abstract: Of all the signals and cues that orchestrate the activities of a social insect colony, the reproductives' fertility pheromones are perhaps the most fundamental. These pheromones regulate reproductive division of labor, a defining characteristic of eusociality. Despite their critical role, reproductive fertility pheromones are not evenly expressed across the development of a social insect colony and may even be absent in the earliest colony stages. In the ant Camponotus floridanus, queens of incipient colonies do not produce the cuticular hydrocarbons that serve as fertility and egg-marking signals in this species. My dissertation investigates the consequences of the dramatic change in the quantity of these pheromones that occurs as the colony grows. C. floridanus workers from large, established colonies use egg surface hydrocarbons to discriminate among eggs. Eggs with surface hydrocarbons typical of eggs laid by established queens are nurtured, whereas eggs lacking these signals (i.e., eggs laid by workers and incipient queens) are destroyed. I characterized how workers from incipient colonies responded to eggs lacking queen fertility hydrocarbons. I found that established-queen-laid eggs, incipient-queen-laid eggs, and worker-laid eggs were not destroyed by workers at this colony stage. Destruction of worker-laid eggs is a form of policing, and theoretical models predict that policing should be strongest in incipient colonies. Since there was no evidence of policing by egg-eating in incipient C. floridanus colonies, I searched for evidence of another policing mechanism at this colony stage. Finding none, I discuss reasons why policing behavior may not be expressed in incipient colonies. I then considered the mechanism that accounts for the change in workers' response to eggs. By manipulating ants' egg experience and testing their egg-policing decisions, I found that ants use a combination of learned and innate criteria to discriminate between targets of care and destruction. Finally, I investigated how the increasing strength of queen-fertility hydrocarbons affects nestmate recognition, which also relies on cuticular hydrocarbons. I found that queens with strong fertility hydrocarbons can be transferred between established colonies without aggression, but they cannot be introduced into incipient colonies. Queens from incipient colonies cannot be transferred into incipient or established colonies. / Dissertation/Thesis / Ph.D. Biology 2012

A integridade das sociedades de insetos depende da capacidade dos indivíduos em discriminar aqueles que não pertencem às suas colônias; essas interações de reconhecimento são mediadas por hidrocarbonetos presentes na camada de cera da cutícula e atuam como mensageiros que codificam informação sobre o meio interno (colônia) e externo (ambiente). Os hidrocarbonetos são produzidos por células dérmicas e o perfil desses compostos recebe influência tanto de fontes endógenas quanto exógenas e isso o torna suscetível às mudanças temporais. A caracterização desses compostos em insetos-praga é importante, pois provêm subsídios tanto para pesquisa básica como para o manejo e controle de pragas. Nesse contexto, o presente estudo abordou as variações dos hidrocarbonetos cuticulares em relação às subcastas morfológicas das operárias (condição endógena) e utilização do substrato foliar (variável exógena) na formiga-cortadeira Atta sexdens, conhecida como uma das principais pragas herbívoras da região Neotropical. Os compostos foram extraídos em solvente apolar hexano e as amostras foram analisadas em sistema de cromatografia a gás acoplado à espectrometria de massas (CG-MS). Foram identificados 30 hidrocarbonetos com cadeia carbônica entre 19 e 40 átomos distribuídos em três classes de compostos, sendo a dos hidrocarbonetos ramificados a mais representativa, especificamente os trimetialcanos. Foram encontradas variações significativas para todas as variáveis analisadas, sugerindo que as subcastas possuem assinaturas químicas semelhantes quanto à diversidade de compostos, mas com grandes diferenças na proporção relativa dos mesmos. Em relação ao substrato foliar, colônias que tiveram acesso ao mesmo recurso apresentam uma composição química cuticular mais semelhante em relação àquelas que tiveram acesso a um recurso diferente. Complementarmente, os testes comportamentais mostraram que, na presença de uma coespecífica intrusa, as operárias residentes são muito mais agressivas em relação àquelas que utilizaram outro tipo de substrato foliar. / The integrity of insect societies relies on the ability of individuals to discriminate between nesmates from non-nestmates. These interactions are mediated by hydrocarbons spread all over the cuticle that act as messengers coding information about the colony and external environment. Cuticular hydrocarbons are produced by dermic cells and its composition is influenced by both endogenous and exogenous sources which makes them subject to temporal changes. Thus, this study approached the variations on this chemical profile in relation to the worker subcastes and type of foliar substrate used by leafcutter ant Atta sexdens, this species is known as one of the major neotropical herbivore pests and this kind of study is important because it can provide subsides in researches related to both basic biology as to pest control methods. The compounds were extracted using apolar solvent (hexane) and the samples were analyzed using gas chromatography - mass spectrometry technique (GC-MS). It was found 30 hydrocarbons which carbon chains varies between 19 and 40 atoms separated in three classes of compounds and the branched hydrocarbons as the most abundant one, especially the trimethyalcanes. The statistical analysis revealed significant differences on all the variables analyzed suggesting that subcastes possess similar chemical signatures as to the variety of compounds but with great differences in relative proportions thereof. Furthermore, the results show that foliar substrate has influence on the composition of cuticular hydrocarbons and concomitantly affect the interspecific recognition, thus colonies that had access to the same substrate has similar cuticular composition comparing to those who do not. Additionally, behavior tests showed that in the presence of a conspecific intruder the resident workers tend to be much more aggressive in relation to those who had fed on a different foliar substrate.

Orientadores: Woodruff Whitman Benson, Jose Roberto Trigo / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-06T14:03:47Z (GMT). No. of bitstreams: 1 Dall'Aglio-Holvercem_ChristianeGoncalves_D.pdf: 1459643 bytes, checksum: 5ed521529fb7303e231df7d9f2ebbda3 (MD5) Previous issue date: 2006 / Resumo: Este trabalho discute alguns aspectos da ecologia da formiga lava-pés Solenopsis invicta e seus parasitóides, as moscas do gênero Pseudacteon (familia Phoridae), e da taxonomia de S. invicta e S. saevissima, ambas nativas da América do Sul. A alometria de colônias de S. invicta no sudeste do Brasil foi estudada através de medições das dimensões do ninho acima do solo, seguidas por escavação e medida da profundidade, biomassa e número de operárias da colônia, assim como da largura média da cabeça das operárias. Análise de regressão dos dados para 28 colônias forneceu novas relações alométricas que podem ser usadas para estimar o volume do ninho (acima e abaixo da superfície do solo), a biomassa de formigas, e o número de operárias a partir do volume do ninho acima do solo. A biomassa de formigas por unidade de volume do ninho, o número de operarias por unidade de volume do ninho, o peso médio por operária, e a largura média da cabeça das operárias foram essencialmente independentes do tamanho da colônia. Em contraste, estas quatro quantidades aumentam com o tamanho da colônia entre colônias de S. invicta na América do Norte. O ciclo sazonal e abundância de moscas parasitóides do gênero Pseudacteon na vizinhança de colônias de S. invicta no sudeste do Brasil foi observado ao longo de um período de 3 anos em li áreas de gramíneas nos arredores de Campinas. O número médio N de forídeos atraídos para um ninho de formigas lava-pés ao longo de um período padrão de observação (15 min) foi usado como uma medida da abundância de forídeos. As dimensões do ninho e algumas variáveis meteorológicas foram também registradas em cada ninho de formigas lava-pés. Pseudacteon tricuspis foi a espécie mais comum (70.4%), seguido por P. curvatus (23.8%), P. obtusus (1.9%) e P.litoralis (3.9%). As características mais notáveis das flutuações sazonais da abundância de Pseudacteon são picos abruptos com duração aproximada de um mês (valores máximos de N entre 4 e 10 forídeos/ninho), que ocorreram em abril e junho de alguns anos; para comparação, valores típicos de N estavam entre 0.2 e 2 forídeos/ninho. As populações de forídeos parecem ser destruídas por incêndios e inundações, mas recuperam-se em uma escala de tempo da ordem de meses. Nenhuma correlação significativa foi encontrada entre a abundância de Pseudacteon e variáveis meteoro lógicas, densidade, ou volume de ninhos de S. invicta, sugerindo que a abundância de Pseudacteon não está diretamente ligada a abundância de S. invicta. Flutuações quase idênticas em abundância de Pseudacteon foram observadas ao longo de um período de 6 meses em duas áreas de estudo separadas por cerca de 3 km, o que pode ser causado por dispersão de forídeos e conseqüente uniformização de densidades sobre áreas de vários quilômetros quadrados. Os números de forídeos atraídos pelos vários ninhos de S. invicta incluídos neste estudo seguiram uma distribuição de Poisson em baixas abundâncias de Pseudacteon (N < 0,5 forídeos/ninho); a distribuição tornava-se mais e mais contagiosa com o aumento da abundância de Pseudacteon (até o máximo observado, de N = 10 forídeoslninho). Esta observação sugere que a atratividade de um ninho de formigas lava-pés aos parasitóides aumenta com o número de forídeos que foram previamente atraídos para o ninho. Critérios quantitativos eficientes foram desenvolvidos para distinguir entre S. invicta e S. saevissima, que são morfológicamente muito similares e cujas distribuições geográficas na América do Sul se sobrepõe significativamente. Estes critérios resultaram da análise de 68 amostras de formigas lava-pés de 9 localidades espalhadas pelo estado de São Paulo (sudeste do Brasil) através de cromatografia gasosa e espectrometria de massa. 22 hidrocarbonetos cuticulares e 18 alcalóides piperidínicos foram identificados nas amostras estudadas, e suas abundâncias relativas foram analisadas utilizando análises de componentes principais e de funções discriminantes. Estas análises mostraram uma separacão nítida entre as duas espécies em um plano com certas coordenadas (P,Q), que são funções das abundâncias relativas dos hidrocarbonetos cuticulares. As amostras provenientes de uma das localidades (São José do Rio Preto) têm uma composição de hidrocarbonetos que é claramente diferente de todas as outras amostras consideradas neste estudo, mas também apresentam algumas caracteristicas em comum com cada uma das duas espécies. Uma possibilidade é que esta população constitua um hibrido de S. invicta e S. saevissima. As duas espécies também exibem uma separação razoavelmente nítida em um plano com coordenadas (R$), que são funções das abundâncias relativas de alcalóides piperidínicos. Os resultados das identificações de espécies baseados nestes critérios de separação quimica concordam com os resultados baseados em um critério morfológico, a presença ou ausência do dente clipeal mediano. A população "anômala" de São José do Rio Preto é morfologicamente classificada como S. invicta / Abstract: This work discusses some aspects of the ecology of the fire ant species Solenopsis invicta and its phorid parasitoid flies ofthe genus Pseudacteon, and ofthe taxonomy of S. invicta and S. saevissima, both of which are native to South America. The allometry of S. invicta colonies in southeast Brazil was studied by measurement of mound (aboveground) dimensions, followed by excavation and measurement of the colony depth, biomass, number of workers, and mean worker head width. Regression anaIysis of data for 28 colonies yielded new allometric relationships which can be used to estimate nest volume (above and below ground), ant biomass, and number ofworkers ITom mound (aboveground) volume. Ant biomass per unit nest volume, number of workers per unit nest volume, mean worker weight, and mean worker head width were nearly independent of colony size. In contrast, these four quantities increase with colony size among S. invicta colonies in North America. The seasonal cycle of abundance of Pseudacteon phorid flies in the vicinity of S. invicta colonies in southeast Brazil was observed over a 3-year period in 11 grass-covered areas in the outskirts of Campinas. The average number N of phorids attracted to a fire ant nest over a standard observation period (15 min) was used as a measure of phorid abundance. Mound dimensions and meteorological variables were also recorded at each fIre ant nest. Pseudacteon tricuspis was the most common species (84.2%), followed by P. curvatus (12.6%), P. obtusus (2.7%) e P. litoralis (0.5%). The most remarkabIe featores of the seasonal fluctuations of Pseudacteon abundance are sharp month-Iong peaks (peak N values between 4 and 10 phorids/nest) which occurred in Abril and June of some years; for comparison, at other times N was typically between 0.2 and 2 phorids/nest. Phorid populations seem to be destroyed by rifes and floods, but recover on a time scale of months. No significant correlation was found between Pseudacteon abundance and meteorological variables, density, or volume of S. invicta nests, suggesting that Pseudacteon abundance is not determined by, and does not determine, the abundance of S. invicta. Almost identical fluctuations in Pseudacteon abundance were observed over a 6-month period in two study areas separated by about 3 km, which may be due to phorid dispersaI and consequent density uniformization over areas of several square kilorneters. The nurnbers of phorids attracted by the various S. invicta nests inc1uded in this study followed a Poisson distribution at 10w Pseudacteon abundances (N < 0.5 phorids/nest), becoming more and more contagious at high Pseudacteon abundances (up to N= 10 phorids/nest). This observation suggests that the attractiveness oftire ant nests to phorid parasitoids increase with the nurnbeer ofphorids which have been previously attracted to the nest. Efficient quantitative criteria were developed for distinguishing between S. invicta and S. saevissima, which are morphologically very similar and whose geographical ranges in South Arnerica significantly overlap. These criteria resulted from the analysis of 68 tire ant samples from 9 localities across São Paulo state (southeast Brazil) via gas chromatography/mass spectrometry. 22 cuticular hydrocarbons and 18 piperidine alkaloids were identified in the studied samples, and their relative abundances were analyzed using principal component and discrirninant function analyses. These analyses showed a sharp separation between the two species in a plane with certain coordinates (P,Q), which are functions of the relative abundances of cuticular hydrocarbons. The fire ant samples from one of the localities (São José do Rio Preto) had a hydrocarbon composition which was markedly different from ali other samples considered in this study, but had some features in common with each ofthe two species. One possibility is that this population constitutes a hybrid of S. invicta and S. saevissima. The two species also show a reasonably sharp separation in a plane with coordinates (R,S), which are functions of the relative abundances of piperidine alkaloids. The species identification results based on these chernical separation criteria agree with those based on a morphological criterion, the presence or absence of the median clypeal tooth. The "anomalous" population from São José do Rio Preto is rnorphologically c1assified as S. invicta / Doutorado / Ecologia / Doutor em Ecologia

Os hidrocarbonetos cuticulares têm importante papel no processo de reconhecimento dos membros da colônia de insetos sociais. Muitos estudos têm mostrado variações qualitativas e quantitativas nestes compostos entre os insetos adultos. Contudo, abordagens referentes à modulação do perfil destes compostos durante a formação da cutícula são escassas, e se restringem aos estágios larval de holometábolos e de ninfas de hemimetábolos. O principal objetivo dessa pesquisa foi caracterizar o perfil de hidrocarbonetos cuticulares e a expressão de genes potencialmente relacionados à sua biossíntese durante o processo de formação e maturação da cutícula adulta. Os perfis de hidrocarbonetos foram caracterizados por meio de GC/MS e mostraram diferenças quantitativas marcantes que significativamente discriminaram as cutículas pupal, adulta-farata e adulta. Em paralelo, sequências de enzimas que catalisam a desaturação (desaturases) ou elongação (elongases) de lipídeos, disponíveis no banco de dados do NCBI, foram utilizadas para o desenho de primers e estudo da expressão gênica por meio de RT-qPCR. Cinco genes de desaturases, e oito genes de elongases mostraram variação de expressão estatisticamente significante no tegumento de abelhas adultas em comparação com pupas e adultas-faratas. Testes de correlação entre os perfis de expressão gênica e de hidrocarbonetos cuticulares evidenciaram os genes potencialmente envolvidos com a biossíntese destes compostos para a formação e maturação da cutícula. Estes resultados corroboram a hipótese de que nos insetos sociais, a cutícula só amadurece completamente por ocasião do início da atividade de forrageamento. Associando estes dados a análises de evolução molecular das desaturases e elongases, pudemos sugerir as etapas da via de síntese de hidrocarbonetos catalisadas por estas enzimas, e assim eleger genes candidatos a futuro silenciamento mediado por RNA de interferência para pesquisa de função. / Cuticular hydrocarbons are important for recognition of nestmates in social insect colonies. Many studies have shown qualitative and quantitative variations in the cuticular hydrocarbons between adult insects. However, approaches on developmental profiles of these compounds during cuticle formation and differentiation are scarce, and restricted to larval stages of holometabolous and nymphs of hemimetabolous. The main objective of this work was to characterize the cuticular hydrocarbons profiles and the expression of genes potentially involved in the biosynthesis of these compounds during the synthesis and differentiation of the adult cuticle in the honeybee. The hydrocarbons profiles were characterized using GC/MS and showed remarkable quantitative differences, thus discriminating the pupal, pharate-adult and adult cuticles from each other. In parallel, we used annotated sequences of enzymes catalyzing lipid desaturation (desaturases) or elongation (elongases), available in NCBI data bank, for primers design and gene expression analysis using RT-qPCR. Five desaturase genes and eight elongase genes showed statistically significant expression changes in the integument of adult bees in comparison to pupae and pharate-adults. Correlation tests supported roles of some of the desaturase and elongase genes in hydrocarbons biosynthesis for incorporation into adult cuticle. In addition, these results go along with the hypothesis that in social insects the cuticle is just completed when the insect starts forager activity. Taken together, these data and an analysis on the molecular evolution of desaturases and elongases allowed suggesting the steps in the pathway of cuticular hydrocarbons biosynthesis that are catalyzed by these enzymes, and also allowed to elect candidate genes for further functional studies using gene silencing mediated by RNAi.

Cuticular hydrocarbons (CHCs) are found on the outer cuticle of all terrestrial arthropods. Although their primary function is in desiccation prevention, these compounds have also been shown to play a variety of roles in insect chemical communication, from species and sex recognition to providing cues of dominance and attractiveness. However, despite growing evidence of their versatility as cues, our knowledge of how CHCs are used in mating interactions is limited to Drosophila and field crickets. In this thesis I investigate the roles CHCs play in interactions at each stage of the mating process in the broad-horned flour beetle Gnatocerus cornutus. I assess the relative importance of CHCs in influencing male reproductive success and examine the complex interplay between different episodes of selection and the mechanisms of sexual selection acting on males. I use a combination of behavioural assays, experimental manipulations and gas chromatography. First, I identify the role of CHCs as cues of sperm competition risk and intensity, demonstrating how the presence of male-derived CHCs on the cuticles of virgin females elicits males to adjust their pre- and post-copulatory investment (chapter 2), by providing information on the state of their competitive environment. I then go on to look at the stability of CHCs as cues of sperm competition over time, finding that they are highly sensitive to environmental degradation (chapter 3) and do not persist in the habitat substrate of this species. Next, I investigate how male CHCs determine fighting and mating success. By estimating and comparing the strength and form of sexual selection imposed by male-male competition and female mate choice, I show that male CHCs are subject to strong antagonistic sexual selection (chapter 4). By experimentally manipulating male CHC profile, I then attempt to verify the selection gradients estimated for female choice 3 (chapter 5). However, my experimental manipulation fails to verify the importance of male CHCs for female mate choice. Finally, I explore the role of same-sex sexual behaviour (SSB) in determining male reproductive success (chapter 6). I find evidence to suggest that SSB may in fact be a form of aggression in its own right, and demonstrate that SSB and fighting may provide equivalent means for males to overcome female choice and secure a mating advantage. My results indicate that CHCs play key roles as chemical cues throughout the mating process and significantly impact male reproductive success. My thesis reveals the intricate nature of the relationships between mechanisms of sexual selection, alongside highlighting the need to consider both the social and physical environment when investigating the importance of chemical cues. I discuss the implications of these results for the evolution of male CHCs and how my findings can be used to further our knowledge of this field.

Analysis of cuticular wax from Kalanchoe daigremontiana leaves was performed to identify the constituent components within the wax, determine how these changed during leaf ontogenesis, and discover how they were distributed within the cuticle. Analysis of extracted cuticular wax by gas chromatography, mass spectrometry, and comparison with authentic standards led to the identification of triterpenoids including glutinol, friedelin, germanicol, epifriedelanol, glutinol acetate and β-amyrin as well as very long chain fatty acid (VLCFA) derivatives including alkanes, primary alcohols, aldehydes, fatty acids, and alkyl esters. Cuticular wax composition in young K. daigremontiana leaves was dominated by triterpenoids, which made up over 70% of the lipid soluble compounds. During leaf ontogenesis, wax composition changed to include a higher proportion of VLCFA derivatives, which made up approximately 50% of cuticular wax in mature leaves. The most abundant triterpenoids in the wax were glutinol and friedelin, both fairly uncommon pentacyclic triterpenoids with a complex proposed biosynthetic mechanism. Tritriacontane (C33 alkane) was the most abundant compound within the VLCFA derivatives. Cuticular wax accumulation was found to correspond well to leaf growth, with both processes slowing at the same time. Variations in the ratio of friedelin-like compounds to glutinol-like compounds during leaf ontogenesis suggest the presence more than one active triterpenoid synthase enzyme in the leaves of K. daigremontiana. VLCFA compounds were found mainly in the epicuticular wax on both the adaxial and abaxial surfaces, while triterpenoids were relatively more abundant in the intracuticular layer. Two different epicuticular wax crystal forms were observed by scanning electron microscopy (SEM) which can be described as platelets with sinuate margins and twisted ribbons. Based on SEM and chemical data as well as previous reports of crystal composition, it is hypothesized that each crystal type has a unique composition, with the platelets containing one or more triterpenoids and the twisted ribbons containing alkanes and other VLCFA derivatives. Confirmation of this hypothesis will have to await further investigation. This research provides information that will aid in the larger goals of characterizing a glutinol or friedelin synthase and understanding the gradients established within epicuticular and intracuticular wax layers. / Science, Faculty of / Chemistry, Department of / Graduate

Directional selection is prevalent in nature yet phenotypes tend to remain relatively constant, suggesting a limit to trait evolution. The genetic basis of evolutionary limits in unmanipulated populations, however, is generally not known. Given widespread pleiotropy, opposing selection on a focal trait may arise from the effects of the underlying alleles on other fitness components, generating net stabilizing selection on trait genetic variance and thus limiting evolution. Here, I look for the signature of stabilizing selection for a suite of cuticular hydrocarbons (CHCs) in Drosophila serrata. Despite strong directional sexual selection on CHCs, genetic variance differed between high and low fitness individuals and was greater among the low fitness males for seven of eight CHCs. Univariate tests of a difference in genetic variance were non-significant but have low power. My results implicate stabilizing selection, arising through pleiotropy, in generating a genetic limit to the evolution of CHCs in this species.

Chemical substances are transferred between bumblebees of the species Bombus terrestris during their mating behavior. These substances are present mainly on the cuticle. The aim of this thesis was to identify and quantitatify differences in the amount of the compounds among defined groups of bumblebees - mated, non-mated and attempting to mate. Cuticular extracts were obtained by washing of the whole bumblebees bodies in hexane. These extracts were analyzed by using a two-dimensional gas chromatography with the time of fly mass spectrometr (GCxGC-TOF/MS). Cuticular hydrocarbons were identified and semi-quantified. Double bounds locations were determined in heptacosene and nonacosene. Five compounds were differed in the examined groups - β-springene, α-springene, dihydrofarnesyl-oleate, palmityl-palmitate and palmityl- palmitoleate. By comparing of the amount of these compounds differences between defined groups of bumblebees were determined. Analysis of the cutilar extracts showed that mated and non-mated bumblebees can be distinguished. Furthermore, there were statistically significant differences in the amounts of identified compounds in all defined groups of bumblebees. Key words: two-dimensional gas chromatography, Bombus terrestris, cutilar extract

No description available.

Conflicts over reproduction are common in animal societies and they are especially pronounced in groups of totipotent individuals. Workers in the drywood termite Cryptotermes secundus are totipotent. They can gain direct fitness via dispersal as winged sexual or inheritance of the natal nest as neotenic replacement reproductive. In this study we examined the actual conflict behaviour in C. secundus, possible mechanism of conflict-regulation, and factors influencing the conflict intensity. Cuticular hydrocarbons provided the information about nestmates which is required for mechanisms of conflict-resolution to work: they were (i) caste-specific, (ii) honest signal of fertility in neotenic queens, as they are reflecting the underlying JH titres mediated via caste-specific Neofem4 expression, and (iii) may also be informative enough to allow kin discrimination. Kin discrimination in C. secundus depended on social context and the individuals’ developmental potential. Sterile soldiers for example showed nepotistic grooming independent from the social context, but affected by wood resource, in contrast to workers that did not react to varying relatedness. Individuals showed distinct behavioural profiles before and during conflict, reflecting their reproductive potential: most prominent in the distinction were the dominance behaviour butting and proctodeal trophallaxis. The proposed role of trophallaxis as honest signal and inhibitory means in regulating reproductive development in C. secundus could be confirmed. Conflict intensity was greatly influenced by wood resource: workers were more likely to stay and fight for inheritance within nests that are resource rich, while when food gets limited dispersal conflict was most pronounced. Thus, individuals did continually assess the ecological (resources) and societal conditions (presence of reproductives, relatedness) and adjusted their developmental decisions in order to maximize own fitness. The developmental potential is linked to the moulting interval, as only individuals in the sensitive phase are able to react to a changing situation like orphaning. Thus the sensitive phase, besides honest signalling (via cuticular hydrocarbon profiles and trophallaxis) might be an further mechanism potentially regulating conflict in C. secundus, as it restricts the number of individuals capable of becoming neotenic in a ‘fair lottery’ process.