Scientists' warning on climate change and insects

Abstract

Climate warming is considered to be among the most serious of anthropogenic stresses to the environment, because it not only has direct effects on biodiversity, but it also exacerbates the harmful effects of other human‐mediated threats. The associated consequences are potentially severe, particularly in terms of threats to species preservation, as well as in the preservation of an array of ecosystem services provided by biodiversity. Among the most affected groups of animals are insects—central components of many ecosystems—for which climate change has pervasive effects from individuals to communities. In this contribution to the scientists' warning series, we summarize the effect of the gradual global surface temperature increase on insects, in terms of physiology, behavior, phenology, distribution, and species interactions, as well as the effect of increased frequency and duration of extreme events such as hot and cold spells, fires, droughts, and floods on these parameters. We warn that, if no action is taken to better understand and reduce the action of climate change on insects, we will drastically reduce our ability to build a sustainable future based on healthy, functional ecosystems. We discuss perspectives on relevant ways to conserve insects in the face of climate change, and we offer several key recommendations on management approaches that can be adopted, on policies that should be pursued, and on the involvement of the general public in the protection effort.

Full text

REVIEW
Scientists’warning on climate change and insects
Jeffrey A. Harvey
1,2
| Kévin Tougeron
3,4
| Rieta Gols
5
|
Robin Heinen
6
| Mariana Abarca
7
| Paul K. Abram
8
| Yves Basset
9,10
|
Matty Berg
2,11
| Carol Boggs
12,13
| Jacques Brodeur
14
|
Pedro Cardoso
15
| Jetske G. de Boer
1
| Geert R. De Snoo
1
|
Charl Deacon
16
| Jane E. Dell
17
| Nicolas Desneux
18
|
Michael E. Dillon
19
| Grant A. Duffy
20,21
| Lee A. Dyer
22
|
Jacintha Ellers
2
| Anahí Espíndola
23
| James Fordyce
24
|
Matthew L. Forister
22
| Caroline Fukushima
15
| Matthew J. G. Gage
25†
|
Carlos García-Robledo
26
| Claire Gely
27
| Mauro Gobbi
28
|
Caspar Hallmann
29
| Thierry Hance
3
| John Harte
30
|
Axel Hochkirch
31,32
| Christian Hof
6
| Ary A. Hoffmann
33
|
Joel G. Kingsolver
34
| Greg P. A. Lamarre
9,10
| William F. Laurance
27
|
Blas Lavandero
35
| Simon R. Leather
36†
| Philipp Lehmann
37,38
|
Cécile Le Lann
39
| Margarita M. L
opez-Uribe
40
| Chun-Sen Ma
41
|
Gang Ma
41
| Joffrey Moiroux
42
| Lucie Monticelli
18
| Chris Nice
43
|
Paul J. Ode
44,45
| Sylvain Pincebourde
46
| William J. Ripple
47
|
Melissah Rowe
48
| Michael J. Samways
16
| Arnaud Sentis
49
|
Alisha A. Shah
50
| Nigel Stork
51
| John S. Terblanche
16
|
Madhav P. Thakur
52
| Matthew B. Thomas
53
| Jason M. Tylianakis
54
|
Joan Van Baaren
39
| Martijn Van de Pol
48,55
| Wim H. Van der Putten
1
|
Hans Van Dyck
3
| Wilco C. E. P. Verberk
56
| David L. Wagner
26
|
Wolfgang W. Weisser
6
| William C. Wetzel
57
| H. Arthur Woods
58
|
Kris A. G. Wyckhuys
59,60
| Steven L. Chown
61
Correspondence
Jeffrey A. Harvey
Email: j.harvey@nioo.knaw.nl
Handling Editor: Jean-Philippe Lessard
Abstract
Climate warming is considered to be among the most serious of anthropogenic
stresses to the environment, because it not only has direct effects on biodiver-
sity, but it also exacerbates the harmful effects of other human-mediated
†
Deceased.
For affiliations refer to page 22
Received: 2 June 2022 Accepted: 19 July 2022
DOI: 10.1002/ecm.1553
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any
medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.
© 2022 The Authors. Ecological Monographs published by Wiley Periodicals LLC on behalf of The Ecological Society of America.
Ecological Monographs. 2023;93:e1553. https://onlinelibrary.wiley.com/r/ecm 1of37
https://doi.org/10.1002/ecm.1553

threats. The associated consequences are potentially severe, particularly in
terms of threats to species preservation, as well as in the preservation of an
array of ecosystem services provided by biodiversity. Among the most affected
groups of animals are insects—central components of many ecosystems—for
which climate change has pervasive effects from individuals to communities.
In this contribution to the scientists’warning series, we summarize the effect
of the gradual global surface temperature increase on insects, in terms of
physiology, behavior, phenology, distribution, and species interactions, as well
as the effect of increased frequency and duration of extreme events such as hot
and cold spells, fires, droughts, and floods on these parameters. We warn that,
if no action is taken to better understand and reduce the action of climate
change on insects, we will drastically reduce our ability to build a sustainable
future based on healthy, functional ecosystems. We discuss perspectives on
relevant ways to conserve insects in the face of climate change, and we offer
several key recommendations on management approaches that can be
adopted, on policies that should be pursued, and on the involvement of the
general public in the protection effort.
KEYWORDS
arthropods, conservation, ecology, evolution, extreme events, global warming, temperature
INTRODUCTION
Of the many trends that are worrying scientists across the
planet, the loss of biodiversity is among the most serious,
because it may lead to the breakdown of ecological
communities with concomitant, detrimental effects on
critical ecosystem services and functions (Steffen
et al., 2015). Across the biosphere, the most prominent
drivers of biodiversity change and decline are habitat
alteration, overexploitation, (agrochemical) pollution,
biological invasions, and anthropogenic climate change
(IPBES, 2019; Millennium Ecosystem Assessment, 2005;
Venter et al., 2016). The biosphere has already warmed
by 1.1C since industrialization and is projected to
warm a further two to five degrees by 2100 (Figure 1)
unless greenhouse gas emissions are significantly reduced
(IPCC, 2021). Climate change can lead not only to the
extinction of species, but also to profound changes in
their abundances, distributions, and species’assem-
blages, compositions, and interactions with other species
(Pecl et al., 2017; Schleuning et al., 2020; Sinervo
et al., 2010; Steinbauer et al., 2018). Moreover, it is
expected to act in either additive or synergistic ways with
other drivers to exacerbate impacts on biodiversity
(e.g., Boggs, 2016; Halsch et al., 2021; Hulme, 2017;
Raven & Wagner, 2021; Verberk, Durance, et al., 2016;
Verheyen & Stoks, 2019). A growing body of empirical
literature is showing that many populations of insects are
declining rapidly across many parts of the biosphere,
although patterns vary geographically and among differ-
ent taxa or functional groups (Biesmeijer et al., 2006;
Crossley et al., 2020; Didham, Barbero, et al., 2020;
Didham, Basset, et al., 2020; Hallmann et al., 2017;
Janzen & Hallwachs, 2021;S

anchez-Bayo &
Wyckhuys, 2019; van Klink et al., 2020; Wagner, 2020;
Wagner, Fox, et al., 2021; Warren et al., 2021). These
declines are considered to be of profound concern, with
terms like an emerging “insect apocalypse”being increas-
ingly used by the media and even some scientists to
describe this phenomenon (Goulson, 2019; Jarvis, 2018).
Observed trends in the demographics of many taxa—
including important functional groups like pollinators,
nutrient cyclers, and natural enemies, as well as in the
abundance of crop, forest, and urban pests—is currently
considered serious enough to merit profound concern
(Wagner, Fox, et al., 2021). Insects are important compo-
nents of biodiversity (García-Robledo, Kuprewicz,
et al., 2020; Stork, 2018; Wilson, 1987) contributing in
diverse and well-documented ways to aboveground and
belowground diversity, ecosystem functioning, and to var-
ious ecosystem services (Dangles & Casas, 2019).
Important ecosystem services provided by insects are polli-
nation, pest control, and nutrient recycling
(Schowalter, 2013; Schowalter et al., 2018). Insects and
their products also provide resources for higher trophic
level organisms, including humans (Ramos-Elorduy, 2009;
2of37 HARVEY ET AL.

Schowalter, 2013). However, some insect species nega-
tively affect human health and welfare by vectoring patho-
gens or by eating our crops (Schowalter et al., 2018). In
natural (unmanaged) ecosystems, abundances of pathogen
and vector species are controlled through various food
web interactions and habitat conditions, whereas anthro-
pogenic land use changes such as deforestation, habitat
fragmentation, and agricultural development can modify
these interactions with consequences for disease transmis-
sion (Burkett-Cadena & Vittor, 2018; Gottdenker et al.,
2014). In addition, it is predicted that global warming
will affect the length of the transmission season and
facilitate the expansion of the geographical range of the
disease (Woodward et al., 2014). How this will affect
spread and severity of vector-borne diseases is difficult to
predict as it depends on the complex interplay between
many factors, including socio-economic ones, which them-
selves can be affected by global warming (Caminade
et al., 2019; Rogers & Randolph, 2006). Pest incidence and
severity is predicted to increase under conditions of global
warming by, e.g., direct effects of higher temperature on
insect survival, development, and reproduction, and by
expansion of their geographical ranges, which is often
exacerbated by global trade and the introduction of exotic
pests (Lamichhane et al., 2015;Skend
ˇ
zi
cetal.,2021).
Given their generally small body size, and the fact
that the vast majority of species are ectothermic
(Harrison et al., 2012), insects are considered to be espe-
cially susceptible to the direct effects of changing temper-
ature and moisture regimes (Halsch et al., 2021; Harvey,
Heinen, Gols, & Thakur, 2020; Wagner, 2020). Climate
change can, therefore, shape the physiology and behavior
of insects, with concomitant effects on life-cycles,
life-history traits, reproduction, and population persis-
tence (García-Robledo et al., 2016; Wagner, 2020). For
example, the temperature-size rule predicts that insect
size is to some degree plastic, and under warmer condi-
tions, ectothermic species develop faster but become
smaller in body size (Atkinson, 1994; Verberk
et al., 2021). However, there are many exceptions to the
temperature-size rule among insect groups or populations
(Horne et al., 2015). In a phylogenetically controlled
analysis of temperature-size relationships in tropical
insects, the main conclusion is that size differences
among populations are heritable rather than the result of
body size plasticity, and global warming will not inevita-
bly lead to body size decreases (Duffy et al., 2015;
García-Robledo, Baer, et al., 2020). Nevertheless, signifi-
cant direct impacts of climate change on insect
populations are to be expected for many species and
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Forecast change: current (annual means 1986–2005) ► end of century (annual means 2081–2100)
[Access 1.0 RCP8.5 CMIP5 model]
FIGURE 1 Changes to the abiotic environment relevant to insect life history. Each panel is the difference between current and
2100 conditions (T=temperature). Extreme indices are based on those suggested by the COE for climate extremes (https://www.climdex.org).
ECOLOGICAL MONOGRAPHS 3of37

taxonomic groups. However, given the great diversity of
insect life histories, behavioral and ecophysiological
adaptations, habitats and environments globally, there
will inevitably be some exceptions to the generally
negative impacts of excessively high temperatures
(e.g., García-Robledo, Baer, et al., 2020)orincreasesin
growing season length in typically cold environments
(Sinclair, 2014). For instance, warming is enabling, at least
transiently, some species of thermophilic forest and agricul-
tural pests and disease vectors such as mosquitoes to
expand their ranges to higher latitudes (Battisti &
Larsson, 2015;Hilletal.,2011;Jacteletal.,2019; Kovats
et al., 2001;Skend
ˇ
zi
cetal.,2021). The economic costs of
these climate-mediated range expansions on food produc-
tion and human health could be enormous. Responses of
insects and different functional groups to climate change
are thus taking a wide variety of forms in different parts of
the globe, including in both natural and human-dominated
environments such as forests, wetlands, agricultural land-
scapes, and urban environments.
A recent study argues that the combined effects of
climate change and agricultural intensification are
negatively affecting insect biodiversity (Outhwaite
et al., 2022). In this synthesis “warning paper,”we
explore insect responses to climate change and climatic
extremes, what is known about them, what knowledge is
still needed to reduce uncertainty, and what key recom-
mendations scientists can formulate for policy makers
and the general public to reduce the harmful impacts.
We address the effects of gradual changes in climate and
increased climatic variability i.e., frequency of extreme,
abrupt, and punctuated events (Figure 2), and how they
are possibly modulated by other global change drivers.
These changes cannot be ignored, as they are already
having major consequences on insects and will have
implications for insect conservation and management in
the coming decades (Cardoso et al., 2020). Although
much information is available on the ecophysiology of
individuals and the survival of different insect species, it
is necessary to keep in mind that our warning is about
the impact this will have on humankind. This impact will
be mediated by the wider effects of climate change for
the disruption of interaction networks within ecosystems
(Tylianakis & Morris, 2017), and the ecosystem services
range
shifts phenology species
interactions
fitness pathogens foodsource fertility
concomitant and/or
sequential
extreme events decline
/extinction
persistence ecosystem
services
reduction of
stressors
GRADUAL
LONG-TERM
EXTREME EVENTS
INTERVENTION
temperature
extremes drought
extreme
rainfall fire
warming seasonality
management public
participation
CHANGE IMPACT
FIGURE 2 Climate change impacts on insects can be categorized into two major categories: Gradual long-term change and extreme
events that will increase in frequency and severity, while interventions include formal mitigation of change through policy and public
approaches which in turn help to reduce impacts in various ways.
4of37 HARVEY ET AL.

and functions provided by insects. Unfortunately, along
with climate change denial, other incipient forms of sci-
entific denial are becoming more prevalent in recent
years, including extinction denial (Lees et al., 2020).
However, if we fail to recognize the importance of insects
and their vitally important role in the functioning of nat-
ural and managed terrestrial and freshwater ecosystems
(Losey & Vaughan, 2006), or the impacts that climate
change and other anthropogenic stressors are having on
them, then we are essentially limiting our ability to act.
Recognizing and acting upon the clear and striking evi-
dence of climate change effects on insects is critical to
support our collective efforts to build an ecologically sus-
tainable future.
EFFECTS OF LONG-TERM,
ANTHROPOGENIC CLIMATE
CHANGE ON INSECTS
The effect of long term, gradual warming on insect
declines is not always immediately apparent. Discernible
changes in insect population dynamics, their distribu-
tions, phenology, or abundance are usually detected after
critical performance thresholds affecting fecundity, sur-
vival, and other vital rates are surpassed (Harvey,
Heinen, Gols, & Thakur, 2020). Furthermore, warming is
occurring unequally across the biosphere and across
time-scales, with temperate, boreal, and polar ecosystems
at higher latitudes warming at much faster rates than
subtropical and tropical ecosystems at lower latitudes
(Parmesan, 2007; Post et al., 2018). Temperatures along
tropical mountains are increasing 0.017C per year, and
ecosystems are already shifting upslope (Feeley
et al., 2013). Winter is warming faster than any other sea-
son (IPCC, 2014), and nights are becoming warmer
worldwide (Karl et al., 1991).
In ectothermic organisms, warmer temperatures are
generally associated with increased metabolic rates that
may lead to increased physiological costs (Irwin &
Lee, 2003; Williams et al., 2012). While warming stimu-
lates growth, development, and reproduction up to some
optimal temperature, beyond this temperature, metabolic
costs increase disproportionately and thermal injuries
accumulate, eventually leading to developmental failure,
reduced fecundity, impaired dispersal capacity, and, ulti-
mately, decreased fitness and increased mortality
(Gilbert & Raworth, 1996). Exposure to elevated tempera-
tures affects insect growth and development, often
resulting in body size reductions, with negative effects on
fecundity, longevity and dispersal, all of which can reduce
their resilience in the face of climate change and in
the worst-case scenarios lead to population crashes
(Abram et al., 2017; Gardner et al., 2011; Hof et al., 2011;
Sweeney et al., 2018).Asaresult,climatechangemay
reduce genetic diversity through processes including popu-
lation bottlenecks, loss of genetic diversity that is not
related to selection, and outbreeding or inbreeding depres-
sion (Halsch et al., 2021; Pauls et al., 2013), which
threatens the persistence of populations. Insect populations
may display very limited genetic variation in traits associ-
ated with thermal tolerance and it is concerning that in
such populations, local adaptation is already lagging
behind current temperatures (García-Robledo & Baer,
2021a,2021b). Studies with ants show that thermal toler-
ance (or intolerance) varies across different species and
under different thermal niche conditions, with tolerance or
even benefits of marginally higher temperatures on colony
fitness offset beyond critical thermal thresholds (Diamond
et al., 2013). This paper primarily focuses on ecological
responses to climate change, with less attention paid to
physiological responses at the individual (=organism) level.
One of the most notable phenotypic responses to warming
seen in some insects are changes in body color and, in
particular, a reduction in melanization (Brakefield &
de Jong, 2011; Clusella-Trulas & Nielsen, 2020; Kingsolver
et al., 2011;Roulin,2014). Reduced melanization is poten-
tially adaptive, as lighter individuals may be able to more
effectively thermoregulate when exposed to extreme heat
than darker individuals (Roulin, 2014). Consequently,
phenotypic plasticity can to some extent counter other
physiological stresses induced by climate change.
We argue that the most important outcome of climate
warming is that insect communities become destabilized
(Diamond et al., 2016;Pelinietal.,2014; Pureswaran
et al., 2021), and that populations and even entire species
may go extinct unless they alter their geographical distri-
butions and/or adjust their spatial and temporal behav-
ioral activity patterns and seasonal phenologies to the new
climatic conditions (García-Robledo et al., 2016;Halsch
et al., 2021; Harvey, Heinen, Gols, & Thakur, 2020). This
leads to changes in the structure of ecological communi-
ties at given locations in ways that affect species interac-
tions, with potentially severe repercussions on ecosystem
stability and functioning, and subsequently the provision-
ing of ecosystem services.
Effect on species distributions
Species’distributions are strongly determined by climatic
factors and are labile, expanding or contracting during
long-term climatic fluctuations (Hewitt, 2000). Similarly,
the rapid rate of current climate warming strongly deter-
mines how successfully insects are able to track climatic
shifts, since they may or may not keep pace with these
changes. Projections of how global warming will affect
species geographic ranges are based on bioclimatic
ECOLOGICAL MONOGRAPHS 5of37

envelope models (Vieilledent et al., 2016), the response of
insects to extreme temperatures (Sunday et al., 2014), or
simple graphical models of species elevational ranges
(Colwell et al., 2008), which calculate the potential loss of
suitable habitats. The general consensus among these dif-
ferent approaches is that species at lowest latitudes or ele-
vations must migrate to cooler environments to avoid
extinction (Colwell et al., 2008; Sunday et al., 2011,2014)
(Figure 3a,b). Moreover, the ability of insects to track
shifting thermoclines will be affected by various aspects
of their eco-evolutionary dynamics (Miller et al., 2020;
Wellenreuther et al., 2022). Predicting this to inform
management strategies will increasingly require the use
of modeling and genomic sequencing (Wellenreuther
et al., 2022).
Warren et al. (2018) generated bioclimatic models
predicting the effects of slight, moderate and extreme
warming on geographic range losses incurred by 34,000
insect species across the biosphere. They found an expo-
nential effect of temperature on range losses by the year
2100. With an increase of 3.2C, the ranges of almost half
of the insect species will contract by 50% or more, whereas
FIGURE 3 (a) Many insects are showing a range of ecophysiological responses to longer-term climatic changes. For example, the
emperor dragonfly (Anax imperator) has shifted its distribution northward and to higher elevations in Europe since 2000 in response to
warming (Platts et al., 2019). (b) In California and Mexico, the Quino Checkerspot butterfly (Euphydryas editha quino) has responded to
recent warming by moving to higher elevations, and by shifting from its preferred lowland food plant (a Plantago species) to Collinsia
concolor, which is more abundant at higher elevations. Increased warming, however, still threatens this endangered subspecies (Parmesan
et al., 2015). (c) Many recent insect declines, such as the now vulnerable yellow-banded bumblebee (Bombus terricola), have been attributed
to climate extremes, and especially hotter maximum temperatures during the summer (Martinet et al., 2015). (d) Exposure to heat waves can
have significant effects on insect reproduction. Functional responses in the facultative hyperparasitoid, Gelis agilis, are strongly correlated
with ambient temperature, and at high temperatures their ability to exploit hosts is greatly impaired (Chen, Gols, et al., 2019). Photograph of
emperor dragonfly by Tim Bekaert; photograph of Quino Checkerspot butterfly by Andrew Fisher (USFWS volunteer biologist); photograph
of yellow-banded bumblebee by rob Foster (https://www.inaturalist.org/users/264273); photograph of Gelis agilis by Tibor Bukovinszky
(NVWA Wageningen University & Research. Wageningen).
6of37 HARVEY ET AL.

this drops to 18% of insects at +2C, and 6% of insects at
+1.5C (Warren et al., 2018). Insects, like other organisms,
are responding to warming by shifting their distributions
poleward and to higher elevations (Grewe et al., 2013;
Heiser & Schmitt, 2013; Parmesan & Yohe, 2003).
However, range retractions at lower latitudes or altitudes
have received less attention (but see Kerr et al., 2015;
Merrill et al., 2008), as the disappearance of a population
is a more gradual process. Moreover, as climate suitability
continues to decline in these so-called trailing edge
populations, more and more insect species, such as butter-
flies, will accumulate an extinction debt (Devictor et al.,
2012;Thomasetal.,2004).
Despite the fact that insect extinction is a demographic
process, only two studies have determined the effect of
global warming on insect population dynamics, range
shifts, and fitness (Deutsch et al., 2008; García-Robledo &
Baer, 2021b). Deutsch et al. (2008) combined life table
analysis with global temperature records to estimate
changes in fitness at current and future temperatures. One
limitation of this study is that it only includes fitness esti-
mates for pantropical crop pests and tropical biocontrol
agents reared in the laboratory, which are usually tolerant
to high temperatures or adapted to laboratory conditions.
Nevertheless, their results support the hypothesis that
tropical insect species are at higher risk of extinction than
temperate insects (Deutsch et al., 2008). For example, the
leaf beetle Cephaloleia belti includes high- and
low-elevation mitochondrial haplotypes locally adapted to
cold and warm temperatures. Both haplotypes interbreed
in a hybridization zone at mid elevation. Demographic
models, combined with long-term temperature datasets,
show that, with an increase of just 2C, trailing-edge
populations will decline, and middle elevations will
become refuges to global warming (García-Robledo &
Baer, 2021a,2021b).
As long as population losses at lower latitudes or ele-
vations are compensated by range expansions at other
boundaries of the range, most species may be able to per-
sist (Deutsch et al., 2008). Range shifts most likely will be
accompanied by loss of genetic diversity; however, not all
species are able to perform such range shifts, as their
capability to do so depends on several factors, including
active or passive dispersal abilities, population dynamics,
genetic diversity, width of the thermal safety margins,
habitat availability, intra- and interspecific interactions,
and co-shifting of food sources (Amundrud &
Srivastava, 2020; Berg et al., 2010; Bybee et al., 2016;
Deutsch et al., 2008; Hof et al., 2011; Menzel &
Feldmeyer, 2021). Thus, different times of response to cli-
mate change in insects are evident, and the existence of a
time-lag in species response implies caution in predicting
species’occurrence shifts following climate change. For
instance, Rom
an-Palacios and Wiens (2020) found that
“hot”years coincided with increased rates of local extinc-
tion, and suggested that more than half of insect species
may not be able to adjust their distributions rapidly
enough to avoid extinction. However, Sunday et al.
(2012) reported that terrestrial ectotherms at the warmest
(or “trailing”) edge of their ranges, including insects, do
not appear to be delimited by insufficient heat tolerance,
suggesting that factors other than temperature
(e.g., drought, competition, light) may shape their
range boundaries and thus, warming may not result in
consistent shifts in these trailing edges (Spence &
Tingley, 2020). Studies with damselflies show plasticity to
warming at range margins and that gene switching for
thermal tolerance appears to be under strong selection
(Dudaniec et al., 2018; Lancaster et al., 2015), suggesting
that some species can thus adapt rapidly to shifting ther-
moclines. However, historical and ongoing habitat loss
and fragmentation and isolation as a consequence of
human land use changes is a major current threat to
insects that exacerbates climate change impacts by ham-
pering range expansions, especially across uneven land-
scapes with elevational gradients (Samways et al., 2020;
Yadav et al., 2018,2021). It is also important to acknowl-
edge that range shifts in response to warming may lead
to the elimination of native competitors in the new range
where there is strong niche overlap (or vice-versa). This
area is in urgent need of further investigation.
The cryosphere (e.g., glaciers, permafrost) covers
10% of the Earth’s surface but is declining as a result of
long-term warming trends (Pörtner et al., 2019; Zemp
et al., 2019). Glacial retreat is indeed an iconic symbol of
climate warming, and leaves habitats that are rapidly
colonized by different biotic communities. Receding ice
threatens many insects that are restricted to narrow habi-
tat zones in front of glaciers (Gobbi et al., 2021). Studies
report that cold-adapted and hygrophilous insects are
moving to higher elevations in response to warming but
with a reduction in their broader spatial distributions
(Moret et al., 2016; Valle et al., 2020). Altitudinal shifts
are not only triggered by the thermal requirements of
cold-adapted species, but also by the increasing competi-
tion of warm-tolerant species coming from lower eleva-
tions, and this covers a wide range of taxa in both
terrestrial and aquatic systems (Brighenti et al., 2021;
Cauvy-Fraunié & Dangles, 2019; Ficetola et al., 2021;
Pizzolotto et al., 2014). Climate warming in glaciated
alpine areas produces both “winners”and “losers”
(Cauvy-Fraunié & Dangles, 2019). The likely reason is
that the response of insects to increasing temperature
varies among species, communities and geographic
area (Ficetola et al., 2021). Winners tended to be general-
ist/invasive species, good dispersers, generally colonizing
ECOLOGICAL MONOGRAPHS 7of37

from downstream or downslope, such as grasshoppers
(Yadav et al., 2018); conversely, the losers are often spe-
cialist species, adapted to cold habitats, among which
some were restricted to isolated glacier-influenced ecosys-
tems (Cauvy-Fraunié & Dangles, 2019). The spatial and
temporal dynamics of the insect species assemblages
linked to ice-related landforms is revealing a rapid turn-
over of species with the substitution of cold-adapted spe-
cies with more eurythermal species (Gobbi et al., 2021).
It is important to temper the primarily negative
effects of warming on insects by also stressing that some
species—at least transiently—may benefit in response to
milder winters and warmer conditions that enhance sur-
vival or voltinism (Marshall et al., 2020; Musolin, 2007;
Tougeron et al., 2017). Winter is considered a major con-
trol agent of forest and crop pests and trends toward
warmer winters or reduced numbers of deep frosts are
leading to increased numbers of pest outbreaks
(Cannon, 1998; Pureswaran et al., 2019). Moreover, since
1970 more species of generalist moths in Great Britain
have increased in abundance than decreased in abun-
dance (38% vs. 31%), presumably in response to warming
(Wagner, Fox, et al., 2021). Moreover, caterpillar abun-
dances in cloud forests in Ecuador increased between
2001 and 2019, because of an increase in sunny days as a
result of climate change (Wagner, Grames, et al., 2021). It
needs to be stressed that positive responses to warming
may be transient, especially as climatic extremes are
increasing, which imposes immense short-term stresses
on insect populations (Harvey, Heinen, Gols, &
Thakur, 2020), or else if warming continues unabated,
pushing many species beyond their thermal optima for
reproduction and survival (Buckley & Kingsolver, 2021).
Effects of warming may also be disproportionate up the
food chain, with more deleterious physiological effects on
predators or parasitoids than on their prey or hosts
(Jeffs & Lewis, 2013; Tougeron et al., 2019). Under this
scenario, herbivores may benefit from enemy release
(Moore et al., 2021). The net effect will still likely have
negative consequences for food webs and communities,
as the loss of top-down control might lead to all kinds of
unpredictable, potentially destabilizing, effects over time.
Effect on phenology
In addition to range shifts, another well-documented aspect
of insect response to climate change is a change in phenol-
ogy. Elevated temperatures increase the duration of the
growing season, which together with faster developmental
rates allows some insect species to increase the number of
generations in a given year (i.e., voltinism) (Bradshaw &
Holzapfel, 2001). Even apparently modest temperature
rises can result in increases in seasonal or annual genera-
tions (Parmesan & Yohe, 2003). For example, Altermatt
(2010)showedthat44butterflyandmothspeciesincreased
the number of generations after 1980 in response to rising
temperatures. While warmer temperatures generally allow
for extra generations in the summer, a higher voltinism
may result in smaller adults being produced (Horne
et al., 2015; Verberk et al., 2021). Moreover, the addition of
extra generations later in the season is also affected by the
overwintering strategy and photoperiod (i.e., shortening
day length and the initiation of diapause), which is a domi-
nant seasonality cue for many temperate insects (Lindestad
et al., 2019; Marshall et al., 2020;Tougeron,2019). Both
parameters are therefore important to consider when
projecting effects of climate change, and extreme tempera-
tures, on patterns in insect voltinism and body size (Bale &
Hayward, 2010; Forrest, 2016; Verberk et al., 2021).
Ecological consequences of changes in overwintering
strategies, and more generally in activity timing, are still
far from being well-understood (Williams et al., 2015). In
some cases, an extended period of warm temperatures that
promotes development may delay winter diapause induc-
tion. This may create a developmental trap resulting in the
production of a complete or partial additional generation
in the autumn that cannot survive or enter diapause (“the
lost generation hypothesis”(van Dyck et al., 2015; Kerr
et al., 2020). In addition, diapause maintenance and termi-
nation are disrupted because of warm winters. For exam-
ple, many insects, like other organisms, require a period of
chilling during diapause before they can resume develop-
ment (Lehmann et al., 2017; Stålhandske et al., 2017). If a
chilling cue is not received, individuals may emerge later
or even not at all (Bale & Hayward, 2010;Tougeron
et al., 2019). In line with this, forest experiments
conducted by Fitzgerald et al. (2021) in North Carolina
support the “cool-season sensitivity”hypothesis, showing
that arthropods adapted to cooler conditions endured
stronger negative effects of warming during warm winters.
Alternatively, MacLean et al. (2017) found that exposure
to warmer conditions in winter actually benefitted acorn
ants by “priming”them metabolically for early activity in
spring. Therefore, the effects of warmer winters on insects
is likely to vary considerably among different taxa.
Exposure to repeated heat waves in summer, or
warmer spells during winter (or both), may also generate
inappropriate cues that lead insects into developmental
traps (e.g., resuming development in the middle of winter)
(Boggs, 2016; Forrest, 2016). For example, unseasonably
warm autumn conditions are causing the wall brown but-
terfly Lasiommata megera, to alter life-cycle decisions by
breaking diapause at the end of the second generation.
This makes the third generation highly susceptible to win-
ter mortality, and as a result the butterfly is declining
8of37 HARVEY ET AL.

rapidly across much of western Europe (van Dyck
et al., 2015). In other cases, a partial or complete loss of
winter diapause may result from successive years with per-
missive winter temperatures, which have been shown to
result in the activity of some aphid parasitoids across the
entire year (Tougeron et al., 2017). Therefore, individuals
that remain active throughout the winter rely on physiolog-
ical and behavioral thermotolerance to survive, including
rapid cold hardening, or the induction of transient and
easily reversible resting stages (Diniz et al., 2017). Warmer
winter conditions have also been shown to affect colony
phenology in honeybees, leading to mismatches with their
floral resources (Nürnberger et al., 2019). Furthermore,
warmer periods in winter also stimulate colony brood
rearing activity and this benefits their main parasite, inva-
sive Varroa destructor mites (Nürnberger et al., 2019;
Vercelli et al., 2021). Seasonal changes in temperature
therefore also need to be placed in the context of facilitat-
ing pathogens or parasites.
Winter conditions tend to be much more variable geo-
graphically and locally than summer conditions
(Bonan, 2004) and this can affect latitudinal variation in
biotic processes much more strongly than conditions dur-
ing the growing season. Trends toward decreasing winter
snow depth, increasing numbers of winter thaws, or later
snowfalls and earlier winter melts are also being observed
in many temperate parts of the world under climate
warming (Fontrodona Bach et al., 2018; McCabe &
Wolock, 2010). The “subnivium,”defined as the physical
interface between the snowpack and ground, is an impor-
tant refuge that protects overwintering insect species
from extremely cold temperatures by providing insula-
tion during diapause (Zhu et al., 2019). During periodic
thaws, or when spring melt occurs early, insects are
exposed to cold temperatures on bare soils that may be
lethal (Bale & Hayward, 2010; Williams et al., 2015). The
absence of snow cover also subjects diapausing stages to
elevated rates of moisture loss and, as a consequence,
desiccation. Loss of snow cover also alters insect metabo-
lism leading to a potential phenological mismatches with
key resources (Renner & Zohner, 2018). Moreover, dia-
pausing insects are more exposed to winter-active preda-
tors when snow cover disappears for even short periods
(Cooper, 2014). However, the longer-term effects of sea-
sonal changes in temperature and precipitation on insect
demographics are thus far little-studied.
Effect on species interactions
Insects, like most organisms, are embedded in complex
communities, and their fate depends on interactions with
other species. In general, predation and other multitrophic
interactions often result from long co-evolutionary
processes that are specific to a particular environment
under relatively stable climatic conditions. However, tem-
perature changes may differentially affect the biology of
each of the component species of a system. This has been
demonstrated in multitrophic systems involving plants,
their herbivores, natural enemies (parasitoids, predators
and pathogens), and hyperparasitoids (Agosta et al., 2017;
Bale et al., 2002; Bannerman et al., 2011; Moore
et al., 2021;Pardikesetal.,2022; Romo & Tylianakis, 2013;
Schreven et al., 2017;Tougeronetal.,2018). Studies show
that the effects on survival, development, and reproduction
are generally negative, e.g., plants growing under condi-
tions of extreme temperatures, increased CO
2
,and/or
reduced rainfall could become suboptimal nutritional
resources for herbivores, indirectly influencing natural
enemy fitness and associated biocontrol services (Han
et al., 2019,2022).Adoublenegativeeffectmayoccur
whentemperatureincreasepromotesherbivorouspest
populations while negatively impacting host plant defenses
(Wang et al., 2021). Temperature increases may also induce
slight shifts in feeding preference; e.g., with phytophagous
insects feeding more on native plants versus invasive plants
when temperature increases (Liu et al., 2021). Interactions
between plants and insect pollinators are typically also
known to be weakened by climate change (Tylianakis
et al., 2008).Theeffectsarelikelytobeevenmoreimpor-
tant in higher trophic levels that depend on the host speci-
ficity and the capacity of the lower trophic levels to adapt
to these changes and also because they are less numerous
(Monticelli et al., 2022; Thackeray et al., 2016; van Baaren
et al., 2010). These effects may be exacerbated in
species-poor communities e.g., on islands or in increasingly
homogeneous, chemically disrupted agro-landscapes
(Raven & Wagner, 2021). Moreover, many vertebrates
(e.g., small mammals, birds, reptiles, amphibians) depend
on insects as food, and the loss of insects in a warming
world is therefore likely to have enormous effects on
broader ecological communities.
One major consequence of changes in the distribution
and phenology of insects is the occurrence of potential
spatial and temporal mismatches among interacting spe-
cies. Differential responses between trophic levels may,
for instance, decouple the timing of generations among
interacting species (Damien & Tougeron, 2019; Gérard
et al., 2020; Thackeray et al., 2016). This may be espe-
cially prevalent among specialized species that are
constrained by the responses of their prey or hosts, or in
species that already have narrow environmental niches
(Damien & Tougeron, 2019; Tylianakis et al., 2008). The
mismatch between insect food availability in spring and
the breeding success of long-distance migratory birds is a
well-studied example (Both et al., 2006). These mis-
matches may subsequently destabilize and reduce the
efficacy of crucial interactions such as between plants
ECOLOGICAL MONOGRAPHS 9of37

and pollinators (Peralta et al., 2020), or herbivores and
natural enemies (Bale et al., 2002; Harvey, 2015; Singer &
Parmesan, 2010). Mismatches may be countered by
strong selective pressure leading to readjustments in the
phenology, distribution, or physiology of interacting spe-
cies (Klapwijk et al., 2010; Schleuning et al., 2020).
Moreover, climate warming may lead to formation of
new interactions among species that were previously
disassociated in space or time (Shah et al., 2020). Novel
interactions may arise as a result of differences among
species responses and thus adaptation to warming
(including their value as an interaction partner,
e.g., nutritional quality), because of differential responses
of spatial or temporal range, or when existing interaction
partners go extinct and switches to new partners are
required (Tylianakis & Morris, 2017).
Thermal tolerance mismatches may occur intraspe-
cifically via sexual conflicts in response to different tem-
peratures, as has been demonstrated in damselflies
(Svensson et al., 2020) and whiteflies (Ma et al., 2014),
or interspecifically, where tightly interacting species,
such as parasitoids and their hosts, exhibit differing sen-
sitivities to high temperatures (Abarca & Spahn, 2021;
Furlong & Zalucki, 2017; Wetherington et al., 2017).
This has been shown with thermal response curves for
pathogen growth and host defense (Thomas &
Blanford, 2003), and may lead to distinctly non-linear
responses to even small temperature changes. These
“thermal mismatch”effects could possibly result in a
destabilization of the dynamics that could lead to the
extinction of part of the system, and could create an
imbalance in how mass and energy transfer in food webs
(Thakur, 2020a). Studies using simulated warming to
mimic natural conditions have shown that exposure to
higher temperatures may also decrease the interaction
strength and the fitness of each of the interacting species
(Sentis et al., 2013). However, in some instances, temper-
ature can shape insect communities directly through dif-
ferences in species’thermal performance capabilities,
and not involve any effect of biotic interactions (Thierry
et al., 2021).
Warmer temperatures lead to higher metabolic and
feeding rates, which tends to increase the strength of tro-
phic interactions in the short term by making consumers
more dependent on their resources (Rall et al., 2012;
Sentis et al., 2012). However, energetic losses often
increase faster at higher temperatures than feeding rate,
leading to metabolic mismatches (Bideault et al., 2021).
This is especially problematic at very high temperatures,
where feeding rates either asymptote or even tend to
decrease (Sentis et al., 2012). This, in turn, influences bio-
mass distribution in food chains, such as in terrestrial
ecosystems where warming favored herbivore-heavy
webs (de Sassi & Tylianakis, 2012), or in some aquatic
systems where it favored top-heavy (predator dominated)
food chains (Kratina et al., 2012;O’Connor et al., 2009).
By modeling empirical data of thermal dependence on
key biological rates, Bideault et al. (2021) predicted that
warming is expected to favor top-heavy webs in both ter-
restrial and aquatic ecosystems. This highlights how tem-
perature can have cascading effects from physiology to
species interactions and community properties. In addi-
tion, the indirect effects of high temperatures on insect
communities may be mediated by changes to food web
structure and stability in communities where invasive
organisms are considered problematic. For example,
Sentis et al. (2021) showed that experimental communi-
ties with less connectivity, shortened food chains and
reduced temporal variability were more susceptible to
invasions under warmer conditions. Moreover, the
authors found that, under warmer conditions, in invaded
communities, species diversity decreased, network con-
nectivity increased and most top predators were lost lead-
ing to their replacement by meso-predators. Such
changes in insect communities can have detrimental con-
sequences for food web stability. This loss of predators
and herbivore regulation can be further exacerbated by
drought (Romo & Tylianakis, 2013). Such changes in
insect communities can have detrimental consequences
for food web stability.
Insects live in close association with microbial com-
munities residing within cells, on the cuticle, inside the
gastrointestinal tract or in the hemolymph. These
microbiomes comprise mutualists that for instance facili-
tate digestion, increase immunity or aid in detoxification
of plant metabolites (Feldhaar, 2011). Microbes may also
host antagonists (e.g., pathogens) that are detrimental to
insect health (Degli Esposti & Martinez Romero, 2017;
Gurung et al., 2019). It is likely that climate-driven
impacts on insect immunity will alter interactions
between hosts, symbionts and pathogens. For instance,
insects exposed to higher and more variable temperature
regimes show higher levels of immunity than those
exposed to lower and less variable temperature regimes
(Bozinovic et al., 2013; Catal
an et al., 2012; Van Dievel
et al., 2017), whereas other studies report opposite find-
ings (Karl et al., 2011). Mutualistic bacteria provide vari-
ous ecological benefits, such as resistance to
environmental stress, pathogen infections or natural ene-
mies, yet their impact remains poorly understood at the
level of ecological communities (Ferrari & Vavre, 2011;
Oliver et al., 2010). Due to the reduction in size of their
genome during coevolution with their hosts, mutualistic
endosymbionts have little potential to cope with a chang-
ing environment, including heat stress, and can therefore
be considered the Achilles heel of insects in a context of
10 of 37 HARVEY ET AL.

increasing temperature (Renoz et al., 2019). Some evidence
demonstrates that the interplay between insect hosts and
their mutualistic and antagonistic microbial partners is
temperature-dependent. For instance, gut microbiome
composition can shift across seasons and is concurrent
with changes in cold tolerance (Ferguson et al., 2018).
Benefits or drawbacks provided by symbionts can be
altered by increasing temperatures (Bensadia et al., 2006;
Higashi et al., 2020; Tougeron & Iltis, 2022), in part
because some symbionts reside in highly heat-sensitive
organs (e.g., bacteriocytes, Shan et al., 2017) and reversely,
alteration in the gut or intracellular microbial community
can influence insect resistance to temperature variation
(Henry & Colinet, 2018; Iltis, Tougeron, et al., 2021;
Jaramillo & Castañeda, 2021). Moreover, the microbiome
can be totally restructured during diapause in arthropods
(Mushegian et al., 2018; Mushegian & Tougeron, 2019),
but how modifications in seasonal strategies following cli-
mate change actually affect microbial communities merit
further investigations.
Aquatic insects face substantially different ecophysiolog-
ical problems compared with terrestrial insects, and likely
will experience climate change in fundamentally different
ways. Like terrestrial insects, aquatic insects are threatened
by rising temperatures and changes in patterns of tempera-
ture variation, among other factors (Birrell et al., 2020). The
underlying challenges, however, differ in part because water
has much higher heat capacity than air, such that—at least
in lotic habitats—local thermal variation is restricted in
space and time relative to atmospheric fluctuations, and
individuals have limited opportunities for behavioral ther-
moregulation. Consequently, aquatic insects have likely
evolved narrower thermal performance curves (Birrell
et al., 2020;Sundayetal.,2012).
Water also has a much lower oxygen capacity than
air, which magnifies the difficulties of supporting aerobic
respiration. Indeed, the oxygen problem may underlie
low observed heat tolerances of aquatic insects: in
warming waters, oxygen availability decreases modestly
but demand (from metabolism) often rises rapidly, and
oxygen shortage may lower heat tolerance (Frakes
et al., 2021; Verberk et al., 2011; Verberk, Overgaard,
et al., 2016). Consequently, aquatic ectotherms, including
aquatic insects may be more sensitive to rising mean tem-
peratures and more prone to reaching upper critical
limits (Rubalcaba et al., 2020; Verberk, Durance,
et al., 2016). Indeed, aquatic insects generally have lower
tolerance limits to heat than terrestrial insects (Chown
et al., 2015). These observations, and several recent
papers, also underscore the importance of flow in deliver-
ing oxygen to the body surface (Birrell et al., 2020; Frakes
et al., 2021). Low flows stemming from changes in hydro-
logical cycles or seasonality may compound the
challenges of higher temperatures and low oxygen.
Interestingly, a similar high temperature-hypoxia rela-
tionship certainly applies also to particular terrestrial
insects such as leaf gallers or cambium miners
(Pincebourde & Casas, 2016). Many aquatic insects spend
part of their lives under water as eggs, larvae, or pupae
before emerging as winged, terrestrial adults. These early
life stages are considered more vulnerable to heat par-
tially because of oxygen limitation inherent to living in
water (e.g., Verberk & Bilton, 2015), and also their lim-
ited mobility prevents them from behaviorally exploiting
gradients in temperature and oxygen availability (eggs
and pupae (Kingsolver et al., 2011). Thus, the response of
aquatic juveniles to rising temperatures will strongly
affect the presence and persistence of terrestrial adults.
These physical and physiological considerations may
have important consequences for how climate change
affects aquatic communities. Aquatic insects in temperate
mountains, for example, experience wider seasonal fluc-
tuations in temperature than do related species in tropi-
cal mountains at similar elevations. Temperate species
have wider thermal breadths and, in some cases, a
greater ability to acclimate to warmer temperatures
(Shah et al., 2017). Such patterns have been seen in other
aquatic taxa such as tadpoles (Gutiérrez-Pesquera
et al., 2016) and marine ectotherms (Sunday et al., 2011).
These studies suggest that tropical aquatic insects are
especially vulnerable to rising temperatures (Tewksbury
et al., 2008).
EFFECTS OF SHORT-TERM,
UNPREDICTABLE CLIMATE
EXTREMES ON INSECTS
No universal definition of extreme climatic events thus
far exists (Van de Pol et al., 2017) reviewed definitions
used in ecology). The term “climatic extremes”has been
used to describe meteorological phenomena (e.g., heat
waves, droughts, extreme rainfall events) as well as their
consequential physical impacts (e.g., flooding, fires, desic-
cation, tornadoes, hurricanes). Furthermore, extreme cli-
matic events can be defined by their climatological
extremeness or by the extremeness of the biological
impacts caused by a climate-related event. Finally, there
is little consensus on the threshold for extremes (1%, 5%
or 10% frequency or a certain percentile of a reference
period) and how other attributes should be factored in,
such as the temporal duration, magnitude, timing, spatial
scale of events, or the biological level of organization it
affects (Meehl & Tebaldi, 2004; Russo et al., 2014). This
lack of consensus is not surprising because extremes are
simply events in the tails of statistical distributions,
ECOLOGICAL MONOGRAPHS 11 of 37

which for both climate factors and ecological responses
are generally unimodal, making any definition of an
extreme climatic event an arbitrary cutoff. Debate about
the precise definition of climatic extremes, however, must
not distract from the fact that a large and growing num-
ber of studies—all examining the impacts of events in the
tails of their distributions—have shown that extreme cli-
matic events have major impacts on insects. In this sec-
tion, we summarize what is known about the impacts on
insects of four types of extreme events: temperature
extremes, droughts, rainfall events, and fire.
Heatwaves and extreme temperatures
With climate change, heatwave conditions are increasing
in frequency, intensity and duration (Christidis
et al., 2015; Frich et al., 2002; Meehl & Tebaldi, 2004;
Perkins et al., 2012). These extreme conditions may occur
in exposed micro-habitats (Gols et al., 2021), and can be
especially stressful for biological functions, particularly in
ectothermic species (Williams et al., 2016) (Figure 3c,d).
One way to measure vulnerability to extreme heat is to
compare heat tolerance limits with habitat temperatures
(Hoffmann et al., 2013; Pincebourde & Casas, 2019), and
such analyses have shown that terrestrial insects may fre-
quently be exposed to lethal temperatures when they are
exposed to the sun, highlighting the need for behavioral
thermoregulation (Sunday et al., 2014). A methodological
complication with measuring heat tolerance is that the
duration of heat stress matters (Terblanche et al., 2007):
prolonged exposure to mild heat stress may be equally
detrimental as short exposure to intense heat, which is
the typical approach used to establish heat tolerance. For
example, a recent study suggests that such heat tolerance
limits may underestimate actual vulnerability, as heat
injury accumulates over time at temperatures below
those found to be lethal in rapid ramping assays
(Rezende et al., 2020, but see Dowd et al., 2015 for a
rejoinder). A recent meta-analysis shows that exposure to
climatic extremes (focusing on heat waves) generally
harms insect fitness in terrestrial ecosystems (Thakur
et al., 2021). This has led in some cases to local extinc-
tions as observed for several French populations of the
butterfly Parnassius apollo, which experienced extreme
warmth during winter followed by cooler, normal tem-
peratures during spring (Nakonieczny et al., 2007).
Heat stress has clear implications for survival. In
addition, heat stress (even a short exposure of <1 h
at 36C) can have significant consequences for reproduc-
tive processes and fertility. Heat-induced sterility has
been documented in a range of insect orders, including
Diptera, Hymenoptera, Coleoptera, Hemiptera, and
Odonata (reviewed in Walsh et al., 2019). Importantly,
both the characteristics of heat waves (i.e., intensity,
duration, and amplitude) and the insects experiencing
them can affect the outcome of these events, as suscepti-
bility varies among ontogenetic life stages (Abarca
et al., 2019; Bowler & Terblanche, 2008; Sales et al., 2021)
and across taxa (e.g., Quinn et al., 1994; Verberk
et al., 2018). Exposure to heat stress during development
can impact adult reproductive trait expression and mat-
ing behavior (Vasudeva et al., 2021). Heat stress during
adulthood also has consequences for male reproductive
function and success; experimental exposure to extreme
thermal conditions reduces sperm function and impairs
male reproductive competitiveness (Sales et al., 2018).
These consequences of heat stress can also have conse-
quences for offspring (Hoffmann et al., 2013; Roux
et al., 2010), with transgenerational damage being seen in
the reproductive fitness and lifespan of sons from either
fathers or sperm exposed to heatwave conditions in
Tribolium flour beetles (Sales et al., 2018). Although
heat-induced fertility impacts generally affect males more
than females, females appear unable to protect stored
sperm from heat-induced damage (Sales et al., 2018;
Walsh, Mannion, et al., 2021; Walsh, Parratt, et al., 2021).
These impacts can have potential consequences for the
operational sex ratio of heat-stressed populations (Walsh,
Parratt, et al., 2021). Similarly, behavioral changes in sex
allocation during simulated heatwaves have also been
reported (Moiroux et al., 2014). In turn, these impacts
may drive changes in mating behavior (e.g., propensity
for remating, Vasudeva et al., 2021) and processes linked
to natural and sexual selection (Walsh, Parratt,
et al., 2021). On the other hand, maternal exposure to
heat can have adaptive transgenerational effects that
involve e.g., tolerance and acclimation, and thus, gener-
ate thermal resistance (Zizzari & Ellers, 2014).
Ultimately, insect reproductive sensitivity to heat can dic-
tate global species distributions according to upper ther-
mal fertility limits in both sexes (Parratt et al., 2021),
indicating that climate change will have important wider
impacts on insect biodiversity.
Negative effects of extreme temperature events on
insects may also be linked with the stresses they induce
in plants with which many insects are intimately associ-
ated (Pincebourde et al., 2017). For example, larvae of the
moth Lobesia botrana fed on low-quality plants induced
by heat stress were negatively affected in terms of devel-
opment and immunity (Iltis, Louâpre, et al., 2021). The
major concern is that the increasing intensity of heat
waves is pushing many insect species and/or their food
plants beyond their adaptive limits, exposing them to
conditions that they may not have experienced in their
evolutionary history, particularly given the lack of
12 of 37 HARVEY ET AL.

adaptive capacity of insects to heat extremes (Harvey
et al., 2021; Harvey, Heinen, Gols, & Thakur, 2020; Ma,
Ma, & Pincebourde, 2021). Short-term (daily) tempera-
ture fluctuations can alter the shape of thermal perfor-
mance curves, reducing the optimum and critical
thermal maximum temperatures relative to those
predicted using mean temperatures alone (Kingsolver &
Buckley, 2018; Paaijmans et al., 2013).
Insect microbiomes are often driven by environmen-
tal microbiomes, for instance those associated with the
host plant or in the soil (Hannula et al., 2019). The effects
of extreme climatic events on environmental
microbiomes (Jansson & Hofmockel, 2020) may pose an
additional pathway through which climate extremes may
affect insect performance. For example, the bacterial
endosymbiont, Wolbachia, is known to impact the capac-
ity of mosquitoes to transmit a range of arboviruses and
parasites, and is being actively deployed as a biocontrol
agent in a number of locations globally. However, com-
plex interactions with Wolbachia appear to have highly
variable effects on malaria parasite infection under
extreme temperatures (Murdock et al., 2014). More
recently, lab studies examining thermal knockdowns
have suggested that Wolbachia can reduce the thermal
tolerance of the primary dengue vector, Aedes aegypti
(Ware-Gilmore et al., 2021). Moreover, infection with
dengue virus also reduced thermal tolerance. These
results demonstrate the potential for complex effects of
temperature variation (including temperature extremes)
on host–microbe interactions, with impacts varying
across environments. In particular, in cases where endo-
symbionts confer resistance to heat shock, this can allow
rapid evolution of heat tolerance by the host (Harmon
et al., 2009).
Extremely high temperatures are not only occurring
during the daytime. Many insects are nocturnal and are
sensitive to abiotic conditions that may differ signifi-
cantly from those experienced by diurnal insects.
Importantly, they may be highly sensitive to tempera-
tures that deviate considerably from normal. Night
warming and extremely high minimal nighttime temper-
atures are also threatening the persistence of some insect
populations and are also affecting interspecific interac-
tions (Higashi et al., 2020; Ma et al., 2020; Ma, Bai,
et al., 2021). However, the asymmetry between night and
daytime warming and extremes is thus far little studied,
especially in terms of its impact on diurnal insects whose
fitness-related traits, along with other overlooked behav-
iors such as sleep (Tougeron & Abram, 2017)or
thermally-gated developmental programs such as
molting, may be sensitive to heat exposure during the
night. For example, successive exposure to extremely
warm nights reduced adult performance (longevity and
fecundity) of the grain aphid Sitobion avenae over subse-
quent days (Zhao et al., 2014). In a study including
diurnal and nocturnal ant species, ant communities from
warmer habitats such as semi-deserts and subtropical dry
forests were more tolerant to high temperatures than ants
from cooler environments such as tropical rain and mon-
tane forests (García-Robledo et al., 2018). In all habitats,
nocturnal ants displayed lower thermal tolerance than
diurnal ants (García-Robledo et al., 2018). In addition,
night-time warming may have distinct effects from day-
time warming on top-down control of plants by herbi-
vores (Barton & Schmitz, 2018), and could interact with
other aspects of global changes such as light pollution to
disrupt predator–prey interactions (Miller et al., 2017).
Considering that insects have to keep up with an
increasingly variable thermal environment, which include
cold and heat shocks, it is crucial to examine the cumula-
tive impacts of fluctuating temperatures on the response of
insects to thermal extremes (Hance et al., 2007;Jeffs&
Leather, 2014). The cumulative effects of stressful condi-
tions on physiological performance is becoming increas-
ingly recognized (Cardoso et al., 2020;Didham,Barbero,
et al., 2020; Didham, Basset, et al., 2020; Harvey, 2015;
Kaunisto et al., 2016) and highlights the potentially much
higher vulnerability of insects to stressful conditions.
Climatic variability over different temporal and spatial
scales may notably limit the evolutionary responses of
insects to longer-term, incipient warming (Buckley &
Kingsolver, 2021; Kingsolver & Buckley, 2015).
Phenotypic plasticity and bet-hedging may be critical
strategies for the persistence of insect populations and
species in response to immediate, intense and more or
less predictable temperature changes (Sgrò et al., 2016).
Phenotypic plasticity in response to thermal variation is
known to protect insects by eliciting changes in a range
of important biological traits (e.g., Vasudeva et al., 2021;
Verberk et al., 2018). Phenotypic plasticity is expected
when climatic variation is at least partially predictable,
but fitness-related traits may be compromised when they
are more stochastic (Liefting et al., 2009). Bet-hedging is
an adaptive strategy in temporally unpredictable environ-
ments (Hopper, 1999). For instance, in environments
with highly unpredictable cold and heat extreme events,
it was demonstrated that several phenotypes can be
expressed among the progeny of a single individual (e.g.,
diapausing versus active individuals, sexual versus
asexual morphs) (Le Lann et al., 2021). Each of these
phenotypes may have advantages over the others
depending on thermal conditions. This diversified
bet-hedging strategy may ensure the survival of a part of
the progeny at each generation and an overall higher fit-
ness over generations compared to thermal specialists
producing a single phenotype. However, tightly
ECOLOGICAL MONOGRAPHS 13 of 37

interacting insect species such as hosts and their parasit-
oids can respond very differently to changes in the mean
and variance in temperature due to trade-offs, evolution-
ary history, and genetic background with parasitoids
being usually more sensitive to thermal stresses (Hance
et al., 2007; Le Lann et al., 2021). Moreover, phenotypic
plasticity and bet-hedging in response to heat waves can
depend on interactions with other species. Using an
aphid–ladybeetle system, Sentis et al. (2017) experimen-
tally investigated the effects of predators and heat shocks
on aphids and showed that heat shocks inhibit pheno-
typic and behavioral responses to predation (and vice
versa), and that such changes may alter trophic
interactions.
Seasonally variable effects of climate
extremes with other anthropogenic
stresses
Seasonal changes in the frequency of extreme tempera-
tures can also disrupt different stages of insect life-cycles.
For instance, if diapause is not initiated because of expo-
sure to warmer winter temperatures, there is a risk of pre-
cocious death of active stages in response to unpredictable
extreme cold spells. Indeed, even as mean winter tempera-
tures increase, the frequency and intensity of short-term
cold periods is also increasing, perhaps as a result of
changes in the strength of the jet stream, facilitating a
breakdown in the polar vortices (Tomassini et al., 2012).
Several consecutive days where temperatures are 10 or
more degrees below normal during an otherwise warm
winter can have negative effects on populations and mod-
ify the relative abundances of competing species (Andrade
et al., 2016;Tougeronetal.,2018), due to their differences
in cold tolerance (Le Lann et al., 2011). For example, cold
spells occurring during winter seem to be an important
factor that determines aphid-parasitoid-hyperparasitoid
community composition in the following spring
(Tougeron et al., 2018). Cold temperatures during winter
months are critical for most insects in temperate biomes
(Hahn & Denlinger, 2011), but climate change may alter
the frequency of such cold events.
The impacts of climatic variability in both terrestrial
and aquatic environments cannot be seen in isolation
and are compounded by other stressors including habitat
loss, removal of refugia, and chemical pollution
(Cavallaro et al., 2019; Liess et al., 2021). Aquatic insects
are especially vulnerable to pollutants; under increased
temperature variability, damselflies’bioenergetic
responses (balance between energy gains and losses) are
more likely to be negative (Verheyen & Stoks, 2020).
These impacts are further aggravated by climate change
mediated reductions in body size (Verheyen &
Stoks, 2019). Similarly complex interactions are expected
to occur in farmland soils or above-ground habitats, but
wait to be characterized.
Drought
Drought is another climatic extreme that threatens
insects. In several different regions, the duration and
intensity of prolonged (acute) droughts is increasing and
is concomitant with above average temperatures, heat
waves and often fire (Dai, 2011; Williams et al., 2022).
Pulsed droughts, on the other hand, may also be
prolonged but are briefly broken by intense rainfall
events (Harris et al., 2018). Both types of drought can
have directly negative physiological effects on insects, or
induce effects on plant communities and insects that
depend on them for food and shelter up to the terminal
end of the food chain (Gutbrodt et al., 2011; Han
et al., 2022; Jactel et al., 2012; Ploughe et al., 2019).
The effect of drought stress on insects is complex and
depends on multiple factors. For instance, insects feeding
on trees may respond to drought quite differently than
insects feeding on smaller plants such as forbs, sedges
and grasses (Gely et al., 2021). During the summer,
drought episodes can decrease herbivorous insect
populations on small plants because these are more
prone to water stress, and this in turn will lead to a scar-
city in food resources that in turn has severe conse-
quences in terms of population dynamics and of
interspecific interactions. For example, one consequence
of desiccation (and thereby loss) of plant tissues is an
increase in competition for hosts or prey among higher
trophic levels. By contrast, insects feeding on trees are
often “buffered”against drought, owing to the fact that
trees contain much greater root and shoot biomass and
can generally endure more intense periods of drought
than smaller plants. Nevertheless, drought stress can still
generate chemical, physiological, and chemical changes
in plants, irrespective of their mass (Anderegg
et al., 2015; Gely et al., 2020; Jactel et al., 2012).
Drought stress can alter foliar and root concentrations
of primary metabolites (e.g., nutrients, such as amino
acids and sugars) and secondary metabolites
(e.g., defensive allelochemicals) and this can affect the
growth and development of insect herbivores (Han
et al., 2016; Sconiers & Eubanks, 2017). A recent review
(Gely et al., 2020) provided a framework that linked
water stress from increased drought severity to insect per-
formance. They predicted that different herbivore guilds
will show different but predictable responses to drought
stress with most guilds being negatively affected, many
14 of 37 HARVEY ET AL.

wood borers being a favorable exception, at least in the
short term. There have been a few whole-forest drought
manipulation experiments. Insect responses to an experi-
mental drought in a tropical rainforest of North
Queensland, Australia, showed variable responses among
different feeding guilds (Gely, 2021). In the experimen-
tally droughted area, there was significantly more wood
borer damage to trees than in the control area (Gely
et al., 2021). Many ant species in Australian rainforests
take nectar from extrafloral nectaries whereas some also
rely on aphid honeydew. Food sources are reduced in
areas under drought, and stable isotope analysis indicates
that many ant species are becoming increasingly preda-
tory (Gely, 2021), which will have impacts on food webs
in these forests.
Droughts can affect reproduction, as some insect eggs
require water for development (Rohde et al., 2017).
Similarly, drought can change plant signaling and the qual-
ity of floral rewards for pollinators leading to reductions in
pollinator attraction and plant reproduction (Descamps
et al., 2018; Rering et al., 2020). Even a single severe
drought can alter plant-insect communities. Following a
severe drought in 1995 in the United Kingdom, the total
abundance of butterflies increased, but this was accompa-
nied by substantial changes in community composition,
particularly in more northerly, wetter sites. Specialist, vul-
nerable species were lost while generalist, widespread spe-
cies increased, likely because of enhanced opportunities for
recruitment from the larger regional populations. A year
later, communities had yet to return to equilibrium
(De Palma et al., 2017), signifying that episodic droughts
can lead to greater extinction risk, likely both in terms of
species and genetic diversity. A similar finding was reported
for butterflies in Arizona (Wagner & Balowitz, 2021).
Single, severe droughts may push the last remaining
ephemeral populations toward extinction, with a concomi-
tant loss of genetic diversity.
While the physiological and ecological mechanisms
associated with responses to extreme drought are
multi-faceted and not well understood, the consequences
are increasingly apparent. A recent mega-drought in
western North America had negative and long-lasting
effects on montane butterfly communities that were com-
parable in magnitude to the combined effects of decades
of habitat loss and degradation at lower elevations
(Halsch et al., 2021). In the case of dragonflies in the
Cape Floristic Region, which is subject to periodic
droughts, adults, even among localized endemic species,
temporarily use artificial ponds to pull through periods of
extreme drought (Deacon et al., 2019). In the same
region, dragonfly adults remained faithful to the pond
margins and continued to forage there until rains
returned, while water beetles soon departed from ponds
when major droughts continued (Jooste et al., 2020).
These responses indicate that freshwater insects can have
different behavioral responses to periodic droughts.
Conversely, less volant taxa and insect which have histor-
ically evolved in perennially humid to wet communities,
such as the faunas of cloud and rain forests would be
expected to be especially challenged by droughts
(Janzen & Hallwachs, 2021; Wagner, 2020). Climatic
extremes, like drought, generate “winners”and “losers”
among insects, based on changes in plant quality and
non-linear effects up the food chain, although, when put
into the context of other anthropogenic stresses, the
longer-term prognosis for insects is negative (Harvey,
Heinen, Gols, & Thakur, 2020).
Extreme rainfall, floods
Climate warming is also leading to an increase in the fre-
quency and intensity of rainfall events, such as those
occurring during thunderstorms and hurricanes (Armal
et al., 2018;Brooks,2013; Frame et al., 2020;
Guhathakurta et al., 2011). Extreme rainfall and accompa-
nying flooding can have both direct and indirect effects on
insects. The direct effects constitute displacement and
drowning. Heavy rainfall dislodges insects from plants
with small or less well-attached species being particularly
vulnerable (Beirne, 1970; Chen, Harvey, et al., 2019;
Moran et al., 1987). Indirectly, insects can be affected by
rainfall and flooding through changes in the abiotic envi-
ronment. Flooding and subsequent soil waterlogging
induces a number of alterations in important soil physico-
chemical properties like soil pH, redox potential and oxy-
gen level that in turn can lead to hypoxia or anoxia
(Ashraf, 2012), affecting soil-dwelling insects in particular.
Insects, including many soil-dwelling and riparian insects
have evolved various mechanisms to withstand short-term
hypoxia or anoxia (Harrison et al., 2018;Hoback&
Stanley, 2001; Woods & Lane, 2016), but these capacities
can be exceeded by longer-term soil flooding. Wet soil may
also force subterranean insects to the soil surface where
they are more vulnerable to attack by their natural ene-
mies (Beirne, 1970).
Changes in soil conditions can lead to changes in
above-ground primary and secondary plant metabolism
that affects the performance of insects feeding on them
(Ayres, 1993). At the same time, rain changes microcli-
matic conditions such as temperature and humidity
which are both important environmental variables affect-
ing insect performance. The sudden drop in temperature
during heavy downpours may reduce feeding activity and
thus extend development time (Chen, Harvey,
et al., 2019). Increased humidity may favor conditions for
ECOLOGICAL MONOGRAPHS 15 of 37

growth of some insects (e.g., aphids and grasshoppers),
but it also may promote infection with pathogenic viruses
and fungi (Beirne, 1970). Other indirect effects of extreme
rainfall on insects can occur through disturbance of the
insect’s habitat. Flooding occurring in the aftermath of
extreme rainfall events may cause the death of small
forbs and increase tree mortality. The effects of this on
insects are not always negative, as some insects thrive on
the woody debris left after severe storms, the regrowth of
shoots, or by colonizing new plants (Gandhi et al., 2007
and references within).
Poff et al. (2018) measured the response of
stream-dwelling insects to an extreme flooding event in a
mountainous area in northern Colorado. The resilience of
the aquatic insects in response to this event depended on
life history traits of the insects: taxa with mobile larvae
and terrestrial adult stages, at the time of the event,
were more persistent than those without these specific life
stages (84% vs. 25% taxa persistence). Some species were
extirpated altogether. After the floodwaters retreated,
genetic diversity declined in some species but increased in
others, suggesting rapid recolonization by some species
(Poff et al., 2018). Similarly, the soil microarthropod com-
munity (Collembola and Acari) of a grassland in the flood-
plain of the Saale river in Germany recovered in terms of
species richness and density within 3 months after a severe
summer flooding (Gonz
alez-Macé & Scheu, 2018). These
results suggest that communities are, to some extent, resil-
ient to these extreme disturbances. Not all members of a
community are equally resilient, however, and the genetic
diversity of populations may change in response to these
events. Some desert stream insects exhibit adaptive behav-
ior to escape flooding and use rainfall preceding flash
floods as a cue to crawl vertically away from the stream
(Lytle & White, 2007). However, this flood avoidance
behavior was only found in insects originating from
populations collected in streams where rain is a reliable
predictor of imminent flooding (Lytle & White, 2007).
A recent study showed that in a protected Costa Rican
tropical forest, parasitism frequency correlated negatively
with precipitation anomalies (i.e., extreme wet events),
suggesting a weakening of trophic interaction strength
(Salcido et al., 2020). On Barro Colorado Island (BCI,
Panama), the variation in the ambient temperature and
precipitation appear to affect the populations of certain
species of assassin bugs (Lucas et al., 2016). Entomologists
also observed that populations of some large Saturniidae
showed a significant increase over time (Basset
et al., 2017). The peaks in saturniid abundance were most
conspicuous with increasing average precipitation on BCI
(Anderson-Teixeira et al., 2015). These studies contrast
with recent findings indicating insect decline in both tropi-
cal and temperate regions. We clearly lack sufficient insect
monitoring data (Basset & Lamarre, 2019) to either refute
or support claims of global insect decline with respect to
tropical regions (Janzen & Hallwachs, 2019). How com-
munities may respond to unpredictable and recurring
extreme rainfall and flooding events, and how this may
affect community structure and functioning, especially
when they increase in frequency and intensity, is largely
unknown.
Fire
Droughts and modified patterns of precipitation have led
to alterations in global fire regimes in terms of extent,
duration, seasonality, and severity (Jain et al., 2021;
Nimmo et al., 2021). While fire is a lethal threat to many
animals, the scientific community is only beginning to
consider it as an integral component of climate change
and an evolving force affecting the response of organisms
to it (Nimmo et al., 2021; Whelan, 1995). Results from
studies on how fire affects insects vary due to differences
in weather, burn intensity, focal taxa studied, and season
of burn (Banza et al., 2021; Dell et al., 2017; Pryke &
Samways, 2012a,2012b; Saunders et al., 2021;
Swengel, 2001). Arthropods possess complex life histo-
ries, and responses are typically taxon-specific (Joern &
Laws, 2013), which limits the body of literature on inver-
tebrate responses to fire, and hinders the capacity to pro-
pose effective conservation policies in response to
extreme fire events (Saunders et al., 2021). More mecha-
nistic studies are crucially needed to ameliorate our abil-
ity to anticipate the consequences of changing fire
regimes.
Many of the insect taxa associated with early succes-
sional series and fire-adapted communities require peri-
odic burns for their persistence. Indeed, many wood
boring beetles and their natural enemies may be attracted
to fires. The impact of fire on arthropods varies from neg-
ative to neutral to positive with some taxa being highly
vulnerable, e.g., Araneae, while others are not, e.g.,
Coleoptera (Kral et al., 2017). Strong recovery of the her-
baceous understory can boost general arthropod abun-
dance (Campbell et al., 2007). Even for species that are
fire-dependent, positive effects of fires can be reversed
when fire regimes are dramatically altered. For example,
localized decreased species richness and/or abundance
after fires have been observed in South Africa (Pryke &
Samways, 2012a) and Australia (Andersen & Müller,
2000), although in South Africa at least there can be
rapid recovery as, e.g., pollinators expand outward from
fire refugia (Adedoja et al., 2019). In any investigation,
it is important to consider not only that burning has dif-
ferential ecological effects based on ecosystem sensitivity
16 of 37 HARVEY ET AL.

or dependency on fire, but also the variant spatial distri-
bution of fire across these different landscapes in terms
of extent (i.e., 10 vs. 10,000 ha
2
), fire frequency, fuel loads
within fire perimeters, and distance to refuges (Pryke &
Samways, 2012a). For example, some fire-resistant plants
with tightly packed leaf bases have been shown to pro-
vide refuges for insects and other arthropods even during
intense fires (Brennan et al., 2011). The importance of
these refuges in the resilience of insect communities
needs further investigation.
Alterations in fire regimes due to global change are
likely to be complex; for example, these changes can con-
tribute to phenological asynchronies in herbivore–enemy
interactions. Parasitoids have latent post-fire recovery
and temporal changes of seasonal burns may affect the
availability of holometabolous hosts at specific life stages
(Koltz et al., 2018). Similarly, Dell et al. (2019) found that
frequent fire resulted in a loss of specialized trophic inter-
actions, and this pushed trophic webs toward generaliza-
tion, including increases in the abundance of generalist
feeding Orthoptera and Lepidoptera. As a result, shorter
burn regimes can generate periodic pest outbreaks. If
these (and other orders) are more efficient at dispersing
during large wildfires, and more rapidly recolonize
post-burn, they could affect community structure and
function. Fires may also have far-reaching consequences
for aquatic insects, especially those that rely on terrestrial
environments during part of their life cycle. For example,
the eggs of some lentic taxa lie dormant in topsoil layers,
making them particularly vulnerable (Blanckenberg
et al., 2019).
Since little is known about longer-term effects of cli-
matic extremes and related events on insects, one effec-
tive method to measure demographic changes is to utilize
technological advances in insect identification, such as
eDNA metabarcoding (Jinbo et al., 2011). This would be
extremely useful immediately in the weeks, months, and
years after an extreme event, such as fire.
INSECT CONSERVATION UNDER
CLIMATE WARMING AND
CLIMATIC EXTREMES
In this paper, we have highlighted individual, population,
and community-level responses to climate change, but
landscape or ecosystem consequences have remained
largely undiscussed, whereas at these levels the conse-
quences are most influential. The balance should there-
fore shift toward these effects in terms of policy-making,
scientific research, and conservation approaches. Indeed,
the effects of climate change on insects are numerous
and often lineage-specific. They may vary across life
stage, physiological state, as well as across local biotic
and abiotic conditions and, thus, appear to be idiosyn-
cratic. Of course, many research topics seem idiosyncratic
until we begin to understand them better. If we want to
understand and mitigate the impacts of extreme climatic
events, and climate change in general, on insect biology
and insect declines, we need more research (and associ-
ated funding and political will) on the impacts of climate
change not only on the basic biology of insects, but also
on integrative aspects at the scale of the ecosystem
(Hof, 2021). The decline in insect abundance and biomass
we are now facing—and can expect in the future given
the effects of climate change described above and still
other stressors—will have far-reaching community-level
effects due to the fact that insects form the major part of
the second trophic level in many ecosystems. As insects
provide a critical contribution to ecosystem functioning
and hence ecosystem services, loss of insect biomass,
abundance, and diversity will therefore disrupt trophic
cascades, including declines of flowering plants and the
erosion of terrestrial food webs (Wilson, 1987). For exam-
ple, the large number of insects during the breeding sea-
son are a crucial component of nestling diets of many
bird species, and a decline in insect availability can
severely reduce nestling survival and fitness (Tallamy &
Shriver, 2021). Likewise, losses in biomass of up to 80%
in important pollinator taxa will inevitably have disrup-
tive consequences for pollination (Hallmann et al., 2017,
2021). Extreme climatic events affect many insect species
in the community simultaneously, exacerbating the dis-
ruptive ecological consequences.
Perhaps more indirectly, but equally disruptive, will be
the effects of large-scale insect decline at the ecosystem
level. Significant losses of insect abundance and diversity
may threaten ecosystem resilience through reduced func-
tional diversity (Ant˜
ao et al., 2020;Jonsson&
Malmqvist, 2000;Seymouretal.,2020). Functional redun-
dancy provides “insurance”against the loss of a few species
(Naeem et al., 2012; Naeem & Li, 1997). With the current
estimated rates of insect decline, functional diversity may
quickly approach the lower threshold of full functional
niche occupancy, meaning that further losses will jeopar-
dize ecosystem functioning. In addition to knowledge
about how species respond to climate extremes (response
traits), we also need to know how they affect ecosystem
processes (effect traits), in order to understand how com-
munity attributes are related to ecosystem functioning and
resilience (de Bello et al., 2021; Suding et al., 2008).
This response-to-effect trait framework is increasingly
adopted by land managers as it can guide landscape
actions and local measures to preserve insect functional
diversity. Among the conservation approaches that can be
undertaken, we can consider direct and relatively local
ECOLOGICAL MONOGRAPHS 17 of 37

approaches aimed at reducing the effects of climate change
on insect biodiversity through appropriate environmental
management, and global policy approaches involving the
general public. It is clear that climate change is harmful to
insects and biological processes involving insects at the
individual, population, community, and ecosystem levels.
We need to act now to minimize these impacts; we know
how to do it, but the decision-making and requisite
funding keep getting pushed down the road or onto the
shoulders of future generations.
Management approaches
To protect insects from climate change, and climatic
extremes, it is necessary to go beyond traditional surveys
that record insect presence-absence and understand their
physiological and behavioral tolerance to environmental
extremes. Insects have both physiological and behavioral
thermoregulation capacities that can prevent exposure to
harmful temperatures (Abram et al., 2017) or buffer them
against the damaging effects of extreme temperatures
once exposed (Ma, Ma, & Pincebourde, 2021). The envi-
ronmental elements that can act on them, and that can
be manipulated, are at both the landscape (macro scale)
and at the micro-habitat level (microclimates), but it is
important to know how management fits with the main-
tenance of other ecosystem services, as well as how each
individual taxon may respond (Oliver & Morecroft, 2014;
Tougeron et al., 2022). The conservation implications of
microclimatic diversity at fine scales are just beginning to
be explored—but a general conclusion from studies to
date is that insects will be more resilient to climate
change when they consist of intact communities with
high structural complexity and high levels of plant spe-
cies diversity, which together will generate diverse micro-
climatic refugia (Pincebourde et al., 2016; Woods
et al., 2015). Microclimates can be influenced by land-
scape properties (Oliver & Morecroft, 2014). For instance,
hedgerows, woodlots, sown vegetation, and flower strips
may represent microclimatic refuges for agrobiodiversity
in the face of extreme climatic events (Lenoir et al., 2017;
Thakur et al., 2020). The windbreak and antifreeze roles
of hedges has been widely studied and confirmed in agri-
cultural landscapes. Wooded and closed areas generally
have lower temperature amplitudes than open areas over
a daily scale but are also colder on average, which can
affect insect thermoregulatory abilities (Alford
et al., 2017; Tougeron et al., 2016). Similarly, field-level
crop diversification and cover cropping has been shown
to be promising (Pan et al., 2020). Mountains and other
sites of topographic complexity may provide microhabitat
diversity to animals challenged by climate change
(Forister et al., 2021; Halsch et al., 2021; Loarie
et al., 2009). Importantly, it is necessary to understand
the extent to which landscape properties can affect insect
tolerance to thermal extremes.
Little attention has thus far been paid to identifying
climate refugia, at least over short temporal scales.
Demonstrating the occurrence and role of some habitat
types or landforms in slowing the declines of some spe-
cies during contemporary climate change could have
great impact on active climate-adaptation strategies.
Despite the recognized importance of climate change
refugia, the ability to quantify their potential for facilitat-
ing species persistence remains elusive. Keppel et al.
(2015) developed a flexible framework for prioritizing
refugia, based on their potential to maintain biodiversity
in the face of climate change. For instance, the
highest-capacity climate-change refugia in Tasmanian
plants is primarily in cool, wet, and topographically com-
plex environments. This result agrees with studies
performed in mountain areas by Brighenti et al. (2021),
Tampucci, Gobbi, et al. (2017), and Tampucci, Azzoni,
et al. (2017) that demonstrated the role of several cold
rocky landforms (i.e., a surface mantle of rocky debris
and interiors composed of ice and rock; e.g., glaciers, rock
glaciers, debris-covered glaciers, ice-core moraines) as
potential warm-stage refugia for cold-adapted aquatic
and ground-dwelling insect species. However, under-
standing how the area and isolation of refugia mediate
changes in taxonomic, functional, and phylogenetic
insect diversity caused by climate change is a key step in
prioritizing the conservation of specific refugial sites that
optimize conservation value. For instance, the access to
such climate refugia through potential dispersal corridors
can rescue insect diversity (Thakur, 2020b). As pointed
out by Morelli et al. (2016), the physical and ecological
diversity of landscapes managed by public agencies sug-
gest that they already contain climate change refugia;
thus, these agencies need tools to detect and prioritize cli-
mate change refugia for management. Moreover, the role
of mammalian ecosystem engineers, which can quite dra-
matically influence the structure and composition of hab-
itats, can also generate localized refugia which benefit
insects during climatic extremes (Thakur et al., 2020).
Thus, management strategies that prioritize conservation
of large vertebrates will benefit smaller organisms,
including insects. One of the crucial factors dealing with
extreme climatic and weather events is to understand the
functional value of the topographic landscape. At sea
level, increasingly impactful high tides inevitably will
lead to direct loss of coastal habitats, by flooding, salt
intrusion, and erosion of dune crests and cliff faces.
Inland, ecological resilience can be gained by ensuring
that large-scale networks of conservation corridors over
18 of 37 HARVEY ET AL.

various elevations are in pace so that not only is more
habitat available, simply reducing the risk of population
loss through more land area being available, but also
because there is a greater chance that refuges are avail-
able (Samways & Pryke, 2016).
The transformation of industrial agriculture toward
agroecology also allows to bring structural diversity in
the landscape that can lead to a better resilience of
insect communities, but also of their biodiversity and the
ecosystem services they provide, e.g., biological control
(Altieri et al., 2015).
For freshwater insects there are several management
options to alleviate the impact of climate warming.
These include measures to improve or safeguard water
oxygenation by ensuring flow or improving water quality,
since low levels of oxygenation may exacerbate heat
stress for these insects. Given that pesticidal pollutants
amplify climate change impacts in aquatic settings
(e.g., Verheyen & Stoks, 2020) with cascading effects over
space and time (Brühl et al., 2021), their phase-down
should rapidly be pursued. Locally, warming can further
be mitigated by increasing shading or increasing ground-
water tables (e.g., by reducing drainage in catchments),
which restores the influence of cool, ground water.
A combination of these measures over larger spatial
scales will also result in a mosaic of different thermal
regimes and this landscape heterogeneity may help
aquatic insects find temporary refuge from heat events.
Prescribed burning is used in many countries to man-
age forests and woodlands. Where the effects of fire are
confined to relatively small areas, recolonization by
ground-dwelling invertebrates from adjacent unburned
areas can be rapid, with communities returning to nor-
mal by the following season (Nunes et al., 2000,2006),
and fires can also enhance habitat diversity (see Fire).
One potential solution to climate-driven changes on
insect assemblages is to counter the effects of extreme
fires with prescribed, managed fires in ecosystems that
rely on fire to maintain healthy structure and function.
In doing so, burning ameliorates effects of future fires by
increasing pyrodiversity: as fire moves across a region,
the resulting landscape includes a fine-scale mosaic of
burned and unburned patches, creating not only refugia
for insects, but conditions that promote spatial heteroge-
neity of resources and enhanced conditions for insect
communities (Kim & Holt, 2012; Koltz et al., 2018;
Ponisio et al., 2016). In longleaf pine ecosystems where
regular application of prescribed fire is extensively used,
juvenile and non-flying insects have been shown to climb
into the canopy where microclimate conditions are more
favorable for their survival (Dell et al., 2017). Land man-
agement practices that have excluded burning on
fire-evolved landscapes have created high fuel loads
which result in extreme fire events, eroding pyrodiversity
and resulting in concomitant reductions in insect biodi-
versity (Berlinck & Batista, 2020; Ponisio et al., 2016).
Ultimately, management of habitats across various
scales in response to climate change and climatic extremes
needs to consider that insects face numerous anthropo-
genic stresses that do not necessarily operate indepen-
dently (Harvey, Heinen, Gols, & Thakur, 2020;Wagner,
Fox, et al., 2021). For example, systemic insecticides trans-
locate to (extra-)floral nectar or honeydew (Calvo-Agudo
et al., 2019), negatively impacting a broad suite of flower
visitors and thereby deepen population-level impacts of
both stochastic or climate-related events. These diverse
stressors interact and therefore should not be mitigated in
isolation. Thus, it is vitally important that factors such as
habitat loss and fragmentation, invasive species, intensive
agricultural practices, various forms of pollution
(e.g., synthetic pesticides and fertilizer), and other stresses
are fully integrated into conservation management
approaches (Harvey, Heinen, Armbrecht, et al., 2020;
Hof, 2021; Pryke & Samways, 2012a). Only in this way will
declines in insects be stabilized or reversed.
Policy making and public participation
It is pertinent to the preservation of insect diversity and
all biodiversity, in general, that drastic changes are made
in the way humans see and treat our resource-limited
planet. We need a massive-scale mobilization with trans-
formative action to address the climate crisis. We echo
the call made by Ripple et al. (2021) to change course in
six areas, including a progressive reduction (and eventual
elimination) in the use of fossil fuels; curbing short-lived
air pollutants such as black carbon (soot), methane,
and hydrofluorocarbons; restoring and permanently
protecting Earth’s ecosystems to restore biodiversity and
accumulate carbon; switching to mostly plant-based
diets; moving away from indefinite gross domestic prod-
uct growth to ecological economics with a circular econ-
omy; and stabilize the human population.
It must be stressed that halting upward trends in ris-
ing carbon dioxide concentrations and global surface
temperatures will take decades, if not more, and there-
fore requires an immediate enforcement of efforts to halt
the drivers of climate change at the global level. The
Paris Agreement, along with COPs 1-26, which are global
efforts to tackle the climate problem with 196 partner
countries, a promising start. Agreements such as these
should, however, have clearly delineated goals within a
strict time-frame, and should strive for immediate imple-
mentation and a much higher degree of accountability.
The current division between land set aside for nature
ECOLOGICAL MONOGRAPHS 19 of 37

reserves and land assigned to agricultural production or
urban development, is far from balanced, and this urgently
needs to be addressed by regional governing bodies.
Existing natural areas need to be strictly preserved; our
planet can no longer afford to lose more pristine habitat.
We need to rethink and revise agriculture, with a strong
emphasis on ecological intensification of production sys-
tems. We can ecologically improve agricultural lands,
through optimization of the ecological matrix, and the cre-
ation of networks by interspersing corridors and stepping
stones of habitat within the agricultural landscape. This
will not only benefit insect species diversity via the provi-
sioning of habitat, but also might serve a crucial role in
mitigation of negative effects of climatic change and
extremes on insects through the creation of climate
refugia. Industrial agriculture in its current form is not
sustainable for the preservation of biodiversity. Unguided
pesticide application and over-use of industrial fertilizers
have many non-target side effects and pollute our ecosys-
tems (Bernhardt et al., 2017); whenever possible, their use
should be avoided and replaced with environmentally
sound alternatives. More strategic and targeted approaches
need to be adopted to ensure the productivity of the agri-
cultural system, while minimizing the detrimental effects
of excess fertilizer and pesticide inputs on (insect) biodi-
versity. Ecological intensification of the agricultural land-
scape has been unequivocally shown to benefit both
agricultural yield and diversity (Gurr et al., 2016). The con-
cept of ecological intensification should be further
extended and incorporated into our landscape and city
planning. Road verges, public green spaces, and local gar-
dens can form important habitats and refugia, which will
benefit insects and related animals, especially under cli-
mate change and climatic extremes.
Specific levels of action to directly protect insects can
range from global political interventions to that of individ-
ual choices and behavior. Although the conservation of bio-
diversity is a systemic challenge, every person can play a
role through their individual actions. Seen through the eye
of an insect, even small individual actions can make a huge
difference. In this context, it is necessary to invest in popu-
larizing the role of insects in ecosystems. Interesting experi-
ments like the use of charismatic species prove useful for
public awareness. Children should also be taught in ele-
mentary classes the vitally important role that insects play
in a healthy, functioning biosphere (Oberhauser &
Guiney, 2009). Also, scientific progress alone is unlikely to
result in desirable outcomes (Wyckhuys et al., 2022)and
needs to be paired with enabling policies, broad
awareness-raising, and stakeholder education. The evi-
dence is clear and the onus is on governing bodies to act
now. With species and habitats being lost every day, a
refusal or delay to act is not a wise choice.
Individual choices and behavior: What can
you do?
Although the most impactful actions are those that should
be implemented by governing institutions, decisions made
at smaller scales by individuals can still make a large dif-
ference for insect conservation (Cosquer et al., 2012;
MacDonald et al., 2015). This is especially relevant in the
context of climatic extremes. Most people live in cities,
which because of a lack of primary resources and suitable
habitats can be a hostile place for many organisms
(Bugnot et al., 2019; Parris et al., 2018). Furthermore, cit-
ies, which are dominated by concrete, tend to form strong
heat islands, which can exacerbate the effects of climate
extremes, especially those associated with rising tempera-
tures (Ramamurthy & Bou-Zeid, 2017). The high propor-
tions of sealed soil surface area may also increase the local
impact of precipitation extremes. It has become evident
that individuals can and are willing to play an important
role in making cities more suited to insect life and other
wildlife (MacDonald et al., 2015). Fortunately, the solu-
tions are generally low-cost. A good place to start is in
your garden or balcony, but even an appropriately
designed windowsill can be relevant. Four ingredients are
essential for insect survival in the face of climatic
extremes: suitable microclimate refugia; access to a water
source; sufficient nutrition; absence of pesticides
(Deguines et al., 2020). Many urban gardens are sealed-off
and neatly organized. None of insects’primary require-
ments for reproduction and survival are met in most gar-
dens. The solutions, however, are remarkably simple.
Sealed areas in a garden, as well as traditionally
well-maintained lawns, should be reduced to a minimum.
Exposed soil, and the plants that grow in it, provide the
most important microclimate needs, sources of moisture,
and nutrients. The choice of what to grow and where also
makes a difference. A highly diverse mixture of native
plants provides the most heterogeneous habitat, and sup-
ports the highest diversity of interactions. Cultivated plant
varieties should be avoided, as although they may appear
highly attractive, they often provide little nutritional
rewards for insects. Pollinators appear to prefer gardens
made up primarily of native plants with a few exotics pre-
sent (Salisbury et al., 2015) Many urban and suburban gar-
deners rely heavily on pesticides, paying little attention to
the label, ignoring recommended application rates and
possible collateral damage. Pesticide use should be avoided
altogether. Sowing native wildflower mixtures, even in
pots, can play a role in fulfilling the basic needs of local
insect diversity. Mowing should be limited, preferably
until after the flowering season and hence peak of insect
abundance. Leaving plant material, such as leaf litter,
standing senescing biomass, and a compost pile are other
20 of 37 HARVEY ET AL.

potential microrefugia that can make a difference.
Insect-friendly gardening reduces individual carbon foot-
prints and increases the rewards in the form of floral
abundance, which is appreciated by insects and (most)
humans alike. An insect-friendly garden is a beautiful gar-
den, but a beautiful garden is not necessarily an
insect-friendly one. We all can make a difference for the
preservation of insect diversity, especially in cities, through
the choices we make (Figure 4).
SUMMARY
Over the past several decades, increasing evidence is showing
that many insect taxa are experiencing rapid declines in both
temperate and tropical ecosystems. Whereas attribution to
any specific factor in explaining these declines is elusive,
there is little doubt among most researchers that
human-induced climate change is playing an important role.
Here it is crucial to distinguish between the effects of more
gradual, incipient warming and the effects of short-term
exposure to climatic extremes, the latter of which pushes
many species to (and beyond) their thermal tolerance.
However, placing overemphasis on any single factor is also
problematic. It is important to recognize that habitat loss and
fragmentation, chemical and organic pollution, invasive spe-
cies and other human-mediated changes to the environment,
which are broadly connected to human land use, are cur-
rently recognized as the main drivers of the declines of
insects and other invertebrate and vertebrate taxa as well.
Importantly, climate change will amplify the effects of other
factors, in particular human land use, and hamper the ability
of insects to avoid or adapt to multiple anthropogenic
stresses. This is because migration to new habitats tracking
climatic changes will not be possible if land use has already
converted these places into unsuitable habitat. Similarly,
land use can pose great barriers to dispersal. Species do not
exist in isolation, but communities and ecosystems are char-
acterized by a bewildering array of multitrophic interactions
that embody a labyrinth of complexity. Warming may dif-
ferentially affect species in food webs, leading to phenologi-
cal mismatches or the loss of key resources. The loss of
•Increase structural
complexity
•Denser vegetaon
•Reduce (sealed) pavement
•Increase natural soil cover
•Include (Semi-)permanent
water bodies
•Wild nave plant resources
year-round
•Balance between
herbs/shrubs/trees
•Keep lier and compost
•Rely on natural pest control
•Manual/minimal weeding
Refugia
Moisture
Nutrients
Pollutants
•Minimal vegetaon with
low diversity
•Sealed surfaces and
concrete (heat islands)
•Regularly disturbed soils
•Lack of water bodies
•Tidy environment
•Regular pescide usage
•Ferlizer applicaon
Insect destrucve
environmental features
Insect-supporve
environmental features
Provision of support to insects during
climate extremes
Crical
aenon
points
FIGURE 4 Local environmental characteristics can either harm or benefit insects (left panel) and this is especially notable when
insects are exposed to climatic extremes such as droughts and heatwaves. Intensively managed landscapes often lead to the simplification of
habitats, reducing plant diversity and thus limiting access to key resources for insects. This ultimately results in declining insect diversity.
Ecologically targeted management strategies (right panel) can rectify this by paying particular attention to several criteria that enhance
ecological communities from the bottom-up, with attention paid to both soil and above-ground processes, which benefits a wide range of
insects across different trophic levels. Images: WikiMedia commons. Users: Lawn: Paul Frederickson CC BY-SA 2.5; pavement: Michiel1972
CC BY-SA 3.0; garden: Fluteflute CC BY-SA 3.0; pesticide: Roy Bateman CC BY-SA 3.0; flower meadow: Ian Knox CC BY-SA 2.0; open
pavement: Titus Tscharntke CC BY-SA 3.0; tree/shrub/herb vegetation: Daderot CC BY-SA 3.0; leaf litter: Ceridwen CC BY-SA 2.0.
ECOLOGICAL MONOGRAPHS 21 of 37

insects also works its way up the food chain, and may be
playing an important role in the widespread decline of their
consumers, such as insectivorous birds in temperate
biomes. The broader ecosystem-level effects of insect
declineandtheroleplayedbyclimatewarmingthusneed
further attention. By conserving insect communities, and
by restoring the ecological balance in farming landscapes,
human welfare can be improved and substantial down-
stream societal benefits can be reaped. Given that climate
change continues unabated and climatic extremes in partic-
ular pose an immediate, short-term threat to insects, with
long-term consequences for ecosystems, it is essential to
also consider the importance of managing and restoring
habitats that make them as “climate-proof”as possible and
enable insects to find refuges in which they can “ride out”
extreme climatic events. At larger scales, corridors should
be maintained that enable insects to disperse over time to
more climatically suitable habitats. Most importantly, there
are means of safeguarding insect populations for posterity,
and we need to take the initiative to implement them. Our
contribution to the scientists’warning series thus highlights
the increasing threat that climate change and attendant
short-term climatic extremes pose to insects and other
ectotherms in terrestrial and freshwater ecosystems.
ACKNOWLEDGMENTS
We dedicate this paper to the memory of Edward
O. Wilson and Tom Lovejoy, and to co-authors Matthew
J. G. Gage and Simon Leather, who passed away during
the process of its preparation.
CONFLICT OF INTEREST
The authors declare no conflict of interest.
AFFILIATIONS
1
Department of Terrestrial Ecology, Netherlands Institute
of Ecology (NIOO-KNAW), Wageningen,
The Netherlands
2
Department of Ecological Sciences, Vrije Universiteit
Amsterdam, Amsterdam, The Netherlands
3
Earth and Life Institute, Ecology & Biodiversity,
Université catholique de Louvain, Louvain-la-Neuve,
Belgium
4
EDYSAN, UMR 7058, Université de Picardie Jules
Verne, CNRS, Amiens, France
5
Laboratory of Entomology, Wageningen University,
Wageningen, The Netherlands
6
Department of Life Science Systems, School of Life
Sciences, Technical University of Munich, Terrestrial
Ecology Research Group, Freising, Germany
7
Department of Biological Sciences, Smith College,
Northampton, Massachusetts, USA
8
Agriculture and Agri-Food Canada, Agassiz Research
and Development Centre, Agassiz, British Columbia,
Canada
9
Smithsonian Tropical Research Institute, Panama City,
Republic of Panama
10
Department of Ecology, Institute of Entomology, Czech
Academy of Sciences, Ceske Budejovice, Czech Republic
11
Groningen Institute of Evolutionary Life Sciences,
University of Groningen, Groningen, The Netherlands
12
School of the Earth, Ocean and Environment and
Department of Biological Sciences, University of South
Carolina, Columbia, South Carolina, USA
13
Rocky Mountain Biological Laboratory, Gothic,
Colorado, USA
14
Institut de recherche en biologie végétale, Département
de sciences biologiques, Université de Montréal,
Montréal, Québec, Canada
15
Laboratory for Integrative Biodiversity Research
(LIBRe), Finnish Museum of Natural History Luomus,
University of Helsinki, Helsinki, Finland
16
Department of Conservation Ecology and Entomology,
Faculty of AgriSciences, Stellenbosch University,
Stellenbosch, South Africa
17
Geosciences and Natural Resources Department,
Western Carolina University, Cullowhee, North
Carolina, USA
18
Université Cote Azur, INRAE, CNRS, UMR ISA, Nice,
France
19
Department of Zoology and Physiology and Program in
Ecology, University of Wyoming, Laramie,
Wyoming, USA
20
School of Biological Sciences, Monash University,
Melbourne, Victoria, Australia
21
Department of Marine Science, University of Otago,
Dunedin, New Zealand
22
University of Nevada Reno –Ecology, Evolution and
Conservation Biology, Reno, Nevada, USA
23
Department of Entomology, University of Maryland,
College Park, Maryland, USA
24
Department of Ecology and Evolutionary Biology,
University of Tennessee, Knoxville, Knoxville,
Tennessee, USA
25
School of Biological Sciences, University of East Anglia,
Norwich, UK
26
Ecology and Evolutionary Biology, University of
Connecticut, Storrs, Connecticut, USA
27
Centre for Tropical Environmental and Sustainability
Science, College of Science and Engineering, James Cook
University, Cairns, Queensland, Australia
28
MUSE-Science Museum, Research and Museum
Collections Office, Climate and Ecology Unit, Trento,
Italy
22 of 37 HARVEY ET AL.

29
Radboud Institute for Biological and Environmental
Sciences, Radboud University, Nijmegen,
The Netherlands
30
Energy and Resources Group, University of California,
Berkeley, California, USA
31
Department of Biogeography, Trier University, Trier,
Germany
32
IUCN SSC Invertebrate Conservation Committee
33
Bio21 Institute, School of BioSciences, University of
Melbourne, Melbourne, Victoria, Australia
34
Department of Biology, University of North Carolina,
Chapel Hill, North Carolina, USA
35
Laboratorio de Control Biol
ogico, Universidad de Talca,
Talca, Chile
36
Center for Integrated Pest Management, Harper Adams
University, Newport, UK
37
Department of Zoology, Stockholm University,
Stockholm, Sweden
38
Zoological Institute and Museum, University of
Greifswald, Greifswald, Germany
39
University of Rennes, CNRS, ECOBIO [(Ecosystèmes,
biodiversité, évolution)] - UMR 6553, Rennes, France
40
Department of Entomology, The Pennsylvania State
University, State College, Pennsylvania, USA
41
Climate Change Biology Research Group, State Key
Laboratory for Biology of Plant Diseases and Insect Pests,
Institute of Plant Protection, Chinese Academy of
Agricultural Sciences, Beijing, China
42
Avignon University, UMR IMBE, Avignon, France
43
Department of Biology, Texas State University,
San Marcos, Texas, USA
44
Department of Agricultural Biology, Colorado State
University, Fort Collins, Colorado, USA
45
Graduate Degree Program in Ecology, Colorado State
University, Fort Collins, Colorado, USA
46
Institut de Recherche sur la Biologie de l’Insecte, UMR
7261, CNRS, Université de Tours, Tours, France
47
Department of Forest Ecosystems and Society, Oregon
State University, Oregon, USA
48
Netherlands Institute of Ecology (NIOO-KNAW),
Department of Animal Ecology, Wageningen,
The Netherlands
49
INRAE, Aix-Marseille University, UMR RECOVER,
Aix-en-Provence, France
50
W.K. Kellogg Biological Station, Department of
Integrative Biology, Michigan State University,
East Lansing, Michigan, USA
51
Centre for Planetary Health and Food Security, School
of Environment and Science, Griffith University, Nathan,
Queensland, Australia
52
Institute of Ecology and Evolution, University of Bern,
Bern, Switzerland
53
York Environmental Sustainability Institute and
Department of Biology, University of York, York, UK
54
Bioprotection Aotearoa, School of Biological Sciences,
University of Canterbury, Christchurch, New Zealand
55
College of Science and Engineering, James Cook
University, Townsville, Queensland, Australia
56
Department of Animal Ecology and Physiology,
Radboud University, Nijmegen, The Netherlands
57
Department of Entomology, Department of Integrative
Biology, and Ecology, Evolution, and Behavior Program,
Michigan State University, East Lansing, Michigan, USA
58
Division of Biological Sciences, University of Montana,
Missoula, Montana, USA
59
Chrysalis Consulting, Hanoi, Vietnam
60
China Academy of Agricultural Sciences, Beijing,
China
61
Securing Antarctica’s Environmental Future, School of
Biological Sciences, Monash University, Melbourne,
Victoria, Australia
DATA AVAILABILITY STATEMENT
No data were collected for this study.
ORCID
Jeffrey A. Harvey https://orcid.org/0000-0002-4227-7935
Kévin Tougeron https://orcid.org/0000-0003-4897-3787
Rieta Gols https://orcid.org/0000-0002-6839-8225
Robin Heinen https://orcid.org/0000-0001-9852-1020
Mariana Abarca https://orcid.org/0000-0002-6944-2574
Yves Basset https://orcid.org/0000-0002-1942-5717
Matty Berg https://orcid.org/0000-0001-8442-8503
Carol Boggs https://orcid.org/0000-0001-7601-6277
Jacques Brodeur https://orcid.org/0000-0001-6148-7877
Pedro Cardoso https://orcid.org/0000-0001-8119-9960
Geert R. De Snoo https://orcid.org/0000-0002-5471-0425
Charl Deacon https://orcid.org/0000-0003-4601-2739
Jane E. Dell https://orcid.org/0000-0003-2791-2184
Michael E. Dillon https://orcid.org/0000-0002-7263-
5537
Grant A. Duffy https://orcid.org/0000-0002-9031-8164
Lee A. Dyer https://orcid.org/0000-0002-0867-8874
Jacintha Ellers https://orcid.org/0000-0003-2665-1971
Anahí Espíndola https://orcid.org/0000-0001-9128-
8836
James Fordyce https://orcid.org/0000-0002-2731-0418
Matthew L. Forister https://orcid.org/0000-0003-2765-
4779
Caroline Fukushima https://orcid.org/0000-0001-7909-
0173
Carlos García-Robledo https://orcid.org/0000-0002-
5112-4332
Claire Gely https://orcid.org/0000-0002-4089-5318
ECOLOGICAL MONOGRAPHS 23 of 37

Mauro Gobbi https://orcid.org/0000-0002-1704-4857
Caspar Hallmann https://orcid.org/0000-0002-4630-
0522
Thierry Hance https://orcid.org/0000-0001-5569-5020
Axel Hochkirch https://orcid.org/0000-0002-4475-0394
Christian Hof https://orcid.org/0000-0002-7763-1885
Ary A. Hoffmann https://orcid.org/0000-0001-9497-
7645
Greg P. A. Lamarre https://orcid.org/0000-0002-7645-
985X
William F. Laurance https://orcid.org/0000-0003-4430-
9408
Blas Lavandero https://orcid.org/0000-0002-2423-7016
Philipp Lehmann https://orcid.org/0000-0001-8344-
6830
Cécile Le Lann https://orcid.org/0000-0002-3949-4066
Joffrey Moiroux https://orcid.org/0000-0002-0132-3763
Lucie Monticelli https://orcid.org/0000-0002-0745-3905
Chris Nice https://orcid.org/0000-0001-9930-6891
Paul J. Ode https://orcid.org/0000-0001-7153-1077
Sylvain Pincebourde https://orcid.org/0000-0001-7964-
5861
Melissah Rowe https://orcid.org/0000-0001-9747-041X
Arnaud Sentis https://orcid.org/0000-0003-4617-3620
Alisha A. Shah https://orcid.org/0000-0002-8454-7905
Nigel Stork https://orcid.org/0000-0001-7812-2452
John S. Terblanche https://orcid.org/0000-0001-9665-
9405
Madhav P. Thakur https://orcid.org/0000-0001-9426-
1313
Matthew B. Thomas https://orcid.org/0000-0002-7684-
0386
Jason M. Tylianakis https://orcid.org/0000-0001-7402-
5620
Joan Van Baaren https://orcid.org/0000-0002-8552-
9645
Martijn Van de Pol https://orcid.org/0000-0003-4102-
4079
Hans Van Dyck https://orcid.org/0000-0002-2013-6824
Wilco C. E. P. Verberk https://orcid.org/0000-0002-
0691-583X
David L. Wagner https://orcid.org/0000-0002-7336-
3334
William C. Wetzel https://orcid.org/0000-0001-5390-
6824
H. Arthur Woods https://orcid.org/0000-0002-3147-
516X
Kris A. G. Wyckhuys https://orcid.org/0000-0003-0922-
488X
Steven L. Chown https://orcid.org/0000-0001-6069-5105
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How to cite this article: Harvey, Jeffrey A.,
Kévin Tougeron, Rieta Gols, Robin Heinen,
Mariana Abarca, Paul K. Abram, Yves Basset, et al.
2023. “Scientists’Warning on Climate Change and
Insects.”Ecological Monographs 93(1): e1553.
https://doi.org/10.1002/ecm.1553
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