids is associated with butterfly declines at four sites in the region that have been monitored for four decades. A negative association between butterfly populations and increasing neonicotinoid application is detectable while controlling for land use and other factors, and appears to be more severe for smaller-bodied species. These results suggest that neonicotinoids could influence non-target insect populations occurring in proximity to application locations, and highlights the need for mechanistic work to complement long-term observational data.
Chapter
Nestled amid the Appalachian Mountains of southwestern Virginia and northeastern Tennessee lies the Clinch Valley, the nation’s leading hot spot for imperiled aquatic organisms. The Clinch River is the only undimmed headwater of the Tennessee River basin, which in turn is the nation’s most biologically diverse drainage system. The surface waters of the Clinch run rich indeed: They are home to at least 29 rare mussels and 19 rare fish. Underground, the region’s limestone bedrock is honeycombed by more than a thousand caves and uncounted underground springs and streams. This little-known world is filled with a menagerie of rare beetles, isopods, and other subterranean insects. These underground realms have yielded more than 30 species new to science in just the past few years. The Clinch Valley is largely rural and sparsely populated. Most residents make their living directly from the land, either mining coal, harvesting timber, grazing cattle, or planting crops. These rural lifestyles have maintained much of the region in a relatively natural state, and more than two-thirds of the Clinch Valley remains forested. The forested hills mask a history of ecologically unsound land use practices, however, that have degraded the legendary quality of the region’s waterways. Virtually anything released in the valley flows downhill into the streams and rivers. Among the greatest threats to the valley’s extraordinary aquatic life are heavy metals leaching from abandoned coal mines, sediment eroding from cutover slopes, and nutrients released by streamside-grazing cattle. These and other threats have already taken a toll on the region’s extraordinary biological richness. Where once there were 60 kinds of freshwater mussels, only about 40 remain. Coastal southern California, in contrast, is one of the most densely populated regions in the nation. It, too, is a hot spot for imperiled species. Its dry Mediterranean climate and varied topography have favored the evolution of a host of unique plants and animals. Altogether, some 86 imperiled species are found along the coast and in the mountains of this nationally significant center of biodiversity. Certain areas stand out even by California standards as having a truly extraordinary diversity of rare species.
Article
Count-based PVA allows researchers to assess patterns of population change through time and to evaluate future persistence. We combined state-space models and citizen science data to evaluate viability of the western population of monarch butterflies over 36 years. A key feature of our analysis was combining irregular sampling from multiple sites to obtain a single estimate of total abundance using state-space models. The average population growth rate was negative, u = − 0.0762 (λ = 0.927), average abundance in the 2000s was < 5% of average abundance in the 1980s, and current quasi-extinction risk is 72% within 20 years. Despite wide confidence intervals in some parameter estimates, western monarch monitoring data provide unambiguous evidence for dramatic population declines. To obtain viable populations, managers could target historic abundance and high enough growth rates to avoid near-term extinction.
Article
Addressing population declines of migratory insects requires linking populations across different portions of the annual cycle and understanding the effects of variation in weather and climate on productivity, recruitment, and patterns of long-distance movement. We used stable H and C isotopes and geospatial modeling to estimate the natal origin of monarch butterflies (Danaus plexippus) in eastern North America using over 1000 monarchs collected over almost four decades at Mexican overwintering colonies. Multinomial regression was used to ascertain which climate-related factors best-predicted temporal variation in natal origin across six breeding regions. The region producing the largest proportion of overwintering monarchs was the US Midwest (mean annual proportion = 0.38; 95% CI: 0.36-0.41) followed by the north-central (0.17; 0.14-0.18), northeast (0.15; 0.11-0.16), northwest (0.12; 0.12-0.16), southwest (0.11; 0.08-0.12), and southeast (0.08; 0.07-0.11) regions. There was no evidence of directional shifts in the relative contributions of different natal regions over time, which suggests these regions are comprising the same relative proportion of the overwintering population in recent years as in the mid-1970s. Instead, interannual variation in the proportion of monarchs from each region covaried with climate, as measured by the Southern Oscillation Index and regional-specific daily maximum temperature and precipitation, which together likely dictate larval development rates and food plant condition. Our results provide the first robust long-term analysis of predictors of the natal origins of monarchs overwintering in Mexico. Conservation efforts on the breeding grounds focused on the Midwest region will likely have the greatest benefit to eastern North American migratory monarchs, but the population will likely remain sensitive to regional and stochastic weather patterns.
Book
Winner of the Marsh Book of the Year Award 2012 by the British Ecological Society. In A Resource-Based Habitat View for Conservation Roger Dennis introduces a novel approach to the understanding of habitats based on resources and conditions required by organisms and their access to them, a quantum shift from simplistic and ineffectual notions of habitats as vegetation units or biotopes. In drawing attention to what organisms actually use and need in landscapes, it focuses on resource composition, structure and connectedness, all of which describe habitat quality and underpin landscape heterogeneity. This contrasts with the current bipolar view of landscapes made up of habitat patches and empty matrix but illustrates how such a metapopulation approach of isolated patchworks can grow by adopting the new habitat viewpoint. The book explores principles underlying this new definition of habitat, and the impact of habitat components on populations, species' distributions, geographical ranges and range changes, with a view to conserving resources in landscapes for whole communities. It does this using the example of butterflies - the most alluring of insects, flagship organisms and key indicators of environmental health - in the British Isles, where they have been studied most intensively. The book forms essential reading for students, researchers and practitioners in ecology and conservation, particularly those concerned with managing sites and landscapes for wildlife.
Article
Summary. Recent work by Reiss and Ogden provides a theoretical basis for sometimes preferring restricted maximum likelihood (REML) to generalized cross-validation (GCV) for smoothing parameter selection in semiparametric regression. However, existing REML or marginal likelihood (ML) based methods for semiparametric generalized linear models (GLMs) use iterative REML or ML estimation of the smoothing parameters of working linear approximations to the GLM. Such indirect schemes need not converge and fail to do so in a non-negligible proportion of practical analyses. By contrast, very reliable prediction error criteria smoothing parameter selection methods are available, based on direct optimization of GCV, or related criteria, for the GLM itself. Since such methods directly optimize properly defined functions of the smoothing parameters, they have much more reliable convergence properties. The paper develops the first such method for REML or ML estimation of smoothing parameters. A Laplace approximation is used to obtain an approximate REML or ML for any GLM, which is suitable for efficient direct optimization. This REML or ML criterion requires that Newton–Raphson iteration, rather than Fisher scoring, be used for GLM fitting, and a computationally stable approach to this is proposed. The REML or ML criterion itself is optimized by a Newton method, with the derivatives required obtained by a mixture of implicit differentiation and direct methods. The method will cope with numerical rank deficiency in the fitted model and in fact provides a slight improvement in numerical robustness on the earlier method of Wood for prediction error criteria based smoothness selection. Simulation results suggest that the new REML and ML methods offer some improvement in mean-square error performance relative to GCV or Akaike's information criterion in most cases, without the small number of severe undersmoothing failures to which Akaike's information criterion and GCV are prone. This is achieved at the same computational cost as GCV or Akaike's information criterion. The new approach also eliminates the convergence failures of previous REML- or ML-based approaches for penalized GLMs and usually has lower computational cost than these alternatives. Example applications are presented in adaptive smoothing, scalar on function regression and generalized additive model selection.
Article
Long-distance migration can lower infection risk for animal populations by removing infected individuals during strenuous journeys, spatially separating susceptible age classes, or allowing migrants to periodically escape from contaminated habitats. Many seasonal migrations are changing due to human activities including climate change and habitat alteration. Moreover, for some migratory populations, sedentary behaviors are becoming more common as migrants abandon or shorten their journeys in response to supplemental feeding or warming temperatures. Exploring the consequences of reduced movement for host–parasite interactions is needed to predict future responses of animal pathogens to anthropogenic change. Monarch butterflies (Danaus plexippus) and their specialist protozoan parasite Ophryocystis elektroscirrha (OE) provide a model system for examining how long-distance migration affects infectious disease processes in a rapidly changing world. Annual monarch migration from eastern North America to Mexico is known to reduce protozoan infection prevalence, and more recent work suggests that monarchs that forego migration to breed year-round on non-native milkweeds in the southeastern and south central Unites States face extremely high risk of infection. Here, we examined the prevalence of OE infection from 2013 to 2016 in western North America, and compared monarchs exhibiting migratory behavior (overwintering annually along the California coast) with those that exhibit year-round breeding. Data from field collections and a joint citizen science program of Monarch Health and Monarch Alert showed that infection frequency was over nine times higher for monarchs sampled in gardens with year-round milkweed as compared to migratory monarchs sampled at overwintering sites. Results here underscore the importance of animal migrations for lowering infection risk and motivate future studies of pathogen transmission in migratory species affected by environmental change.