Biological control needs evolutionary perspectives of ecological interactions.

While ecological interactions have been identified as determinant for biological control efficiency, the role of evolution remains largely underestimated in biological control programs. With the restrictions on the use of both pesticides and exotic biological control agents (BCAs), the evolutionary optimization of local BCAs becomes central for improving the efficiency and the resilience of biological control. In particular, we need to better account for the natural processes of evolution to fully understand the interactions of pests and BCAs, including in biocontrol strategies integrating human manipulations of evolution (i.e., artificial selection and genetic engineering). In agroecosystems, the evolution of BCAs traits and performance depends on heritable phenotypic variation, trait genetic architecture, selection strength, stochastic processes, and other selective forces. Humans can manipulate these natural processes to increase the likelihood of evolutionary trait improvement, by artificially increasing heritable phenotypic variation, strengthening selection, controlling stochastic processes, or overpassing evolution through genetic engineering. We highlight these facets by reviewing recent studies addressing the importance of natural processes of evolution and human manipulations of these processes in biological control. We then discuss the interactions between the natural processes of evolution occurring in agroecosystems and affecting the artificially improved BCAs after their release. We emphasize that biological control cannot be summarized by interactions between species pairs because pests and biological control agents are entangled in diverse communities and are exposed to a multitude of deterministic and stochastic selective forces that can change rapidly in direction and intensity. We conclude that the combination of different evolutionary approaches can help optimize BCAs to remain efficient under changing environmental conditions and, ultimately, favor agroecosystem sustainability.

Prey life-history influences the evolution of egg mass and indirectly reproductive investment in a group of free-living insect predators.

The balance between risk and benefit of exploiting resources drives life-history evolution in organisms. Predators are naturally recognized as major drivers of the life-history evolution of their prey. Although prey may also influence the life-history evolution of their predators in the context of an evolutionary arms race, there is far more evidence of the role of predators than of prey.The goal of this study was to investigate the role of prey in life-history evolution of predators using ladybird beetle predators of aphids and coccids. These particular ladybirds and their prey were chosen because literature shows that the pace of life of aphids is faster than that of coccids and this difference is reflected in the life histories of the ladybirds that specialize on feeding on aphids or coccids.Thirty-four species of ladybird predators of aphids and eight of coccids belonging to five different tribes were collected and reared in the laboratory. The females were weighed as well as their eggs, and their reproductive investment estimated as the number of ovarioles. Phylogenetic relatedness was controlled for in the statistical analyses.Controlling for female mass revealed that ladybird predators of aphids lay bigger eggs than ladybird predators of coccids. This difference is not influenced by phylogenetic relatedness but only by the type of prey eaten. We suggest that ladybird predators of coccids lay smaller eggs because neonate larvae do not have to search, catch, and subdue prey. Both types of ladybirds have a similar reproductive investment relative to their body mass when phylogeny is controlled for.Recognizing the influence of prey on the life-history evolution of predators is important for understanding food web dynamics. From an applied perspective, this fine evolutionary tuning of prey-predator relationships should be used to guide and increase the efficiency of biological control programs.

New synonym of Nephus (Nephus) voeltzkowi Weise (Coleoptera: Coccinellidae), with comments on the origin of a Neartic population and its possible asexual status.

Coccinellidae (Coleoptera), commonly known as ladybirds or ladybugs, are a highly diversified family comprising nearly 6000 described species (Vandenberg 2002) distributed in 2 subfamilies and 24 tribes (Seago et al. 2011). The genus Nephus Mulsant, 1846, present worldwide, is currently placed in the vast Coccidulini tribe (Seago et al. 2011). There are different classifications for Nephus: Gordon (1976, 1985) considered five Nephus subgenera (Depressoscymnus Gordon, Nephus Mulsant, Scymnobius Casey, Sidis Mulsant, and Turboscymnus Gordon), while Fürsch (1987) considered nine Nephus subgenera, and later (Fürsch 1996) excluded Diomus Mulsant, 1850 as subgenus, leaving Bipunctatus Fürsch, 1987, Depressoscymnus Gordon, 1976, Geminosipho Fürsch, 1987, Nephus Mulsant, 1846, Parascymnus Chapin, 1965, Scymnobius Casey, 1899, Sidis Mulsant, 1850 and Turboscymnus Gordon, 1976. Gordon and González (2002) elevated Scymnobius to genus.

Phenotypic Response to Light Versus Shade Associated with DNA Methylation Changes in Snapdragon Plants (Antirrhinum majus).

The phenotypic plasticity of plants in response to change in their light environment, and in particularly, to shade is a schoolbook example of ecologically relevant phenotypic plasticity with evolutionary adaptive implications. Epigenetic variation is known to potentially underlie plant phenotypic plasticity. Yet, little is known about its role in ecologically and evolutionary relevant mechanisms shaping the diversity of plant populations in nature. Here we used a reference-free reduced representation bisulfite sequencing method for non-model organisms (epiGBS) to investigate changes in DNA methylation patterns across the genome in snapdragon plants (Antirrhinum majus L.). We exposed plants to sunlight versus artificially induced shade in four highly inbred lines to exclude genetic confounding effects. Our results showed that phenotypic plasticity in response to light versus shade shaped vegetative traits. They also showed that DNA methylation patterns were modified under light versus shade, with a trend towards global effects over the genome but with large effects found on a restricted portion. We also detected the existence of a correlation between phenotypic and epigenetic variation that neither supported nor rejected its potential role in plasticity. While our findings imply epigenetic changes in response to light versus shade environments in snapdragon plants, whether these changes are directly involved in the phenotypic plastic response of plants remains to be investigated. Our approach contributed to this new finding but illustrates the limits in terms of sample size and statistical power of population epigenetic approaches in non-model organisms. Pushing this boundary will be necessary before the relationship between environmentally induced epigenetic changes and phenotypic plasticity is clarified for ecologically relevant mechanisms with evolutionary implications.

A new species of Nephus (Nephus) (Coleoptera, Coccinellidae) described from Reunion Island.

We report here a new species belonging to Nephus (Nephus) Mulsant. Nephus (Nephus) apoloniasp. nov. was collected in the Reunion Island (Mascarene Archipelago, Indian Ocean). We describe this new species and redescribe and illustrate three other Nephus species already known from Reunion: Nephus (Nephus) oblongosignatus Mulsant, 1850, Nephus (Geminosipho) reunioni (Fürsch, 1974) and Nephus (Nephus) voeltzkowi Weise, 1910. Furthermore, we present a phylogenetic tree for these four species and calculate the genetic distances between them, using high-throughput DNA sequencing of the mitochondrial genome. The similar external morphology of N. apolonia sp. nov. and N. voeltzkowi very probably explains why individuals from the first species have been mistakenly identified as the latter and were not recognized as different until now. Other than external and genitalia traits, the present study provides molecular evidence confirming these are indeed two different species.

Intraspecific difference among herbivore lineages and their host-plant specialization drive the strength of trophic cascades.

Trophic cascades - the indirect effect of predators on non-adjacent lower trophic levels - are important drivers of the structure and dynamics of ecological communities. However, the influence of intraspecific trait variation on the strength of trophic cascade remains largely unexplored, which limits our understanding of the mechanisms underlying ecological networks. Here we experimentally investigated how intraspecific difference among herbivore lineages specialized on different host plants influences trophic cascade strength in a terrestrial tri-trophic system. We found that the occurrence and strength of the trophic cascade are strongly influenced by herbivores' lineage and host-plant specialization but are not associated with density-dependent effects mediated by the growth rate of herbivore populations. Our findings stress the importance of intraspecific heterogeneities and evolutionary specialization as drivers of trophic cascade strength and underline that intraspecific variation should not be overlooked to decipher the joint influence of evolutionary and ecological factors on the functioning of multi-trophic interactions.

The nature of the arena surface affects the outcome of host-finding behavior bioassays in Varroa destructor (Anderson & Trueman).

Varroa destructor, an acarian parasite of the Western honey bee Apis mellifera L., is a serious threat to colonies and beekeeping worldwide. The parasite lifecycle occurs in close synchrony with its host development. The females have to discriminate between different developmental stages of the host and trigger an appropriate behavioral response. Many studies have focused on these behavioral aspects, whether it is the choice of a precise host stage or the reproduction of female mites. Behavioral tests often require laboratory settings that are very different from the mite's environment. Our first experiment was designed to study the impact of the surface of test arena on the mite behavior. We found that plastic from Petri dishes commonly used as test arenas disturbs the female mites and can cause death. We searched for a substrate that does not harm mites and found that gelatin-coated plastic Petri dishes responded to these expectations. We then investigated the host choice behavior of phoretic mites confronted with larval stages of the bee on gelatin-coated arenas to watch if the well-documented orientation towards 5th instar larva was observable in our conditions. Pupal stages were included in the host choice experiments, initially to act as neutral stimuli. As white-eyed pupae were revealed attractive to the mite, several pupal stages were then included in a series of host choice bioassays. These additional experiments tend to show that the positive response to the white-eyed pupa stage depends on cues only delivered by living pupae. Further investigation on the nature and impact of these cues are needed as they could shed light on key signals involved in the parasite lifecycle.

Different phenotypic plastic responses to predators observed among aphid lineages specialized on different host plants.

The role of intraspecific variation in the magnitude and direction of plastic responses in ecology and evolution is increasingly recognized. However, the factors underlying intraspecific variation in plastic responses remain largely unexplored, particularly for the hypothesis that the herbivores' phenotypic response to predators might vary amongst lineages associated with different host plants. Here, we tested whether plant-specialized lineages of the pea aphid, Acyrthosiphon pisum, differed in their transgenerational phenotypic response to ladybird predators (i.e., the asexual production of winged offspring by wingless mothers). In a full factorial laboratory experiment, we found that six aphid clonal lineages each specialized either on alfalfa or clover significantly differed in their transgenerational phenotypic response to predators. Some lineages produced an increased number of winged aphids in predator presence while others did not respond. Aphid lineages specialized on alfalfa had stronger phenotypic responses to predators than those specialized on clover. Although we tested only six aphid lineages from two biotypes, our results imply that intraspecific variation in prey phenotypic response of herbivores to predators differs amongst lineages specialized on different host plants. Our findings therefore raise the question of the influence of plant specialization in shaping herbivore phenotypic responses, and highlight the need to consider multi-trophic interactions to understand the causes and consequences of intraspecific variation in complex phenotypic traits.

The evolution of chemical defenses along invasion routes: Harmonia axyridis Pallas (Coccinellidae: Coleoptera) as a case study.

The evolution of increased competitive ability (EICA) hypothesis (Blossey & Nötzold, 1995) postulates that escaping from coevolved enemies increases invaders fitness by energy reallocation from defenses and immunity to growth and reproduction. In this context, we evaluated the evidence of evolutionary change in invasive populations of Harmonia axyridis Pallas (Coccinellidae: Coleoptera). We measured egg defenses-cocktail of hydrocarbons on the egg's surface flagging egg toxicity and the concentration of the main alkaloid harmonine-in individuals from three populations along the invasion route (Japan: native, United States: introduced more than 30 years ago, South Africa: introduced in the early 2000s) in a common garden experiment. Our results support the EICA hypothesis: We found changes along the invasion route in the profiles of the hydrocarbons coating the eggs' surface and a decrease in the concentration of harmonine in eggs from the most recent invasive South African population compared to the long established in the United States and the native Japanese ones.

Evolution without standing genetic variation: change in transgenerational plastic response under persistent predation pressure.

Transgenerational phenotypic plasticity is a fast non-genetic response to environmental modifications that can buffer the effects of environmental stresses on populations. However, little is known about the evolution of plasticity in the absence of standing genetic variation although several non-genetic inheritance mechanisms have now been identified. Here we monitored the pea aphid transgenerational phenotypic response to ladybird predators (production of winged offspring) during 27 generations of experimental evolution in the absence of initial genetic variation (clonal multiplication starting from a single individual). We found that the frequency of winged aphids first increased rapidly in response to predators and then remained stable over 25 generations, implying a stable phenotypic reconstruction at each generation. We also found that the high frequency of winged aphids persisted for one generation after removing predators. Winged aphid frequency then entered a refractory phase during which it dropped below the level of control lines for at least two generations before returning to it. Interestingly, the persistence of the winged phenotype decreased and the refractory phase lasted longer with the increasing number of generations of exposure to predators. Finally, we found that aphids continuously exposed to predators for 22 generations evolved a significantly weaker plastic response than aphids never exposed to predators, which, in turn, increased their fitness in presence of predators. Our findings therefore showcased an example of experimental evolution of plasticity in the absence of initial genetic variation and highlight the importance of integrating several components of non-genetic inheritance to detect evolutionary responses to environmental changes.

Overwintering aggregations are part of Hippodamia undecimnotata's (Coleoptera: Coccinellidae) mating system.

Aggregation during diapause is a common phenomenon in arthropods that nevertheless remains poorly understood. The most commonly claimed benefit is that survival is higher in aggregations but animal aggregations could also be driven by sexual selection. In this perspective, we investigated whether aggregations in insects could be part of their mating system. We studied the overwintering aggregations of the ladybird Hippodamia undecimnotata (Schneider), an aphidophagous species from Southern and Eastern Europe as well as Asia. We collected ladybirds at three aggregation sites in Southwest France, during two overwintering periods (2013-2014 and 2014-2015). We checked their reproductive status by counting the viable sperm cells in the sperm storage organs of both males and females, and by assessing the ovarian status of females. We also investigated if mating behaviour occurred in these aggregations. We found that males have a high quantity of viable sperm cells (70-95%) in their reproductive organs throughout the overwintering periods. In contrast, although most females (85-95%) had empty spermatheca at the onset of the aggregations in autumn, the majority (65-91%) had numerous viable sperm in their spermatheca at the time of dispersal from the aggregation in early spring. Furthermore, frequent copulations were observed towards the end of the overwintering period, few weeks before dispersal. These results suggest that finding sexual mates may have been involved in overwintering aggregations in H. undecimnotata.

Effect of pollen extract supplementation on the varroatosis tolerance of honey bee (Apis mellifera) larvae reared in vitro.

As the main source of lipids and proteins in honey bees, pollen is a major nutrient provider involved in development and health and has been studied for tolerance stimulation against pathogens and parasites. In the case of Varroa destructor Anderson & Trueman (Acari, Mesostigmata: Varroidae) parasitization, the lack of a complete laboratory system to rear both the bee larva and the acarian parasite limited the studies concerning larval nutrition effects on the bee tolerance and resistance against varroatosis. Due to the development of this complete rearing protocol, we managed to feed young honey bee larvae with pollen supplemented solutions and to study the effect on their later development under parasitism conditions. In our experimental conditions, pollen influences neither the deformity rate, nor the survival of bees both parasitized and unparasitized. However, pollen extract supplementation seems to significantly impact the weight of the spinning bee larvae without having an effect on the physiological weight loss during pupation, so the differences found at the larval stage remain the same as at emergence. Varroa has a deleterious effect on bee pupae and led to a steady increase of the physiological weight loss experienced during metamorphosis. Interestingly, this ponderal loss associated with Varroa parasitization seems to be reduced in the polyfloral pollen supplementation condition. Altogether, this work is to our knowledge the first to study in laboratory conditions the impact of larval nutrition on the tolerance to parasitism. A diverse pollen diet may be beneficial to the bees' tolerance against V. destructor parasitism.

Non-additive effects of simulated heat waves and predators on prey phenotype and transgenerational phenotypic plasticity.

Understanding the effects of extreme climatic events on species and their interactions is of paramount importance for predicting and mitigating the impacts of climate change on communities and ecosystems. However, the joint effects of extreme climatic events and species interactions on the behaviour and phenotype of organisms remain poorly understood, leaving a substantial gap in our knowledge on the impacts of climatic change on ecological communities. Using an aphid-ladybeetle system, we experimentally investigated the effects of predators and heat shocks on prey body size, microhabitat use, and transgenerational phenotypic plasticity (i.e., the asexual production of winged offspring by unwinged mothers). We found that (i) aphids were smaller in the presence of predators but larger when exposed to frequent heat shocks; (ii) frequent heat shocks shifted aphid distribution towards the plant's apex, but the presence of predators had the opposite effect and dampened the heat-shock effects; and (iii) aphids responded to predators by producing winged offspring, but heat shocks strongly inhibited this transgenerational response to predation. Overall, our experimental results show that heat shocks inhibit phenotypic and behavioural responses to predation (and vice versa) and that such changes may alter trophic interactions, and have important consequences on the dynamics and stability of ecological communities. We conclude that the effects of extreme climatic events on the phenotype and behaviour of interacting species should be considered to understand the effects of climate change on species interactions and communities.

Impact of the Phoretic Phase on Reproduction and Damage Caused by Varroa destructor (Anderson and Trueman) to Its Host, the European Honey Bee (Apis mellifera L.).

Varroa destructor is a parasitic mite of the honeybee that causes thousands of colony losses worldwide. The parasite cycle is composed of a phoretic and a reproductive phase. During the former, mites stay on adult bees, mostly on nurses, to feed on hemolymph. During the latter, the parasites enter brood cells and reproduce. We investigated if the type of bees on which Varroa stays during the phoretic phase and if the duration of this stay influenced the reproductive success of the parasite and the damage caused to bees. For that purpose, we used an in vitro rearing method developed in our laboratory to assess egg laying rate and the presence and number of fully molted daughters. The expression level of two Varroa vitellogenin genes (VdVg1 and VdVg2), known to vary throughout reproduction, was also quantified. Results showed that the status of the bees or time spent during the phoretic phase impacts neither reproduction parameters nor the Varroa vitellogenin genes levels of expression. However, we correlated these parameters to the gene expression and demonstrated that daughters expressed the vitellogenin genes at lower levels than their mother. Regarding the damage to bees, the data indicated that a longer stay on adult bees during the phoretic phase resulted in more frequent physical deformity in newborn bees. We showed that those mites carry more viral loads of the Deformed Wing Virus and hence trigger more frequently overt infections. This study provides new perspectives towards a better understanding of the Varroa-honeybee interactions.

The smell of change: warming affects species interactions mediated by chemical information.

Knowledge of how temperature influences an organism's physiology and behaviour is of paramount importance for understanding and predicting the impacts of climate change on species' interactions. While the behaviour of many organisms is driven by chemical information on which they rely on to detect resources, conspecifics, natural enemies and competitors, the effects of temperature on infochemical-mediated interactions remain largely unexplored. Here, we experimentally show that temperature strongly influences the emission of infochemicals by ladybeetle larvae, which, in turn, modifies the oviposition behaviour of conspecific females. Temperature also directly affects female perception of infochemicals and their oviposition behaviour. Our results suggest that temperature-mediated effects on chemical communication can influence flows across system boundaries (e.g. immigration and emigration) and thus alter the dynamics and stability of ecological networks. We therefore argue that investigating the effects of temperature on chemical communication is a crucial step towards a better understanding of the functioning of ecological communities facing rapid environmental changes.

Towards a mechanistic understanding of temperature and enrichment effects on species interaction strength, omnivory and food-web structure.

Revealing the links between species functional traits, interaction strength and food-web structure is of paramount importance for understanding and predicting the relationships between food-web diversity and stability in a rapidly changing world. However, little is known about the interactive effects of environmental perturbations on individual species, trophic interactions and ecosystem functioning. Here, we combined modelling and laboratory experiments to investigate the effects of warming and enrichment on a terrestrial tritrophic system. We found that the food-web structure is highly variable and switches between exploitative competition and omnivory depending on the effects of temperature and enrichment on foraging behaviour and species interaction strength. Our model contributes to identifying the mechanisms that explain how environmental effects cascade through the food web and influence its topology. We conclude that considering environmental factors and flexible food-web structure is crucial to improve our ability to predict the impacts of global changes on ecosystem diversity and stability.

The role of semiochemicals in short-range location of aggregation sites in Adalia bipunctata (Coleoptera, Coccinellidae).

To survive unfavorable periods, ladybird beetles form conspicuous aggregations in specific microsites, with these locations remaining the same year after year. This constancy of location leads to the hypothesis that semiochemicals are involved in the attraction and aggregation of ladybirds to the microsite. In this study, we identified two types of semiochemicals that could play key roles in the attraction and aggregation formation of the two-spotted ladybird, Adalia bipunctata. We first isolated and identified three alkylmethoxypyrazines from A. bipunctata and tested the behavioral responses of diapausing ladybirds to these chemicals in a four-way olfactometer. This revealed that 2-isobutyl-3-methoxypyrazine, on its own or as part of a two-component mixture with 2-isopropyl-3-methoxypyrazine, elicited a positive behavioral response, causing arrestment of diapausing A. bipunctata. As ladybirds are in contact with each other in aggregations, we investigated the role of cuticular hydrocarbons (CHCs) in driving the cohesion and maintenance of aggregation. When an extract of CHCs from diapausing ladybirds was deposited near an alkylmethoxypyrazine source, ladybirds spent more time in the vicinity of the source. We identified a set of CHCs specific to diapausing A. bipunctata. Alkylmethoxyyrazines and CHCs thus deliver information to diapausing ladybirds searching for an aggregation site, as well as mediating several other behaviors throughout the ladybird's life cycle. Chemical parsimony is discussed.

Effects of simulated heat waves on an experimental plant-herbivore-predator food chain.

Greater climatic variability and extreme climatic events are currently emerging as two of the most important facets of climate change. Predicting the effects of extreme climatic events, such as heat waves, is a major challenge because they may affect both organisms and trophic interactions, leading to complex responses at the community level. In this study, we set up a simple three-level food chain composed of a sweet pepper plant, Capsicum annuum; an aphid, Myzus persicae; and a ladybeetle, Coleomegilla maculata, to explore the consequences of simulated heat waves on organism performance, trophic interactions, and population dynamics. We found that (1) heat waves do not affect plant biomass, significantly reduce the abundance and fecundity of aphids, and slightly affect ladybeetle developmental time and biomass, (2) heat waves decrease the impact of ladybeetles on aphid populations but do not modify the effect of aphids on plant biomass, and (3) food chains including predatory ladybeetles are more resistant to heat waves than a simple plant-aphid association, with aphid abundance being less influenced by heat waves in the presence of C. maculata. Our results suggest that more biodiverse ecosystems with predators exerting a strong biotic control are likely to be less influenced by abiotic factors and then more resistant to extreme climatic events than impoverished ecosystems lacking predators. Our study emphasizes the importance of assessing the effects of climatic change on each trophic level as well as on trophic interactions to further our understanding of the stability, resilience, and resistance of ecological communities under climatic forcing.

Using functional response modeling to investigate the effect of temperature on predator feeding rate and energetic efficiency.

Temperature is one of the most important environmental parameters influencing all the biological processes and functions of poikilothermic organisms. Although extensive research has been carried out to evaluate the effects of temperature on animal life histories and to determine the upper and lower temperature thresholds as well as the optimal temperatures for survival, development, and reproduction, few studies have investigated links between thermal window, metabolism, and trophic interactions such as predation. We developed models and conducted laboratory experiments to investigate how temperature influences predator-prey interaction strengths (i.e., functional response) using a ladybeetle larva feeding on aphid prey. As predicted by the metabolic theory of ecology, we found that handling time exponentially decreases with warming, but--in contrast with this theory--search rate follows a hump-shaped relationship with temperature. An examination of the model reveals that temperature thresholds for predation depend mainly on search rate, suggesting that predation rate is primarily determined by searching activities and secondly by prey handling. In contrast with prior studies, our model shows that per capita short-term predator-prey interaction strengths and predator energetic efficiency (per capita feeding rate relative to metabolism) generally increase with temperature, reach an optimum, and then decrease at higher temperatures. We conclude that integrating the concept of thermal windows in short- and long-term ecological studies would lead to a better understanding of predator-prey population dynamics at thermal limits and allow better predictions of global warming effects on natural ecosystems.

Evolution of cannibalism and female's response to oviposition-deterring pheromone in aphidophagous predators.

1. Egg cannibalism by larvae is common in Coccinellidae and is known to be advantageous for the cannibals. Furthermore, larvae of aphidophagous ladybirds usually produce an oviposition-deterring pheromone (ODP), which inhibits oviposition by adult females. It has been proposed that the response to ODP has evolved because of the high costs of cannibalism. However, this has never been formally proved. 2. In this paper, we study the theoretical evolution of this system. We first look at the conditions under which cannibalism and the response to ODP can evolve. Subsequently, we examine the occurrence of polymorphism both in the production of larval tracks and in the sensitivity of females to specific pheromones. 3. The models predict that the amount of cannibalism should not depend on prey density and that evolution should lead to a continuous increase in cannibalism, and consequently larvae should always cannibalize eggs when possible. In response to the cost of cannibalism, ODP recognition can evolve, so that females avoid laying eggs in patches of prey already occupied by conspecific larvae. The result is an arms race between larvae and adult females, which favours a diversification of ODP pheromones. Our models show that: (i) females should be able to recognize mixtures of hydrocarbons rather than a single molecule; and (ii) females should be more sensitive to the tracks of their own offspring than those of non-related larvae.