Interactions between multiple anthropogenic stressors can have unexpected synergistic or antagonistic effects, making it difficult to predict their combined effect using single stressor studies. The interaction between invasive consumers and nutrient enrichment is particularly important as both of these stressors frequently co-occur and their respective bottom-up and top-down effects have the potential to interact across multiple trophic levels. We conducted a mesocosm experiment that crossed an increasing nutrient addition gradient against an increasing zebra mussel invasion gradient. Native zooplankton communities were added to the mesocosms, and after three months we examined how the single stressor effects on available resources and the zooplankton community were altered by their multiple stressor interaction. Added nutrients had no effect on primary producer abundance, but increased the abundance and dominance of the top consumer, which likely increased predation pressure on the producers and so prevented their response to increased nutrients. Zebra mussels reduced total phytoplankton abundance by ~75%, rotifer abundance by ~80%, and shifted communities towards dominance of cladocerans and adult/juvenile copepods. When combined, the top-down control exerted by the mussels interacted antagonistically to prevent any bottom-up influence of nutrient enrichment on the zooplankton community. These results provide insight into the potential outcomes of nutrient and invasive consumer stressor interactions, and illustrate the need for researchers to consider single stressor problems in a multiple stressor context. / Thesis (Master, Biology) -- Queen's University, 2014-01-31 15:20:15.387

Arctic food webs are characterized by multi-year predator-prey cycles. Arctic foxes (Vulpes lagopus) feed primarily on rodents, but also on avian and marine prey when rodents are scarce. I examined temporal variation in the arctic fox diet related to food availability and stress hormones (i.e. cortisol). Lemmings (Dicrostonyx richardsoni), goslings, and goose eggs were important components of the fall and winter diet. Goslings were important in fall, even when rodents were abundant. Lemmings were most important in winter, even when lemming densities were low. Consuming stored eggs did not reduce cortisol concentrations, suggesting that arctic foxes do not prefer stored eggs to lemmings. I also found that prey hormones increased fecal hormone concentrations of captive arctic foxes, introducing an additional caution for hormone studies involving predators. Nonetheless, relationships between stress hormone concentrations and changes in food availability can provide insight regarding the importance of food sources to consumer populations.

A mesocosm experiment with four triplicated treatments (control, clear carbon, pigment and dissolved organic matter) was carried out to investigate the effects of light (by pigment addition) and carbon (C) on phytoplankton communities and on the grazing pressure of microzooplankton on phytoplankton in the Baltic Sea. Phytoplankton concentration and species composition were determined by using an inverted microscope following the Utermöhl method. In order to measure differences in phyto-zooplankton trophic interactions between treatments, a dilution experiment was performed at the end of the mesocosm experiment. Surprisingly, the results show that light attenuation was beneficial to phytoplankton production while carbon enrichment had negative effects on phytoplankton production, the highest phytoplankton production was in the low light treatment and lowest phytoplankton production was in the clear C treatment. Cyanobacteria were the most dominant group, representing over 95% of the phytoplankton community. Diatoms were the least dominant group, representing less than 5% of the phytoplankton community. Microzooplankton grazing pressure was affected by light attenuation and C enrichment. Grazing pressure was highest on cyanobacteria in the clear C treatment, but chrysophytes experienced low grazing pressure and had high growth rates in the Control treatment where microzooplankton fed selectively on chrysophytes. The least abundant group, diatoms, decreased although they experienced no grazing pressure in the Control and clear C treatments. This experiment resulted in two surprising findings. First, increased light intensity reduced phytoplankton biomass, especially in the clear C treatment, and second, cyanobacteria constituted an important food source for microzooplankton grazers. This study adds to the increasing evidence that light effects can be counterintuitive and that cyanobacteria are not necessarily grazing resistant and can meditate nutrient transport to higher trophic levels.

The aim of this thesis was to identify general structuring mechanisms in benthic food webs within the framework of ecological stoichiometry theory. Ecological stoichiometry is defined as the balance of multiple chemical substances in ecological interactions and explicitly considers the combined dynamics of key elements such as carbon (C), nitrogen (N) and phosphorus (P). Ecological stoichiometry theory was developed for pelagic environments, thus it must be tested whether the same mechanisms are applicable to benthic environments. In this thesis, ecological stoichiometry theory was used as a framework to investigate nutrient pathways in benthic littoral ecosystems. I conducted one invertebrate field sampling and six experiments. In the experiments, factors such as grazing, light, nutrients and fish presence were manipulated. The results showed that stoichiometric variability in consumers could mostly be explained by taxa. However, there was some stoichiometric variability due to sampling season, site, and nutrient enrichment. Grazing mostly increased periphyton N and P content, although nutrient recycling effects were dependent on grazer stoichiometry. Grazing changed benthic algal community composition by increasing the proportion of grazing resistant algae species. Additionally, grazing decreased algal diversity, especially under nutrient poor conditions. The manipulation of fish presence revealed that fish affected primary producer biomass and stoichiometry through nutrient recycling. The manipulation of abiotic factors, such as light and nutrient addition could affect periphyton nutrient content, biomass and benthic algal chlorophyll a content. The separate addition of N or P led to an increase of the added nutrient in the periphyton. Increased light intensities led to a decreased cellular chlorophyll a content and increased C:nutrient ratios. This thesis arrives at the conclusion that periphyton-grazer-predator interactions in the benthic are bound by stoichiometric constraints. Nutrient recycling by benthic invertebrates and fish are important mechanism in benthic littoral ecosystems.

La question de la coexistence entre espèces est cruciale dans le contexte des changements globaux, où certaines espèces colonisent de nouveaux écosystèmes, tel le cas des espèces invasives. Dans les habitats discontinus, la coexistence entre espèces invasives et natives gagne à être considérée à la lumière de la théorie des métacommunautés, et en particulier en examinant séparément les processus de dispersion et les interactions locales afin de déterminer leurs effets relatifs. Les approches observationnelles en milieux naturels permettent par ailleurs la prise en compte de la complexité des relations possibles. C’est le travail abordé dans cette thèse à travers l’exemple de l’effet de l’invasion de l’écrevisse de Louisiane sur les amphibiens natifs dans des réseaux de mares. La densité des larves et la distribution des amphibiens ont montré la conccurrence avec l’écrevisse dans les mares. Grâce à la génétique du paysage, la dispersion de l’écrevisse s’est révélée fortement contrainte par le caractère discontinu des habitats, contrairement à celle des amphibiens, bien qu’elle ait été variable parmi les trois espèces étudiées. Les analyses d’isotopes stables ont montré que malgré sa position centrale dans les réseaux trophiques, l’écrevisse n’avait pas de fortes interactions trophiques avec les amphibiens, tout au plus un effet indirect par la modification de l’habitat. Sur la base des mécanismes considérés ici, la coexistence entre les amphibiens natifs et l’écrevisse invasive semble ainsi envisageable. Des suites de ce travail sont envisageables à l’échelle des communautés grâce à l’ADN environnemental. / The question of species coexistence is crucial in the context of global changes, where some species colonize new ecosystems, such as the case of invasive species. In patchy habitats, the coexistence between invasive and native species gains at being considered in the light of the metacommunity theory, and particularly by examining separately the processes of dispersal and local interactions to determine their relative effects. Besides, observational approaches in natural habitats allow to account for the complexity of the possible relationships. It is the work tackled in this thesis through the example of the effect of the red swamp crayfish on native amphibians in pond networks. The density of larvae and the distribution of native amphibians showed the co-occurrence with the crayfish in ponds. Owing to landscape genetics, the dispersal of crayfish was shown to be highly constrained by the patchy nature of habitats, contrary to that of amphibians, although it was variable among the three species considered. The stable isotope analyses showed that, despite its central position in the food webs, the crayfish did not have strong trophic interactions with amphibians, all the more an indirect effect through habitat modification. On the basis of the mechanisms considered here, the coexistence between native amphibians and the invasive crayfish seem possible. Perspectives from this work are envisaged at the community scale using environmental DNA.

Bark beetles regulate forest succession by removing weakened or stressed trees and exposing understory species to light from canopy gaps. Subterranean termites are predominate decomposers of coarse woody debris in southern pine forests; however, little is known about their role in forest health and succession. Both groups of insects rely heavily on fungal symbioses to fill their respective ecological niches in southern pine forests. During recent inspections of southern pine timber, we observed that trees in the early stages of bark beetle attack often had subterranean termites in blue-stained portions of the trunk. The frequency of subterranean termite presence in blue-stained areas of trees increased proportionally to the stage of bark beetle attack. However, practically no research has undertaken the challenge of describing how woody resources created by bark beetles are identified and utilized by subterranean termites before any signs of stress are visible. Therefore, this study examined possible facilitative interactions between subterranean termites, bark beetles and their blue-stain fungal associates, and other invertebrates, and investigated the effect of blue-stain fungi on surface properties of wood. Both native (Reticulitermes spp.) and Formosan subterranean termites exhibited a higher feeding preference for blue-stained sapwood than for unstained sapwood in laboratory assays. Native subterranean termites also consumed blue-stained sapwood at a higher rate than decayed wood. This study was the first to demonstrate that wood containing a non-decay fungus could elicit a feeding response from subterranean termites greater than that observed for decayed wood. Additionally, the surface properties of bark beetle-attacked southern pine were initially reduced by blue-stain fungal infection; however, the process of kiln-drying reversed this effect, resulting in a surface that was more conducive to wood product manufacturing.

Les terres noires (Histosol) du Sud du Québec représentent une ressource agricole non-renouvelable où l’on produit la majorité des laitues (Lactuca sativa L.) au Canada. Les paillis de seigle font partie des pratiques culturales durables proposées pour les protéger. Les impacts de ces paillis sur l’agroécosystème demeurent méconnus. Cette étude visait à mesurer les impacts des paillis de seigle sur la chaîne trophique associée au puceron de la laitue, Nasonovia ribisnigri (Mosley, 1841), un ravageur important. Au cours d’une saison de croissance et avec deux transplants, les impacts des paillis de seigle ont été examinés in situ. Dépistages visuels et pièges-bols jaunes ont mesuré la colonisation des plants par N. ribisnigri, ses ennemis naturels et leurs proies alternatives. En cages d’exclusion, la valeur adaptative et la croissance des populations de pucerons ont été mesurées. Ex situ, les effets résiduels des paillis dans le sol sur le fitness et la croissance des populations de N. ribisnigri ont été mesurés. In situ, les paillis de seigle ont limité la colonisation des laitues par N. ribisnigri et ont recruté d’avantage d’ennemis naturels et de proies alternative sur un des transplants. Le paillis a réduit le fitness des pucerons ainsi que la concentration en acides aminés foliaires et le poids des laitues en début de saison. Ex situ, des effets résiduels du paillis de seigle furent détectés sur la composition du microbiote du sol mais pas sur le fitness des pucerons. Ces résultats démontrent le potentiel des paillis de seigle comme pratique culturale contre N. ribisnigri mais n’éclaircissent pas les mécanismes à l’œuvre et soulignent un risque quant au rendement de la laitue. / Southern Quebec cultivated peatland (Histosols) are a non-renewable resource producing most Canadian lettuce (Lactuca sativa L.). Rye mulches are one of the conservation practices proposed to reach sustainable production, but their impacts on the agroecosystem are still poorly studied. We aimed to assess impacts of rye mulches on the trophic chain around the lettuce aphid Nasonovia ribisnigri (Mosley 1841), an economically significant pest. Through one growing season and two successive transplants, we monitored impacts of rye mulches in situ. Visual scouting and yellow pan traps assessed plant colonization by N. ribisnigri, its natural enemies and alternative preys. Exclusion cage experiments measured aphid fitness and population growth. Ex situ, we attempted to detect if lingering effects of the rye mulch within the soil affected lettuce plants, N. ribisnigri and soil microbiota. In situ experiments showed that rye mulch had the potential to inhibit N. ribisnigri field colonization and recruited natural enemies and alternative preys in one transplant. Mulch reduced aphid fitness as well as lettuce foliar amino acids concentration and weight at the beginning of the season. Ex situ experiments detected lasting effect of rye mulch on microbiota composition but not on aphid nor lettuce fitness after mulch removal. This study highlights the potential of rye mulches as a cultural practice against N. ribisnigri but does not reveal mechanisms at play while underlining agronomic challenges concerning lettuce yields.

The ecology and evolution of a species is often considered only within the context of pairwise interactions even though a species' distribution and abundance may be determined by interactions with many species within and between trophic levels. Multiple herbivores often share the same host and may interact indirectly by altering the relationships between herbivores, their host plants and their parasitoids. However, the relationships between parasitoids and herbivore hosts have typically been studied in isolation of other herbivore species. I examined how the outcomes of species interactions change when multiple relationships are considered. Chapter 1 examined the potentially conflicting selection pressures Manduca sexta exerts on Nicotiana tabacum (tobacco), since M. sexta has pollinating adults but herbivorous larvae. I demonstrated that high nectar amino acids do not affect floral visitation, but increased oviposition of herbivores on leaves. Thus, the relative costs and benefits of nectar rewards may depend on the community of pollinators and their life histories. In the remaining chapters I examined how feeding on tobacco by the aphid Myzus persicae altered the interactions between a parasitoid (Cotesia congregata) and its hornworm host (M. sexta). Chapter 2 demonstrated that aphids reduced hornworm abundance and parasitism. Changes in hornworm abundance were not due to density-dependent changes in moth oviposition, but the proportion of caterpillars attacked by parasitoids was inversely density dependent with hornworm density. Chapter 3 examined whether changes in hornworm abundance and parasitism reflected aphid-induced changes in host plant quality or volatile emissions. Aphids increased hornworm mortality, did not affect parasitoid performance, and increased parasitoid search time. In combination with Chapter 2, results suggest that aphids can mediate parasitoid-caterpillar interactions through changes in host plants that reduce hornworm survival and alter parasitoid behavior. Chapter 4 addressed how the outcome of interactions that are altered by aphids changed with spatial scale, and found no effect spatial scale on hornworm abundance and parasitism. In this system, aphids alter parasitoid-caterpillar interactions through changes in plant quality that reduce hornworm performance and abundance, and in turn, influence parasitoid attack. This work demonstrates that the outcome of multispecies interactions may not be predictable from pairwise interactions. / Ph. D.

A presente pesquisa teve como objetivos aprimorar a amostragem de Spodoptera frugiperda através do uso de armadilhas com feromônio sexual, verificar a capacidade de dispersão de Trichogramma atopovirilia como agente de controle biológico desta praga bem como avaliar a atratividade de Amaranthus sp. às fêmeas de T. atopovirilia. Para aprimorar a amostragem de S. frugiperda, os estudos foram conduzidos em dois campos comerciais de produção de milho, sendo um composto de 36,3 ha (denominado de MIP e com 30 pontos de amostragem) e outro de 10 ha (denominado de convencional e com 10 pontos de amostragem). Nestes campos foram instaladas armadilhas com feromônio sexual, distribuídas de forma aleatória e na proporção de 1 armadilha / ha. As avaliações foram realizadas duas vezes por semana durante todo o desenvolvimento das plantas, considerando-se: número de adultos machos coletados pelas armadilhas, injúria dos insetos às plantas, número de posturas e de larvas (pequenas, médias e grandes). O número de adultos coletados nas armadilhas foi correlacionado com os níveis de injúrias, número de posturas e densidade larval. O número de larvas observadas não apresentou diferença entre áreas e pontos de amostragem. A injúria dos insetos às plantas, número de posturas e de larvas pequenas não apresentaram correlação com os adultos capturados pelas armadilhas. Houve correlação significativa entre o número de larvas grandes (4o e 5o ínstares) e o número de plantas apresentando o cartucho furado ou destruído. Não houve correlação da coleta de adultos nas armadilhas com a infestação ou níveis de injúria, demonstrando que as larvas ainda precisam ser contadas para determinar o momento do seu controle. A capacidade de dispersão de T. atopovirilia foi estudada em 3 fases distintas de desenvolvimento das plantas de milho, quando estas possuíam de 4 a 6 folhas, 8 a 10 folhas e pendoamento. Em cada fase foi realizada uma infestação artificial com posturas (até 24 h) de S. frugiperda nas plantas localizadas em distâncias que variaram de 6 a 24 m do ponto de liberação do parasitóide. Logo após esta infestação, foi realizada somente uma liberação de adultos do parasitóide para cada fase da cultura e sempre no período mais fresco do dia (manhã ou entardecer). O parasitismo foi permitido durante até 48 horas quando as posturas eram recolhidas e acondicionadas em câmara climatizada até a emergência dos adultos. Os resultados indicaram que as plantas em suas diferentes fases de desenvolvimento afetaram a capacidade de dispersão dos parasitóides. As posturas localizadas nos pontos mais distantes somente foram atingidas pelos parasitóides durante a fase de desenvolvimento da cultura em que as plantas de milho estavam menores (4 a 6 folhas). Nas outras fases, os índices de parasitismo foram maiores naquelas posturas localizadas mais próximas do ponto de liberação. A atratividade de quatro espécies de Amaranthus (A. retroflexus, A. viridis, A. hybridus e A. spinosus) às fêmeas de T. atopovirilia foram avaliadas utilizando-se olfatômetro Peterson. Estas plantas foram testadas em período de desenvolvimento vegetativo e reprodutivo. Adotou-se o delineamento inteiramente casualizado com 40 repetições (considerando-se um inseto/ repetição) e cada inseto foi observado durante o tempo máximo de 600 segundos. As espécies mais atrativas foram A. viridis e A. retroflexus, nos estádios vegetativo e reprodutivo, respectivamente. Quando testadas simultaneamente, as espécies de plantas não demonstraram atratividade ao parasitóide. Estes resultados sugerem a viabilidade do uso destas plantas ou seus derivados no manejo de habitats de hospedeiros e aumento de parasitismo em programas de manejo integrado de pragas. / The objectives of this research were the improvement of cross-section Spodoptera frugiperda using pheromone traps to verify the capacity of dispersion of Trichogramma atopovirilia as an agent for biological pest control as well as evaluating the attractiveness of Amaranthus sp. to T. atopovirilia females. To improve cross-section of S. frugiperda, the studies were carried out in two commercial corn field plantations, one made up of 36,3 ha (called MIP and with 30 points of sampling) and another of 10 ha (called conventional and with 10 points of sampling). In these fields pheromone traps were randomly displaced (ratio of 1 trap/ha). The evaluations were realized twice a week during the whole plant development, considering : number of male adults collected by the traps, insect injury to plants, egg masses and larval densities (small, medium and large). The numbers of adults collected were correlated to the insect injury, egg masses and larval density. The larval occurrence did not show difference between areas and sampling points. The injury levels, egg masses and small larva densities did not show correlation to adult caught by traps. There was significant correlation between large larvae (4th and 5th instars) and the number of plants showing whorl holes and/or destroyed ones. There were no correlations observed among adults caught by traps and insect infestation or injury, so demonstrating that larvae should be counted for timing pest control. The capacity of dispersion of T. atopovirilia was studied in 3 distinct phases of corn plant development, when they had 4 to 6 leaves, 8 to 10 leaves and tassel. In each stage was applied an artificial infestation with egg masses (< 24 h) of S. frugiperda on plants located 6 to 24 m from a parasitoid release point. Right after this artificial infestation, was applied only one parasitoid adult release to each distinct corn plant phase , always early in the morning or late in the afternoon. The parasite contamination was allowed during 48 h and then the egg masses were retrieved and taken into a climatic chamber until emerging adulthood. The results showed that plants affected the parasitoid dispersal. The egg masses placed farthest from the release point were parasited only during the 4 to 6-leaf stage. In the others stages, only egg masses next to the release point were parasited. The attractiveness of four species of Amaranthus (A. retroflexus, A. viridis, A. hybridus and A. spinosus) to the females of T. atopovirilia was evaluated by a Petersons olfactometer. These plants were evaluated in both vegetative and reproductive stages. Complete randomized procedure was adopted with 40 replications (considering one insect/replication) and each insect was observed during 600 seconds. The most attractive species were A. viridis and A. retroflexus during vegetative and reproductive stage, respectively. When plants were tested simultaneously, the species did not show attractive properties to the parasitoids. These results suggest the viability of the use of these plants or its derivatives to change host habitat and parasite increase for biological control in integrated pest management.

A agregação em insetos centra-se nas vantagens relativas ao forrageio e no aumento da defesa contra predadores. Além disso, agregações de ovos podem beneficiar-se pelo aumento nas taxas de eclosão larval. Neste trabalho, foram testadas as hipóteses de que agregações de Ascia monuste orseis (Godart, 1819) (Lepidoptera: Pieridae) uma espécie que se alimenta em agregações larvais em plantas da família Brassicaceae - conferem benefícios à performance larval e adulta, na assimilação de alimento pelas larvas, na viabilidade dos ovos e na proteção larval contra inimigos naturais. A possibilidade de um dado comportamento do grupo resultar em maior ataque por inimigos naturais também foi avaliada, bem como um padrão comportamental defensivo em resposta aos inimigos naturais. Para tanto, quatro tratamentos de diferentes tamanhos de agregações larvais foram formados (1, 7, 15 e 30 larvas) para avaliar a performance e a assimilação de alimento em laboratório. Em campo, três tratamentos foram formados (1, 10 e 50 larvas) para testar o efeito do tamanho do tamanho do grupo na predação e parasitismo. A performance do estágio de ovo foi examinada em posturas de tamanhos variados em casa de vegetação. Larvas gregárias desenvolveram-se mais rápido nos ínstares iniciais e tornaram-se fêmeas mais fecundas em comparação às solitárias; entretanto, larvas solitárias apresentaram maior tamanho do que as gregárias. A sobrevivência não diferiu entre os tratamentos em laboratório. Foi observado um menor consumo per capita de alimento por larvas gregárias, sem custos para a assimilação de alimento. A viabilidade dos ovos aumentou com o tamanho da agregação de ovos, comprovando o benefício da agregação larval na fase de ovo. A menor predação per capita em agregações larvais maiores conferiu uma maior proteção às larvas de A. monuste orseis contra predadores e parasitoides, através do efeito da diluição do ataque entre os indivíduos do grupo. O parasitoidismo foi mais expressivo em larvas de primeiros ínstares, enquanto que larvas mais tardias foram mais atacadas por predadores, independente do tamanho da agregação. Por fim, eventos comportamentais que envolvem movimentação da cabeça como exploração e alimentação foram mais perigosos para as larvas de A. monuste orseis em comparação ao repouso e deslocamento, semelhantemente a espécies de hábito solitário. Eventos comportamentais supostamente defensivos foram observados em todos os ínstares e tratamentos. As vantagens da agregação em A. monuste orseis mostraram-se especialmente importantes no estágio de ovo e primeiros ínstares, pela diminuição da mortalidade de ovos e vulnerabilidade larval aos inimigos naturais. Esses benefícios provavelmente sobrepõem-se aos custos, como a competição por interferência observada entre as larvas no final do desenvolvimento. Nossos resultados mostram que o malogro dos ovos e os efeitos dos inimigos naturais constituem fortes pressões seletivas na manutenção da agregação de ovos e larval em A. monuste orseis, a qual confere uma melhor performance do ponto de vista bi-trófico, bem como maior probabilidade de sobrevivência individual sob o ponto de vista tri-trófico. / In insects, the gregarious habit has been shown to improve foraging and defense against predation to both larval and adult stages. Egg clusters could also be beneficial through increased larval hatching. In this study, we tested the hypothesis that egg clusters and larval aggregations of the neotropical butterfly Ascia monuste orseis (Godart, 1819) (Pieridae) - a subspecies that uses Brassicaceae as hosts - lead to several benefits from both bi- and tri-trophic perspectives. Larval and adult performances, food assimilation by the larvae, egg viability, as well as protection against natural enemies were assessed from individuals reared either isolated or aggregated. The behavior of larval aggregations was also examined with respect to predation risk, as well as the corresponding larval defensive behaviors after enemy attack. Four treatments with different larval aggregation sizes were assigned (1, 7, 15 e 30 larvae) to assess larval performance and food utilization in the laboratory. In the field, three treatments were assigned (1, 10 e 50 larvae) to evaluate the effects of group size on predation and parasitoidism. Egg performance was examined through egg clusters of different sizes in a greenhouse. Gregarious larvae developed faster, especially in early instars, and became more fecund females than solitary larvae; however, the latter attained larger body size than the former. Under laboratory conditions, survival did not differ among treatments. Lower food ingestion per capita was observed in gregarious larvae, with no cost in food assimilation. The viability of eggs increased as egg aggregation size increased. The lower per capita predation in larger larval aggregations than smaller groups conferred higher protection to A. monuste orseis larvae against natural enemies, through the dilution effects among individuals of the group. Parasitoidism was more intense in small-sized larvae while late instars were more susceptible to predators, regardless of aggregation size. Similar to species with solitary habit, behavioral events which involved head movements as searching and feeding were more dangerous to A. monuste orseis larvae compared to resting and walking. Presumed defensive behaviors were observed in all instars and treatments. Thus, the benefits of aggregation in A. monuste orseis can be seen especially in the egg stage and in first instars, as it reduces egg mortality and larval vulnerability to natural enemies. These benefits probably overcome some costs, such as interference competition in the late instars. Taken together, the results show that egg failure and top-down effects constitute selective pressures in maintaining egg and larval aggregation in A. monuste orseis, by providing better performance from a bi-trophic perspective and increased probability of individual survival from a tri-trophic perspective when compared to solitary individuals.