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Climate Change Impacts: Insects

Erik Stange

Encyclopedia of Life Sciences

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Climate Change Impacts: Advanced article

Insects . Introduction
Article Contents

Erik E Stange, Norwegian Institute for Nature Research, Lillehammer, Norway . Effects on Physiology

. Species Distributions
Matthew P Ayres, Dartmouth College, Hanover, New Hampshire, USA . Phenology

. Population Dynamics

. Summary

Online posting date: 15th November 2010

Climate exerts powerful effects on the distribution and that projected climate changes will produce a range of
abundance of the earth’s insect species, and we should effects across insect populations’ physiology, distribution
expect climate warming to generate changes for many and abundance. In much the same way that the magnitude
insect populations and the ecosystems they inhabit. A of changes in temperature and precipitation patterns are
globally variable, insect species living in different habitats
substantial scientific literature provides a foundation for
will also vary both in the amount of climate change they
describing how insect species are responding to recent
experience and their sensitivity to such changes. There is a
climatic trends on the basis of insect physiology, and substantial scientific literature addressing climatic effects
predicting generalised species distributions and popu- on a range of different insect species and their populations,
lation dynamics for the future. Warmer temperatures which provides a foundation for describing how insect
generally lead to more rapid development and survival in species are responding to recent climatic trends and pro-
insects in mid- to high latitudes, which can account for viding some generalised predictions for future responses.
detectable and unambiguous shifts in a range of insect This article attempts to summarise the general effects of
species over the past half century. Increased warmth also climate on insects with regard to insect physiology, and
advances the onset of insect life cycles for the many spe- how climate changes could ultimately impact species’ dis-
cies that use thermal cues to match the timing of life his- tributions, phenology and population dynamics. See also:
tory events with the changing seasons. Owing to their
Biological Impacts of Climate Change; Insecta (Insects)
Climate exerts powerful effects on the distribution and
relatively short life cycles, high reproductive capacity and
abundance of the earth’s insect species similar to how it
high degree of mobility, insects’ physiological responses influences the global distribution of vegetation and plant
to warming temperatures can also generate particularly species (Sykes, 2009). Global patterns of temperature,
large and rapid effects on species population dynamics. precipitation, relative humidity, winds and solar radiation
set the physiological limits determining which insect species
are able to exist in certain habitats. Thus, the rapid changes
in these abiotic factors arising from greenhouse gas emis-
sions (IPCC, 2007) can affect the distribution or abundance
Introduction of any insect population directly. Climate changes can also
produce indirect effects on a focal insect species via more
direct effects on abundance of food resources, competitors,
Insects constitute about two-thirds of more than one mil-
enemies and mutualists (Figure 1) and affect the nature of
lion extant described animal species (Chapman, 2006).
biotic interactions among species (e.g. coefficients of
They are conspicuous denizens of terrestrial and aquatic
competition, attack rates and conversion efficiencies).
habitats from the equator to the poles, and display diverse
Understanding how climate change will affect an indi-
life histories at all imaginable trophic levels. All insects are
vidual insect species and the communities and ecosystems
responsive to temperature, so we should expect frequent
in which they live requires consideration of the magnitude
impacts of climate warming on insect populations and the
of expected climate change, the species’ capacity to
ecosystems they inhabit. However, we should also expect
accommodate variation in climate and the plasticity of
species with which the focal species interacts. See also:
Climate Change Impacts: Vegetation
ELS subject area: Ecology

How to cite:
Stange, Erik E; and Ayres, Matthew P (November 2010) Climate
Change Impacts: Insects. In: Encyclopedia of Life Sciences (ELS). John
Effects on Physiology
Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0022555
Insects are highly sensitive to temperature, with insect
metabolic rates tending to about double with an increase of

ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net 1
 Climate Change Impacts: Insects

Predators

+ −

Climate Insect herbivores

+ −

Plants

Figure 1 Climate change might affect distribution or abundance of any species directly (solid arrows), with possible effects on the other populations in the
community, or it could affect interactions among species (dashed arrows). Interactions among species could be physiological (e.g. phenological race
hypothesis and encapsulation) or demographic (e.g. biocontrol of whiteflies by Encarsia).

108C (Gillooly et al., 2001; Clarke and Fraser, 2004). This hardiness too early in the spring under new climates (Dukes
general pronounced effect of temperature on insect phy- et al., 2009).
siology presumably contributed to patterns in geological Climate can also generate indirect effects on a given
time of increased plant damage from insect herbivores in insect species via effects on the physiology of vegetation or
periods of increased temperature (Wilf, 2008). Climatic natural enemies (which may themselves be insects; Figure 2).
warming tends to influence (and frequently amplify) insect Changes in cloud cover, temperature precipitation, soil
species’ population dynamics directly through effects on nutrients and carbon dioxide can all impact the primary
survival, generation time, fecundity and dispersal (Bale and secondary chemistry of plant tissue, which influences
et al., 2002). At least in temperate and boreal regions, nutritional suitability for herbivores. Theory regarding the
increases in summer temperature will generally accelerate physiology of plant resource allocation permits predictions
the development rates of insects (and other poikilotherms) of the magnitude and direction of climatic effects on food
and can increase their reproductive potential (Sharpe, quality for insect herbivores (Herms and Mattson, 1992;
1977; Gillooly et al., 2002). Ayres and Lombardero, 2000). Plant tissue is generally
Climatic warming also reduces the risk to insect popu- most nutritionally suitable for insect herbivores when it has
lations of winter mortality due to extreme cold (Ayres and lower concentrations of plant secondary metabolites
Lombardero, 2000). All insect species have lower lethal (which are frequently antiherbivore defences), and higher
temperatures, and there are frequent records of demo- concentrations of the nitrogen that is often limiting for
graphic impacts (including the determination of poleward animal growth (Mattson, 1980). Reduced solar radiation
distribution) from temperatures that drop below lower (from increased cloud cover) and increased soil nutrients
lethal temperatures. For many insects (those described as (from greater soil microbial activity) tend to reduce con-
freeze-intolerant; Bale et al., 2002), the lower lethal tem- centrations of secondary metabolites; increased atmos-
perature is the temperature at which body fluids suddenly pheric carbon dioxide commonly reduces plant nitrogen
crystallise (the super-cooling point). Some insects are tol- concentrations and sometimes increases secondary
erant of freezing, but these also have relatively discrete metabolites; and modest drought frequently increases
lower lethal temperatures. One general effect of climate secondary metabolite concentrations (even though
warming on insects is to reduce the frequency in many mid- extreme drought can decrease concentrations of secondary
to high-latitude ecosystems of lethally cold temperatures. metabolites). The effects of these physiological responses
However, there will likely be some predictable deviations are best documented for insect herbivores in terrestrial
from this generalisation. Insects that overwinter in the litter systems (Ayres and Lombardero, 2000; Bidart-Bouzat and
layer of forests and grasslands may face higher mortality Imeh-Nathaniel, 2008). However, responses to climatic
rates from the loss of insulating protection provided by effects also extend to aquatic communities of insect detri-
snow (Lombardero et al., 2000). Also, insect cold hardiness tivores, where elevated carbon dioxide influenced aspen
varies seasonally, and diminishes with the approach of trees such that their leaves decomposed more slowly in
spring. If average winter temperatures become warmer but streams (Tuchman et al., 2002). See also: Ecology of
with increased variance in climate extremes (Karl and Invertebrate Nutrition; Plant Defences against Herbivore
Trenberth, 2003; IPCC, 2007), then bouts of cooler tem- Attack
peratures towards the end of the winter could produce Climate is capable of impacting the biological inter-
increased mortality for insect populations that lose cold actions between species. For example, if insect herbivore

2 ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
 Climate Change Impacts: Insects

Migratory arrival times &

Bird predators
energetic requirements

Bird predation rate versus

+ −
larval development

Larval development rate,

Climate Caterpillar herbivores
overwinter survival, etc.

Larval development + −
rate versus leaf maturation

Leaf maturation rate

Hardwood trees
and nutrient content

Figure 2 Effects climate change has on insects within hardwood forest ecosystem, where Lepidoptera are the dominant primary consumer. The effects
arrows’ labels provide some examples of direct effects and effects on trophic interactions, but these examples are not an exhaustive list.

development is generally more sensitive to temperature Species Distributions

than the development of host plant tissue, warmer con-
ditions will allow insects feeding in the early season to It seems to be common for insect herbivores to have geo-
complete more of their development while immature high- graphic distributions that are more limited than those of
quality foliage is still available (Ayres, 1993; Stange, 2009). their host plants (MacLean, 1983). However, life history
Colder conditions, however, may generally favour host traits common among insects – such as high dispersal
plants in this developmental race. Probably, climate ability, short generation times and high fecundity – make it
changes can also influence the biotic interactions between possible for populations of insect herbivores to rapidly
predatory or parasitic insect species and their prey or hosts, colonise habitats once the abiotic conditions become
but surprisingly little is known about this. One theoretical favourable. Since plant distributions cannot usually
possibility is that insect enemies of other insects tend to be expand as rapidly, climatic warming is likely to expand the
favoured in predator–prey interactions because their range of many herbivores further into the poleward portion
mobility, which influences their attack rates, depends on of their host plants’ distributions. Insects feature promin-
muscle performance, which is highly temperature sensitive ently among the documented range expansions that illus-
(Harrison and Roberts, 2000). If the temperature sensi- trate biological responses to recent climate change. The
tivity of insect predators or parasitoids is greater than that many examples are too numerous to provide here but
of their prey, warmer temperatures will generate a greater include the northern range expansions of Lepidoptera
role for top-down control of systems’ herbivory (Berggren (butterflies and moths) from more than 20 years of obser-
et al., 2009). Parasitoids can exert stronger effects on vations in Finland (Mikkola, 1997), Great Britain (Hill
their insect hosts at higher temperatures (Virtanen and et al., 2002) and western Europe (Parmesan et al., 1999). In
Neuvonen, 1999), even though the shortened development all such reviews that we know, poleward range expansions
time of their hosts means parasitoids have a narrower have been common and contractions towards the equator
timeframe when these hosts are vulnerable to attack. have been rare. Of the 35 non-migratory European but-
The spectrum of possible paths to ecological impacts is terfly species with established northern and southern
even richer because physiological responses of plants to extents studied by Parmesan et al. (1999), 22 species have
warming temperatures can simultaneously affect biotic expanded their ranges north by 35–240 km and only two
interactions between mobile parasitoids and their hosts. species have shifted south over the past 30–100 years.
Climate changes that result in lower nutritional quality of Odonates (dragonflies and damselflies) in Great Britain
plant tissue will increase consumption requirements for provide another example of temperate insect range shift.
insect herbivores, thereby increasing the duration of their Between 1960 and 1995, 23 of the 24 temperate species
development and corresponding exposure to natural expanded the northern limit of their distribution with a
enemies (Emmerson et al., 2004). There could be com- mean expansion of 88 km (Hickling et al., 2005).
pensating benefits for those that co-opt plant secondary Similar range changes are also occurring along elevational
metabolites as antipredator defences (Bjorkman and gradients. The winter pine processionary moth (Thaumetopea
Larsson, 1991), which can include encapsulation of para- pityocampa), a defoliating pest species whose larvae feed on
sitoid eggs (Kapari et al., 2006). pine needles during the winter, has extended its upper

ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net 3
 Climate Change Impacts: Insects

elevational boundary by 110–230 m in addition to a northern or more of their life history events to the changing seasons.
expansion of 87 km over the past 30 years (Battisti et al., Climatic warming therefore tends to advance the onset of
2005). Approximately one-third of this altitudinal range insect life cycles relative to Julian date. This plasticity is
expansion occurred following the exceptionally warm sum- generally adaptive because plants at the base of the food
mer of 2003, when warmer night temperatures enhanced the web are similarly responsive to temperature with regard to
flight dispersal abilities of gravid females (Battisti et al., 2006). the timing of leaf-out, leaf maturity, flowering, seed set and
In instances where host plants are not available at higher seed dispersal. Presumably there will be cases of insect
elevations, however, insect populations can experience range populations that become less abundant because of
constrictions. The black-winged white butterfly (Aporia cra- phenological mismatches between themselves and their
taegi) has disappeared from lower elevations of the Sierra de host plants (or prey) as climate change proceeds. Detailed
Guadarrama Mountains in central Spain since the 1970s, predictions will be possible with better understanding of
without a corresponding colonisation of the upper elevations physiological controls across taxa on seasonal rhythms. In
where its host plants are rare (Merrill et al., 2008). Field some cases, earlier emergence or hatching has increased
transplant experiments of eggs demonstrated the negative insect species’ voltinism, or the number of generations
impact that low elevation sites had on the viability of A. completed in a year. Examples include Lobesia botrana
cratagei populations, despite the abundance of host plants. It (Lepidoptera) a pest of grapes in Spain (Martin-Vertedor
is still unclear whether this effect was due to direct effects of et al., 2010) and Ips typographus (Coleoptera) a pine beetle
climate on A. cratagei eggs and larvae, or possible indirect in the European alps (Faccoli, 2009). The spruce beetle,
effects mediated by insect predators. Butterflies in central Dendroctonus rufipennis, has apparently responded to
Spain have constricted their ranges in response to warming, warmer temperatures by shortening the duration of its life
with the lower elevational limits of 16 species rising by an cycle in some regions from 2 years to 1 year, dramatically
average of 212 m – thereby reducing their habitat by increasing the mortality of its host trees (Logan et al.,
approximately one-third over 30 years (Wilson et al., 2005). 2003). Other examples of climate change producing
The effects of climate changes are evident in aquatic com- advanced insect phenologies, include the advanced flight
munities: over the past 25 years, the richness and abundance dates of Ephemeroptera (mayflies) in high altitude streams
of stream headwaters has diminished in response to warming (Harper and Peckarsky, 2006), Odonata in the Netherlands
water temperatures (Durance and Ormerod, 2007). (Dingemanse and Kalkman, 2008) and Lepidoptera in
Compared to mid- to high-latitude ecosystems, there is California (Forister and Shapiro, 2003). There also may be
less empirical knowledge and greater theoretical uncer- a tendency for reduced predation from birds in the early
tainty about the effects of climate change on tropical and growing season because physiological controls on the
subtropical insect populations. One hypothesis is that timing of bird reproduction frequently involve photo-
warmer is generally better for insects and all poikilothermic period and are therefore less responsive than insects or
animals (Frazier et al., 2006), and that low-latitude popu- plants to warming (Both et al., 2009).
lations will increase in abundance. An alternative
hypothesis is that there will be little effect on low-latitude
populations (static abundance) because they tend to be in
the flat part of their temperature response functions and the Population Dynamics
temperature increases are less in degree Celsius than at
higher latitudes and much less in terms of a percentage of Because insect species in general have relatively short life
annual thermal sum. Yet insects’ thermal tolerance is cycles, high reproductive capacity and high degree of
generally proportional to the magnitude of temperature mobility, the physiological responses to warming tem-
variation insects experience, and tropical insects currently peratures can produce large and rapid effects on species
may live in conditions that are quite close to their thermal population dynamics. We see a clear link between warm
optimum (Deutsch et al., 2008). Therefore, a third possi- climate conditions and some recent large-scale insect out-
bility is that there will be a loss of equatorial populations break events: including bark beetles in North America
concomitant with the poleward expansion of populations (Raffa and Berryman, 1987; Berg et al., 2006, Tran et al.,
(also referred to as the climate envelope hypothesis). One of 2007), Lepidoptera (caterpillar) defoliators in Scandinavia
the few documented examples of this involved the com- (Jepsen et al., 2008), aphids in the United Kingdom (Lima
munity of 102 species of tropical Lepidoptera along the et al., 2008) and the winter pine processionary moth in
slopes of Borneo’s Mount Kinabalu, which have moved the Europe (Battisti et al., 2005, 2006). Yet climate change also
geographic centre of their distributions an average of 67 m has the capacity to disrupt outbreak events or limit the
higher over the course of 42 years (Chen et al., 2009). poleward expansion of an insect species. Cold winters that
synchronise emergence of a pine beetle (Dendroctonus
frontalis) can produce low population abundances that can
Phenology subject a population to local extinction (Friedenberg et al.,
2007). Increased voltinism as a result of earlier emergence
Most insect species living at mid- to high latitudes employ dates of Japan’s green northern stinkbug, Nezara viridula
thermal cues (in whole or part) to match the timing of one (Heteroptera), can promote reproduction events that are

4 ENCYCLOPEDIA OF LIFE SCIENCES & 2010, John Wiley & Sons, Ltd. www.els.net
 Climate Change Impacts: Insects

too late into the autumn for the progeny to mature enough Summary
to survive the winter (Musolin, 2007).
Core theory from population ecology provides a solid The literature on insect species’ responses to climatic
general foundation for considering impacts of climate variation provides a solid foundation for predicting general
change on insects. Biological populations change as eR, effect climatic warming will have on insect populations and
where e is the base of the natural logarithms and R the per the communities in which they live. This remains an
capita change in population size from one time step to the extremely active area of research, and new studies continue
next. R at time step t tends to be negatively related to to enhance our understanding of the mechanisms by which
current and sometimes previous abundance (N). climate change affects individuals and ultimately the
dynamics of the population and community in which they
Rt ¼ f ðNt ; Nt 1 ; . . .Þ þ t ½1Š live. We expect climatic warming to generally increase the
abundance and distribution ranges of a majority of insect
If, as is commonly true, the density-dependent component species. However, the evidence supporting this assertion
of this relationship is approximately instantaneous and comes predominantly from studies of mid- to high-latitude
linear with a negative slope (Rt  b02b1
Nt), with b0 rep- populations of terrestrial insect herbivores. It is therefore
resenting the population’s potential growth rate and b1 the extremely important that new research projects continue to
per capita effect changes in a population’s size will have on explore effects climate changes can have on the groups of
its growth rate. This creates a stabilising endogenous insect species that are currently underrepresented in cli-
feedback system that tends to regulate the population mate change literature – including populations from tro-
around an equilibrium abundance (K  b0/b1), with vari- pical latitudes, aquatic habitats and higher trophic levels.
ance in N around K that is a function of the exogenous
(density-independent) variation in R(e). Climatic variation
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