Robby Stoks’s research while affiliated with KU Leuven and other places

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Publications (393)


Figure 3. Species richness (upper row) and Shannon diversity (bottom row) of the bacterioplankton (BPK, left) and gut microbiomes (GUT, right) of the different combinations of D. magna and D. pulex genotypes. Error bars represent one standard error of the mean. For the gut microbiomes, the representation of the genotypes is as follows: KNO(DB) are gut microbiomes from the D. magna genotype KNO that was in co-cultures with D. pulex genotype DB. DB(KNO) are gut microbiomes from the D. pulex genotype DB that were in co-cultures with D. magna genotype KNO. For two treatments (BPK: T2-DB and GUT: OM2(DK)) the error bars are missing, this is because only one replicum was left after rarifying the data (some samples were removed because of the low number of reads).
Figure 5. Relative abundance of the bacterial classes in the inocula (upper row, left), bacterioplankton (BPK, upper row, right) and gut microbiomes (GUT, lower row) of D. magna and D. pulex in mono-and co-cultures. The samples are grouped per species (m, p, mp, m-co and p-co) and genotype (KNO, OM2, T2, DB, DK and GB). m = D. magna monocultures, p = D. pulex monocultures, mp = bacterioplankton of co-cultures of D. magna and D. pulex, m-co = gut microbiome of D. magna co-cultures, p-co = gut microbiome of D. pulex co-cultures.
Figure 6. Venn diagram of the bacterioplankton, showing the unique and shared ASVs for D. magna in monocultures, D. pulex in monocultures and co-cultures. The numbers show the number of ASVs and the relative abundance in percentage of the ASVs per subgroup.
Figure 7. Venn diagram representing the unique and shared ASVs of the gut microbiomes, grouped per species and culture type (based on the rarified data). The numbers show the number of ASVs and the relative abundance in percentage of the ASVs per subgroup.
How does co-occurrence of Daphnia species affect their gut microbiome?
  • Article
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December 2024

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48 Reads

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Species co-occurrence can lead to competitive interactions that influence fitness. Competition is typically assumed to be modulated by species niche, especially food-acquisition related traits. The influence of interspecific interactions on host microbiome communities has rarely been considered, and yet may provide an alternative mechanism regarding the effect of host species co-occurrence on their fitness. Here, we investigated whether the composition of the gut microbial community differs between two Daphnia species (D. magna and D. pulex), and whether the gut microbiome of one species depends on the presence of the other. We hypothesized the stronger filter-feeder D. magna to have a larger effect on the gut microbiome of the weaker filter-feeder D. pulex than vice versa. To this purpose, three D. magna and three D. pulex genotypes were first made axenic and then grown in monocultures or in co-cultures in natural environmental bacterioplankton-enriched water, before assessing the community composition of the gut microbiomes and bacterioplankton. We found that the composition of the gut microbiome of the two Daphnia species did not significantly differ overall. However, subtle differences (i.e. the relative abundance of certain bacteria) between mono- and co-cultures were found at the Daphnia genotype level. For most genotype combinations (six out of nine), the microbiome of D. pulex changed more (i.e. distance in microbiome composition was more sensitive to culture type in D. pulex than in D. magna) when grown in co-cultures with D. magna than in monocultures. This provides limited support for our hypothesis that the stronger filter-feeder has a larger effect on the gut microbiome of the weaker one than vice versa, and that this effect is possibly mediated via the bacterioplankton community.

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Insect responses to seasonal time constraints under global change are facilitated by warming and counteracted by invasive alien predators

October 2024

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56 Reads

In seasonal environments, organisms with complex life cycles not only contend with seasonal time constraints (TC) but also increasingly face global change stressors that may interfere with responses to TC. Here, we tested how warming and predator stress imposed during the egg and larval stages shaped life history and behavioural responses to TC in the temperate damselfly Ischnura elegans. Eggs from early and late clutches in the season were subjected to ambient and 4 °C warming temperature and the presence or absence of predator cues from perch and signal crayfish. After hatching, larvae were retained at the same thermal regime, and the predator treatment was continued or not up to emergence. The late eggs decreased their development time, especially under warming and when not exposed to predator cues. However, the late eggs increased their development time when exposed to predator cues, especially to crayfish cues. The TC decreased survival of late larvae that were as eggs exposed to crayfish cues, indicating a carry-over effect. The TC and warming additively reduced late larvae development time to emergence. Independent of the TC, predator cue effects on development time were stronger during the egg than during the larval stage. The late individuals expressed lower mass at emergence, which mirrored the size difference between field-collected mothers. Warming caused a higher mass at emergence. The late individuals increased their boldness and showed a higher number of moves, whereas warming caused a decreased boldness. There was no predator cue effect on larval behaviour. The results indicate that late individuals compensate for late season egg laying, which is facilitated under warming but counteracted under predation risk, especially when imposed by the crayfish.


How does co-occurrence of Daphnia species affect their gut microbiome?

September 2024

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95 Reads

Species co-occurrence can lead to competitive interactions that influence fitness. Competition is typically assumed to be modulated by species niche, especially food–acquisition related traits. The influence of interspecific interactions on host microbiome communities has rarely been considered, and yet may provide an alternative mechanism regarding the effect of host species co-occurrence on their fitness. Here, we investigated whether the composition of the gut microbial community differs between two Daphnia species ( D. magna and D. pulex ), and whether the gut microbiome of one species depends on the presence of the other. We hypothesized the stronger filter-feeder D. magna to have a larger effect on the gut microbiome of the weaker filter-feeder D. pulex than vice versa . To this purpose, three D. magna and three D. pulex genotypes were first made axenic and then grown in monocultures or in cocultures in natural environmental bacterioplankton-enriched water, before assessing the community composition of the gut microbiomes and bacterioplankton. We found that the composition of the gut microbiome of the two Daphnia species did not significantly differ overall. However, subtle differences between mono- and cocultures were found at the Daphnia genotype level. For most genotype combinations (six out of nine), the microbiome of D. pulex changed more when grown in cocultures with D. magna than in monocultures. This provides limited support for our hypothesis that the stronger competitor has a larger effect on the gut microbiome of the weaker one than vice versa , and that this effect is possibly mediated via the bacterioplankton community.



Ecological consequences of body size reduction under warming

August 2024

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225 Reads

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1 Citation

Body size reduction is a universal response to warming, but its ecological consequences across biological levels, from individuals to ecosystems, remain poorly understood. Most biological processes scale with body size, and warming-induced changes in body size can therefore have important ecological consequences. To understand these consequences, we propose a unifying, hierarchical framework for the ecological impacts of intraspecific body size reductions due to thermal plasticity that explicitly builds on three key pathways: morphological constraints, bioenergetic constraints and surface-to-volume ratio. Using this framework, we synthesize key consequences of warming-induced body size reductions at multiple levels of biological organization. We outline how this trait-based framework can improve our understanding, detection and generalization of the ecological impacts of warming.



Effects of the mesocosm predation treatments on the body size and population densities of the Daphnia magna from the pre‐ and high‐fish periods in the resurrected natural population: (A) Adult size, (B) average size, (C) adult density and (D) juvenile density. The letters above the boxplots represent significant differences among mesocosm predation treatments within each period, while the asterisks represent significant differences between the high‐fish period and pre‐fish period within each mesocosm treatment. Means are based on five replicate mesocosms per treatment combination.
Effects of the mesocosm predation treatments on the bioenergetic variables of the Daphnia magna from the pre‐ and high‐fish periods in the resurrected natural population under the three predation treatments: (A) Energy availability (Ea), (B) energy consumption (Ec) and (C) cellular energy allocation (CEA). Values are corrected for body dry mass (DM). The letters above the boxplots represent significant differences among mesocosm predation treatments within each period, while the asterisks represent significant differences between the high‐fish period and pre‐fish period within each mesocosm treatment. Means are based on five replicate mesocosms per treatment combination.
Effects of the mesocosm predation treatments on the body stoichiometry of the Daphnia magna of the pre‐ and high‐fish periods in the resurrected natural population: (A–C) stoichiometric ratios (C:N, C:P, N:P), and (D–F) elemental contents (%C, %N, %P). Body elemental contents are expressed as percentage of dry body mass (DM). The letters above the boxplots represent significant differences among mesocosm predation treatments within each period, while the asterisks represent significant differences between the high‐fish period and pre‐fish period within each mesocosm treatment. Means are based on five replicate mesocosms per treatment combination.
Clonal frequencies of Daphnia magna in the experimental mesocosm populations under the three mesocosm predation treatments at the start and after selection in the mesocosm experiment: (A) pre‐fish period Daphnia and (B) high‐fish period Daphnia. Clonal frequencies of each period are given based on five replicate mesocosms per treatment. Mean clonal frequencies with 1 standard error are shown.
Rapid evolution of consumptive and non‐consumptive predator effects on prey population densities, bioenergetics and stoichiometry

Predators can strongly influence prey populations not only through consumptive effects (CE) but also through non‐consumptive effects (NCE) imposed by predation risk. Yet, the impact of NCE on bioenergetic and stoichiometric body contents of prey, traits that are shaping life histories, population and food web dynamics, is largely unknown. Moreover, the degree to which NCE can evolve and can drive evolution in prey populations is rarely studied. A 6‐week outdoor mesocosm experiment with Caged‐Fish (NCE) and Free‐Ranging‐Fish (CE and NCE) treatments was conducted to quantify and compare the effects of CE and NCE on population densities, bioenergetic and stoichiometric body contents of Daphnia magna, a keystone species in freshwater ecosystems. We tested for evolution of CE and NCE by using experimental populations consisting of D. magna clones from two periods of a resurrected natural pond population: a pre‐fish period without fish and a high‐fish period with high predation pressure. Both Caged‐Fish and Free‐Ranging‐Fish treatments decreased the body size and population densities, especially in Daphnia from the high‐fish period. Only the Free‐Ranging‐Fish treatment affected bioenergetic variables, while both the Caged‐Fish and Free‐Ranging‐Fish treatments shaped body stoichiometry. The effects of CE and NCE were different between both periods indicating their rapid evolution in the natural resurrected population. Both the Caged‐Fish and Free‐Ranging‐Fish treatments changed the clonal frequencies of the experimental Daphnia populations of the pre‐fish as well as the high‐fish period, indicating that not only CE but also NCE induced clonal sorting, hence rapid evolution during the mesocosm experiment in both periods. Our results demonstrate that CE as well as NCE have the potential to change not only the body size and population density but also the bioenergetic and stoichiometric characteristics of prey populations. Moreover, we show that these responses not only evolved in the studied resurrected population, but that CE and NCE also caused differential rapid evolution in a time frame of 6 weeks (ca. four to six generations). As NCE can evolve as well as can drive evolution, they may play an important role in shaping eco‐evolutionary dynamics in predator–prey interactions.



Microgeographic differentiation in thermal and antipredator responses and their carry-over effects across life stages in a damselfly

February 2024

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65 Reads

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3 Citations

Global warming and invasive species, separately or combined, can impose a large impact on the condition of native species. However, we know relatively little about how these two factors, individually and in combination, shape phenotypes in ectotherms across life stages and how this can differ between populations. We investigated the non-consumptive predator effects (NCEs) imposed by native (perch) and invasive (signal crayfish) predators experienced only during the egg stage or during both the egg and larval stages in combination with warming on adult life history traits of the damselfly Ischnura elegans. To explore microgeographic differentiation, we compared two nearby populations differing in thermal conditions and predator history. In the absence of predator cues, warming positively affected damselfly survival, possibly because the warmer temperature was closer to the optimal temperature. In the presence of predator cues, warming decreased survival, indicating a synergistic effect of these two variables on survival. In one population, predator cues from perch led to increased survival, especially under the current temperature, likely because of predator stress acclimation phenomena. While warming decreased, predator cues increased larval development time with a proportionally stronger effect of signal crayfish cues experienced during the egg stage, indicating a negative carry-over effect from egg to larva. Warming and predator cues increased mass at emergence, with the predator effect driven mainly by exposure to signal crayfish cues during the egg stage, indicating a positive carry-over effect from egg to adult. Notably, warming and predator effects were not consistent across the two studied populations, suggesting a phenotypic signal of adaptation at a microgeographic scale to thermal conditions and predator history. We also observed pronounced shifts during ontogeny from synergistic (egg and early larval stage) toward additive (late larval stage up to emergence) effects between warming and predator stress. The results point out that population- and life-stage-specific responses in life-history traits to NCEs are needed to predict fitness consequences of exposure to native and invasive predators and warming in prey at a microgeographic scale.



Citations (83)


... The pattern of declining body size with warming in many aquatic ectotherms is known as the 'temperature-size rule' (hereafter TSR) (Daufresne, Lengfellner, and Sommer 2009) and is recognised as the 'third universal response to warming' (Forster, Hirst, and Atkinson 2012;Gardner et al. 2011). Warming-induced changes in body size can indirectly affect predator-prey size ratios and alter species interactions (Boukal et al. 2019;Sentis et al. 2024). Size-structured interactions can also cause abrupt shifts in community structure along environmental gradients (Lindmark et al. 2019;Thunell et al. 2021) due to emergent Allee effects associated with abrupt changes in population size structure (de Roos and Persson 2002), and these shifts can be triggered by changing body size (Dijoux and Boukal 2021). ...

Reference:

Differences in Tri‐Trophic Community Responses to Temperature‐Dependent Vital Rates, Thermal Niche Mismatches and Temperature‐Size Rule
Ecological consequences of body size reduction under warming

... This specific warming magnitude is close to or falls within the "very likely" range for projected increases in global mean surface temperatures between 2081 and 2100, according to the scenarios SSP2-4.5 (warming of 2.1-3.5 • C), SSP3-7.0 (warming of 2.8-4.6 • C) and SSP5-8.5 (warming of 3.3-5.7 • C). Moreover, this approach is supported by previous research that has used comparable temperature increases to study biological responses to warming with zebrafish and other aquatic species (Duan et al., 2023;Sulukan et al., 2022;Yang et al., 2020;Xu et al., 2024). Lastly, this temperature increase reflects the variations found in the natural environment of zebrafish, which include daily fluctuations of 5.6 • C, seasonal variations of up to 19 • C, and temperatures ranging from 16.6 • C to 38.6 • C depending on geographical location (López-Olmeda and Sánchez-Vázquez, 2011). ...

Local thermal adaption mediates the sensitivity of Daphnia magna to nanoplastics under global warming scenarios
  • Citing Article
  • June 2024

Journal of Hazardous Materials

... Theory and empirical evidence indicate a trade-off between both traits 3,4 . With some exceptions 5 , an early metamorphosis often comes with the cost of a smaller size 6 . This trade-off can be (partly) offset by an increased growth rate, enabling a short development to be combined with a large size at metamorphosis [7][8][9] . ...

Microgeographic differentiation in thermal and antipredator responses and their carry-over effects across life stages in a damselfly

... [74,75]). In addition, populations can evolve to adapt to novel conditions and changes in environmental variability [94,95]. Although the effects of these and other ecological and evolutionary processes on biodiversity are likely to be complex and often hard to predict [96], there are four ways that they could alter the null expectation of nonlinear dynamics of biodiversity loss driven by clustered warming tolerances. ...

When and how can we predict adaptive responses to climate change?

Evolution Letters

... To resist the harsh environment of the stomach lumen, LB use glutamate and aromatic L-amino acid decarboxylases which help in sustaining the transmembrane pmf to energize metabolism and transport [57,58]. The fact that these bacterial enzymes produce amino acid-derived compounds used as neurotransmitters (NTs) in bilaterian animals [59], such as serotonine, dopamine and GABA [55], both in the central nervous system (CNS) and the enteric nervous system (ENS) [60], suggests a possible feed forward loop [61] where proto-LB residing in the upper gut of some vertebrate ancestor would respond to food intake by generating an acidic environment, facilitating digestion and contributing to gut-brain signaling. ...

Hierarchical eco-evo dynamics mediated by the gut microbiome
  • Citing Article
  • October 2023

Trends in Ecology & Evolution

... Species possess intrinsic adaptive capacities to climate change characterized by three key components (Bellard et al. 2012;Swaegers et al. 2024): (1) adaptation, whereby species with very rapid generation times can adapt via evolutionary mechanisms (Hughes 2000;Swaegers et al. 2024); (2) plasticity, as certain species have the capacity to acclimate and modify their physiological traits, including growth, respiration, and tissue composition (Hughes 2000;Lindholm et al. 2015), and additionally, species may adapt their behavior, such as day-night activity rhythm and feeding locations (Dussault et al. 2004;Melin et al. 2014) and can also adapt their phenology, such as reproductive timing and dormancy cessation (Kannan et al. 2009;Roy & Sparks 2000;Walther et al. 2002); (3) dispersion, with certain species having the capacity to disperse quickly within their current habitat, when it has become unsuitable, to more suitable habitat (Dawson et al. 2011;Edelsparre et al. 2024;Swaegers et al. 2024). ...

Plasticity and associated epigenetic mechanisms play a role in thermal evolution during range expansion

Evolution Letters

... Eggs and larvae commonly share water bodies with top predators such as fish and crayfish to which they respond with changes in life history traits 5,32,33,35,37 . Cues from two different predators were used to induce nonconsumptive effects (NCEs). ...

Latitude‐specific urbanization effects on life history traits in the damselfly Ischnura elegans

... Verheyen et al., 2023). Cold-origin snails had a higher fat and sugar content at 12 °C, which 587 results in a higher CEA and is probably associated with the lower energy investment in 588 reproduction (lower number of egg masses and eggs per egg mass at lower temperatures). ...

Adverse effects of the pesticide chlorpyrifos on the physiology of a damselfly only occur at the cold and hot extremes of a temperature gradient
  • Citing Article
  • March 2023

Environmental Pollution

... This water system is a crucial part of the Fuhe River, the only river among the nine major systems that traverses the main urban area of Baoding City, connecting it with Baiyangdian Lake, the largest freshwater wetland in the North China Plain. Damselfly larvae were collected from June to July 2023 and screened based on physical characteristics (Theys et al., 2023). Identification was further verified by DNA barcoding (see Supplementary File: Paragraph 1). ...

Effects of heat and pesticide stress on life history, physiology and the gut microbiome of two congeneric damselflies that differ in stressor tolerance
  • Citing Article
  • March 2023

The Science of The Total Environment