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Running head: DAMSELFLIES IN AQUATIC ECOTOXICOLOGY
Integrating ecology and evolution in aquatic toxicology: insights from damselflies
Robby Stoks1, Sara Debecker2, Khuong Dinh Van3 AND Lizanne Janssens4
Laboratory of Aquatic Ecology, Evolution and Conservation, University of Leuven, B-3000
Leuven, Belgium
1 E-mail addresses: [email protected]
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Abstract.
Current legislation and ecological risk assessment fails in protecting aquatic biodiversity at
low levels of contaminants. We here address three topics embedded in general stress ecology
and evolutionary ecology that are relevant to arrive at a better evaluation of the risk of low
contaminant levels in aquatic systems: (i) delayed effects of contaminants, (ii) interactions
between contaminants and biotic interactors, and (iii) vulnerability to contaminants under
global warming. We develop these topics by capitalizing on the key insights obtained using
damselflies as model organisms. First, delayed contaminant effects on important fitness-
related effects could be shown both during the larval stage as well as after metamorphosis in
the adult stage. Second, synergistic interactions with bacteria and predation risk have been
demonstrated and advances in the mechanistic understanding of these synergisms with biotic
interactors are presented. Third, the strength of assessing the impact of contaminants under
global warming using a space-for-time substitution approach and the need to consider
temperature extremes is illustrated. These studies using damselflies as model organisms
highlight the relevance to consider contaminant effects after the exposure period and in the
presence of natural stressors such as predation risk and higher temperatures. They further
highlight the need for spatially explicit risk assessment and conservation tools. These insights
are relevant for most aquatic taxa. Indeed most aquatic taxa have a complex life cycle, are
strongly affected by predation risk and by warming and show latitudinal gradients. A better
integration of these topics in ecological risk assessment will be a major challenge for both
scientists and policy makers, but of crucial importance to preserve aquatic biodiversity.
Key words: ecological risk assessment, synergistic effects, global warming, space-for-time
substitution, stress ecology, complex life cycle
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Due to legislation, levels of contaminants such as pesticides are at low, sublethal
levels in large parts of the industrialized world (Kohler and Triebskorn 2013). Despite this,
recent continental-scale studies indicate that also in these regions strong contaminant-driven
losses in aquatic biodiversity are being detected. For example, pesticides at concentrations
that current legislation considers environmentally protective are causing significant effects on
both the species and family richness in European rivers, with losses in taxa up to 42% of the
recorded taxonomic pools (Beketov et al. 2013). Furthermore, there is strong evidence that
organic chemicals threaten the ecological integrity and consequently the biodiversity of
almost half of the European water bodies (Malaj et al. 2014). Ecological risk assessment
(ERA) of contaminants such as pesticides therefore fails in protecting aquatic biodiversity
(Beketov et al. 2013, Malai et al. 2014). This review focuses on three interrelated topics
embedded in general stress ecology and evolutionary ecology that may contribute to a better
evaluation of the negative effects of sublethal contaminant levels: (i) delayed effects of
contaminants, (ii) interactions between contaminants and biotic interactors, and (iii)
vulnerability to contaminants under global warming. While these topics were largely ignored
in traditional ERA, some of their aspects are starting to receive attention in recent ERA
guidelines (e.g. EFSA 2013).
In general, there is increasing appreciation to include fundamental concepts of ecology
(Relyea and Hooverman 2006) and evolution (Jansen et al. 2012) in toxicology to better
understand the impact of contaminants in natural systems. Among aquatic taxa, damselflies
have a long tradition of being used as model organisms to successfully study key topics in
ecology and evolution (Cordoba-Aguilar 2008). We here extend this view by highlighting that
damselflies also have been proven to be useful model organisms to address ecological and
evolutionary questions in aquatic toxicology. One major reason for their use as model
organisms in ecology and evolution is that their natural history and their responses to abiotic
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and biotic factors is exceptionally well known (Corbet 1999). This is especially true with
regard to the three topics under study (as indicated per topic below). Given that many
characteristics of damselflies are shared by most other aquatic taxa, the insights obtained by
using damselflies are likely widely transferable to other groups. Rather than advocating the
use of damselflies as model organisms in aquatic toxicology, we here want to illustrate how
damselflies have been successfully used to address key ecological and evolutionary topics in
aquatic toxicology and thereby provided some important novel insights and proofs-of-
principle to strengthen ERA in aquatic systems.
In this review we will first provide general background information on damselflies in
relation to aquatic toxicology. In the main part, we will illustrate some major insights into
each of the three above-mentioned topics that may contribute to stronger ERA and that were
obtained using damselflies as model organisms.
General background information on damselflies in relation to aquatic toxicology
Damselflies play an important ecological role and are therefore relevant to study in aquatic
toxicology (for examples, see Bried this issue). Any contaminant effects on damselflies likely
will cascade through food webs, potentially across ecosystem boundaries. Damselfly larvae
may contribute strongly to nutrient recycling (Ngai and Srivastava 2006) and, since they are
intermediate predators in aquatic food webs, effects on their densities or traits may affect
lower and higher trophic levels. Damselfly larvae are being preyed upon by predators such as
fish and large dragonfly larvae. They are themselves important predators of zooplankton and
other small invertebrates including two major standard test organisms in ecotoxicology: the
water flea Daphnia magna and Chironomus midges. Moreover, damselfly larvae are
important predators of larvae of vector mosquitoes making them useful biocontrol agents in
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mosquito control programs (Saha et al. 2012). Understanding the vulnerability of damselfly
larvae relative to these taxa is important to understand whether safe levels for standard test
organisms protect also higher levels in the food web and whether pesticide levels targeted to
kill vector mosquitoes in the larval stage do not impair biological control by invertebrate
predators. Studying the impact of contaminants on predator-prey interactions is getting
increased attention in ERA as impaired antipredator mechanisms may increase the negative
effects of contaminants that are not captured in the typical single-species toxicity tests carried
out in isolation (e.g. Brooks et al. 2009, Rasmussen et al. 2013).
During the aquatic larval stage damselflies cannot escape exposure to stressors making
them especially vulnerable to contaminants in aquatic systems. Moreover, damselflies can
accumulate pesticides and trace metals during their larval stage making them suitable for
biomonitoring (Van Praet et al. 2012, 2014a,b). Given their complex life cycle with an aquatic
larval stage where growth occurs and a terrestrial stage where dispersal and reproduction
occurs, they have the potential to link aquatic and terrestrial ecosystems (Stoks and Cordoba-
Aguilar 2012). Contaminants accumulated in the aquatic larval stage may be transported to
the terrestrial system after metamorphosis. For example, Yu et al. (2013) report a
bioaccumulation factor of 138,400 for polychlorinated biphenyls (PCBs) in adult dragonflies
collected at a river affected by the effluent from a wastewater treatment plant. This transfer
can relocate a considerable amount of this contaminant from aquatic to terrestrial habitats.
Furthermore, their complex life cycle makes damselflies useful organisms to study delayed
effects and interactions between stressors across metamorphosis.
Delayed effects of contaminants
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Effects of contaminants may persist or even only become detectable after the exposure period.
Such delayed effects are especially relevant to study for the many pesticides that are applied
in pulses and degrade quickly which may give the false impression that the effects of the
pesticide disappear after the pesticide has been degraded.
Delayed effects of pesticides within the larval stage
One important finding using damselfly larvae was that pulse exposure to some
pesticides may make them afterwards more vulnerable to predation risk (Janssens and Stoks
2012). After a 24h pulse exposure to low, sublethal levels of endosulfan and Roundup,
Enallagma cyathigerum larvae increased activity levels and even further increased mobility
when predation risk was present. Because the endosulfan pulse tended to also reduce escape
swimming speed, this resulted in an increased mortality by predation in larvae pre-exposed to
endosulfan. In contrast, previous exposure to Roundup caused the larvae to swim faster,
thereby offsetting the increased predation risk associated with the increased activity levels.
This study highlights that considering changes in predator–prey interactions may improve
ecological risk evaluations of short sublethal pesticide pulses. Furthermore, the study
underscored the need to effectively consider the outcome of predator–prey interactions.
Delayed effects of pesticides after metamorphosis
Delayed effects across metamorphosis are easily overlooked in the many animals that
have a complex life cycle. In several studies on damselflies it was shown that exposure during
the larval stage affected fitness-related traits in the adult stage (Table 1): metamorphosis
success, body mass, immune function, fat content and water content, flight-related traits, and
lifespan. Such carry-over effects may reflect the suboptimal levels of some traits already
present in the larval stage, such as lowered fat content, that have not been compensated during
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metamorphosis. Yet, suboptimal effects may also only become detectable after
metamorphosis. A striking example was found in Coenagrion scitulum where no effects of the
pesticide chlorpyrifos were observed during the larval exposure period while negative effects
only became apparent during and after metamorphosis (Dinh Van et al. in prep.). Another
related finding was that metamorphosis is the most sensitive stage when exposing E.
cyathigerum to the chemical perfluorooctane sulfonic acid (PFOS) in the egg and larval stage
(Bots et al. 2010). Two reasons may explain why such 'hidden' delayed effects only become
detectable after metamorphosis. Firstly, metamorphosis itself is a stressful event, associated
with high metabolic activity and upregulation of oxidative stress (Gaupale et al. 2012), and
animals may be unable to deal with both types of stress simultaneously. Secondly,
metamorphosis is often combined with a feeding stop that may take several days and thereby
may reduce energy reserves (Stoks et al. 2006). Yet, metamorphosis may as well mask larval
stress in the adult phenotype as the stress of metamorphosis may overrule the previous stress
effects. Indeed, while larval exposure to endosulfan increased the asymmetry levels in the
larval stage of C. puella, this was no longer the case in the adult stage. This could be
explained because asymmetry levels increased strongly during metamorphosis (Campero et al.
2008b).
Interactions between pesticides and biotic interactors
In natural systems animals typically are exposed to several stressors simultaneously
(Sih et al. 2004). This is a huge problem for standard ecotoxicological tests where animals are
exposed to contaminants while keeping abiotic and biotic conditions at optimal levels.
Additive effects of several mild stressors may potentially reach thresholds that impair fitness-
related traits and even reach lethal levels. Especially challenging for ERA are synergistic
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interactions where the negative effects of contaminants are magnified in the presence of
natural stressors. In a review by Holmstrup et al. (2010) it was concluded that contaminants
interacted in a synergistic way with natural stressors in more than 50% of the studies. It is,
however, poorly understood when such synergisms are to be expected. One way forward is to
start exploring the mechanisms underlying synergisms with contaminants. Aquatic toxicology
research using damselflies as model organisms focused on interactions between contaminants
and stress imposed by other species and identified synergistic interactions with non-
pathogenic bacteria (Janssens and Stoks 2013a) and predation risk (Campero et al. 2008c,
Janssens and Stoks 2013d).
Synergistic interactions with non-pathogenic bacteria
Given the general belief that non-pathogenic bacteria do not impair fitness, their
potential to modulate pesticide effects has been ignored. Yet, recently it has been documented
that the nonpathogenic bacteria Escherichia coli can negatively affect fitness-related traits in
butterflies (e.g. Freitak et al. 2007). This was confirmed in damselflies: exposure to non-
pathogenic E. coli reduced growth rate and fat storage in the damselfly E. cyathigerum
(Janssens and Stoks 2013a). This could be explained by a costly upregulation of immune
defence and stress proteins (Janssens and Stoks 2013a, 2014a). Notably, exposure to this
bacterium magnified the effects of the pesticide chlorpyrifos (Janssens and Stoks 2013a).
While chlorpyrifos inhibited its target enzyme acetylcholinesterase, this inhibition was much
stronger in the presence of E. coli. Furthermore, stress proteins that were upregulated in the
presence of chlorpyrifos to maintain cellular homeostasis, instead decreased when the
pesticide was combined with E. coli. Most importantly, upon exposure to chlorpyrifos the
immune defence was impaired and the bacterial load increased drastically. The observed
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synergistic effects between pesticides and widespread non-pathogenic bacteria are likely
common and deserve further attention to improve ERA of pesticides.
Synergistic interactions with predation risk
Given the ubiquity of predators in aquatic systems, a synergism that is getting
increased attention is the one between pesticides and predation risk (Relyea and Mills 2001).
Among invertebrates the physiological mechanisms of predation risk have been best
characterized in damselflies (e.g. Stoks et al. 2005a, b, Slos and Stoks 2008). Building on this,
three important mechanisms underlying synergisms between pesticides and predation risk
were identified using damselfly larvae as model organisms. A first important pathway
operates through interactive effects on resource acquisition and energy allocation. In response
to exposure to endosulfan C. puella larvae showed a growth increase. Yet, the pesticide
interacted with predation, causing a growth decrease (Campero et al. 2008c). The pesticide-
induced growth increase was due to an increased efficiency of assimilating ingested food.
However, when the pesticide was combined with predation risk, food intake, assimilation
efficiency and conversion efficiency all decreased, thereby generating the growth decrease.
Potentially, part of the assimilated food was allocated to detoxification and repair, for
example to stress proteins that are upregulated in pesticide-exposed damselfly larvae
(Janssens and Stoks 2013d).
A second potential mechanism is the inhibition of the enzyme acetylcholinesterase
(AChE), a typical target enzyme of insecticides such as carbamates and organophosphates
(Domingues et al. 2010). Research on the damselfly C. puella showed that exposure to
predation risk can inhibit this enzyme up to 70% (Campero et al. 2008c); which may be lethal
(Domingues et al. 2010). Additive inhibitions of AChE by a pesticide and predation risk
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therefore have the potential to generate synergistic effects on mortality when their combined
inhibition reaches the threshold above which mortality occurs.
A third mechanism identified in damselfly larvae is related to oxidative damage. A
study on E. cyathigerum showed that exposure to the pesticide glyphosate nearly doubled
oxidative damage to lipids, but only in the presence of predation risk (Janssens and Stoks
2013d). This could be explained by the fact that the pesticide inhibited one of the main
antioxidant enzymes in arthropods, superoxide dismutase, but only in the presence of
predation risk. This may be explained by energetic constraints when both stressors were
combined. Additionally, both pesticides (Lushchak 2011) and predation risk (Janssens and
Stoks 2013c) have been shown to increase levels of reactive oxygen species. Oxidative
damage to biomolecules may affect many fitness-related traits, such as swimming speed
(Janssens and Stoks 2014b). Therefore, it may underlie synergistic effects on these traits.
Vulnerability to contaminants under global warming
Besides contamination, global warming has been identified as a major threat to
biodiversity worldwide (Millennium Ecosystem Assessment 2005). Damselflies are ideal
study organisms in this context as they are particularly vulnerable to global warming (Hassall
and Thompson 2008; Hassall this issue). Exploiting the well-known latitudinal patterns in life
history in damselflies allowed advancing our knowledge on how local thermal adaptation
(hence gradual thermal evolution in response to global warming) may shape vulnerability to
contaminants. Note that the actual effects of climate change on contaminants remains difficult
to predict because temperature changes can affect chemical use, uptake, excretion,
biotransformation, fate, transport, bioavailability and the exposure duration to contaminants
(Rohr et al. 2011). We here only discuss the effects of warming on susceptibility or toxicity,
the effect of a chemical at a controlled exposure duration (Rohr et al. 2011).
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Gradual thermal evolution and the future ability to deal with contaminants in situ
There is increasing concern that higher temperatures may magnify the impact of
metals and pesticides such as organophosphates (Noyes et al. 2009, Moe et al. 2013). This is
directly relevant not only for future but also for current ERA as contaminants are typically
tested at room temperature, while along latitudinal gradients species may face higher
temperatures, hence potentially suffer higher toxicity from the same contaminant levels. The
extent to which temperature increases vulnerability to contaminants strongly depends on the
evolution of local thermal adaptation of populations along latitudinal gradients, and the
potential for gradual thermal evolution in situ when temperatures will further rise under global
warming. Surprisingly, large-scale latitudinal variation in vulnerability to contaminants has
very rarely been studied (but see Cherkasov et al. 2010).
Capitalizing on the well-documented latitudinal differentiation in life history in the
damselfly Ischnura elegans (e.g. Stoks et al. 2012), a set of studies addressed these topics
using a space-for-time substitution approach where populations are compared along a natural
temperature gradient. The performance of conspecific individuals from the cooler region
along the gradient can be tested at the current and predicted temperature under global
warming. Importantly, they can also be compared with the performance of organisms at the
warmer region of the gradient that have thermally adapted to the higher local temperature
(Stoks et al. 2014). Specifically, we reared larvae of replicated high-latitude (southern
Sweden) and low-latitude (southern France) populations in common garden experiments. We
did so at 20 °C and at 24 °C, the mean summer water temperatures in the shallow ponds
where I. elegans occurs at the high and low latitude, respectively. The 4 °C difference
matches the predicted temperature increase by 2100 under IPCC (2013) scenario RCP8.5. At
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each rearing temperature larvae were exposed to two types of contaminants: the trace metal
zinc and the pesticide chlorpyrifos. The current vulnerability to these contaminants of low-
latitude larvae at 24 °C probably provides a valid estimate of the expected vulnerability of
high-latitude larvae at a temperature increase of 4 °C given gradual thermal evolution in the
high-latitude populations.
For zinc, we found greater zinc-induced mortality and greater zinc-induced reductions
in activity at 24 °C compared to 20 °C in high-latitude, compared to low-latitude larvae (Dinh
Van et al. 2013, Janssens et al. 2014b). This pattern of local thermal adaptation indicates that
the predicted temperature increase of 4 °C by 2100 may strongly magnify the impact of zinc
at higher latitudes. It also indicates that gradual thermal evolution may mitigate the effects of
warming on the vulnerability to zinc. This urges caution when making predictions about the
effects of contaminants on a given species in a warming world based on studies that
investigated the effects of temperature at a single latitude. Additionally, this study
underscores the critical importance of considering local thermal adaptation along natural
gradients in ERA.
For chlorpyrifos, however, no thermal adaptation with regard to vulnerability to this
contaminant was found: chlorpyrifos only caused mortality at 24 °C and the reductions in
growth rate and food intake were stronger at 24 °C but this to the same extent at both latitudes
(Dinh Van et al. 2014a, b). Instead, there was evidence that the increased toxicity of
chlorpyrifos at 24 °C was stronger in terms of growth reduction in the faster growing low-
latitude populations. This is consistent with energy allocation trade-offs between growth rate
and pesticide tolerance (Sibly and Calow 1989). Another key finding was that while
chlorpyrifos impaired the predator behaviour (food intake) of the damselfly larvae more at 24
°C, this was not the case for their antipredator behaviour (activity and escape swimming
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speed). This suggests that the pesticide-induced changes in food web interactions may be
strongly different at high temperatures.
Heat waves and interactions across metamorphosis
Most studies simultaneously exposed organisms to contaminants and another stressor,
yet, there is increasing concern that stressors may also interact when imposed separately in
time (Segner 2011). Several studies have documented carry-over effects of larval pesticide
exposure on adult traits (Table 1). Yet, few studies have explored how exposure to
contaminants in the larval stage affects the sensitivity of the animals to a different stressor in
the adult stage (e.g. Rohr and Palmer 2005). How heat waves in the adult stage interact with
previous exposure to contaminants in the larval stage is particularly relevant in the context of
global warming. Under global warming an increase in the frequency and severity of extreme
weather events such as heat waves is expected (IPCC 2013). Despite this, temperature
extremes received very little attention in ecotoxicology (but see Kaur et al. 2011).
A study on I. elegans from low and high latitudes showed that a heat wave in the adult
stage interacted across metamorphosis with chlorpyrifos exposure during the larval stage
(Janssens et al. 2014a). Exposure to chlorpyrifos lowered the adult fat content and caused a
decrease in immune function. Yet, these effects were modulated in opposite ways by the heat
wave. When pesticide exposure was followed by a heat wave, the fat reduction was smaller
and the immunosuppression was stronger. As both exposure to the pesticide and the heat wave
caused a reduction in fat content, their joint effect may have been less than additive to avoid
fat levels going below a critical threshold value. In contrast, the pesticide-induced reduction in
immune function can be explained as a physiological cost of the stress exposure: adults that
had been exposed to the pesticide during the larval stage had probably a lower energy content
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and, as a result, they could have prioritized investment in defence against the stressor that was
present at that moment (i.e. heat) over investment in long-term defence (i.e. immune
response).
General conclusions
We have addressed three key topics in aquatic toxicology where damselflies have been
proven to be ideal model organisms to provide new insights and important proofs-of-principle
directly relevant to arrive at a more realistic ERA. First, with regard to delayed contaminant
effects, important fitness-related effects in response to aquatic exposure could be shown both
during the larval stage (for example, on predator-prey interactions) as well as after
metamorphosis in the adult stage. This illustrates that studies focusing only on effects during
the exposure period may underestimate the real impact of contaminants. Second, studies on
damselfly larvae made important contributions to the mechanistic understanding of the
widespread, yet little understood synergism between exposure to pesticides and predation risk.
Third, studies on damselflies illuminated the additional insights that can be obtained by
assessing the impact of contaminants under global warming by using a space-for-time
substitution approach and the need to consider not only increases in mean temperatures but
also temperature extremes. Combining a latitudinal gradient with common-garden warming
experiments revealed the potential of local adaptation in thermal tolerance and in life history
to mitigate the effects of contaminants in a warming world. These studies thereby not only
underscored the relevance of including temperature (Bednarska et al., 2013), and predator-
prey interactions (Brooks et al., 2009; Rasmussen et al., 2013) in ERA of contaminants, but
also highlighted the complexity of contaminant effects on predator-prey interactions being
differentially temperature-dependent pending on the trophic level. Moreover, these studies
illustrated the need for spatially explicit ERA and conservation tools. Indeed, low- and high-
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latitude animals often responded differently to contaminants. This highlights that intraspecific
evolution along natural thermal gradients may shape vulnerability to contaminants.
While studies using damselflies as model organisms added considerably insight in
these topics, they apply to many other aquatic taxa. Indeed most aquatic taxa have a complex
life cycle (e.g. many aquatic insects, amphibians), are strongly affected by predation risk
(Preisser et al. 2005) and by warming and show latitudinal gradients in life history (Stoks et
al. 2014). The reason why studies on damselflies were so successful in addressing these
general topics in aquatic toxicology is that, relative to most other aquatic taxa, a wealth of
background ecological and evolutionary information is available for this group. Further
integration of these topics in ERA of contaminants in aquatic systems is a major challenge for
both scientists and policy makers. Such effort is of crucial importance given that recent
continental-scale studies indicate that current ERA of contaminants and associated legislation
fails to avoid strong aquatic biodiversity losses (Beketov et al. 2013, Malaj et al. 2014).
Acknowledgements
We thank the many colleagues who helped us throughout the years to realize the here
presented research program and two anonymous reviewers and Chris Hassall for their
constructive comments. KDV benefited a PhD fellow of VOSP and an IRO Supplement, LJ
received an IWT PhD fellowship and postdoctoral fellowships from the University of Leuven
and the Research Foundation-Flanders (FWO-Flanders), SD is an FWO PhD fellow. This
research was financially supported by research grants from FWO- Flanders, the Belspo
project Speedy and the KU Leuven Centre of Excellence Funding PF/2010/07.
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TABLE 1. Overview of studies testing for effects of larval exposure to a contaminant on adult
fitness-related traits in damselflies.
Contaminant Damselfly
species
Adult trait Reference
Carbaryl X. zealandica Wing size =
Wing asymmetry ↑
Hardersen and Wratten
(1998)
Carbaryl X. zealandica Wing asymmetry ↑ Hardersen (2000)
Carbaryl X. zealandica Wing asymmetry =
Wing size =
Hardersen et al. (1999)
Chlorpyrifos C. scitulum Wing malformation ↑
Mass ↓ only in ♀
Flight muscle mass ↓ in edge ♂
Fat content ↓ in ♀ and in core
♂
Encapsulation response ↓
in ♂
PO activity ↓ in edge ♂ with
nylon filament
Dinh Van et al. in prep.
Chlorpyrifos E.
cyathigerum
Mass ↓ @ low food
Mass ↓ @ 18 and 24°C
Cold resistance ‘↑’
Janssens and Stoks (2013b)
Chlorpyrifos E.
cyathigerum
Mass ↓
Water content ↓
Fat content ↓ especially @ high
food
Janssens et al. (2014a)
24
529
530
Hsp70 ↑ especially @ high food
PO activity ↓ especially @ low
food in Swedish animals
PO activity ↓ especially @ low
food and heat stress
Flying ability =
Esfenvalerate C. scitulum Mass ↓
Metamorphosis success ↓
especially in edge populations
Dinh Van et al. in prep.
Endosulfan C. puella Mass =
PO activity ↓ especially @ low
food
Hemocyte numbers ↓ especially
@ low food
Campero et al. (2008a)
PFOS E.
cyathigerum
Metamorphosis success ↓
Mass =
Bots et al. (2010)
Zinc I. elegans Metamorphosis success ↓
especially @ low latitudes
Mass ↓
Adult lifespan ↓
Debecker et al. in prep
25
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