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Termites are social insects that live in colonies headed by reproductive castes. The breeding system is defined by the number of reproductive individuals in a colony and the castes to which they belong. There is tremendous variation in the breeding system of termites both within and among species. The current state of our understanding of termite breeding systems is reviewed. Most termite colonies are founded by a primary (alate-derived) king and queen who mate and produce the other colony members. In some species, colonies continue throughout their life span as simple families headed by the original king and queen. In others, the primary king and queen are replaced by numerous neotenic (nymph- or worker-derived) reproductives, or less commonly primary reproductives, that are descendants of the original founding pair leading to inbreeding in the colony. In still others, colonies can have multiple unrelated reproductives due to either founding the colonies as groups or through colony fusion. More recently, parthenogenetic reproduction has shown to be important in some termite species and may be widespread. A major challenge in termite biology is to understand the ecological and evolutionary factors driving the variation in termite breeding systems.

Collective behaviors of social insects are often regulated by pheromones. In subterranean termites, some workers forage for and exploit decaying wood for new food resources while forming tunnels from their nest. Colonizing new food resources requires workers to build and disinfect tunnels and chambers inside the nest and ingest decaying wood; therefore subterranean termite colonies should have mechanisms to establish and maintain groups of workers to perform these functions. Recently, an aggregation pheromone was identified in workers of the termite Reticulitermes speratus , which induces quick attraction of nestmate workers and prolonged aggregation to the site of attraction. In this study, we extended this work to another species of Reticulitermes and identified a worker aggregation pheromone in the termite R. virginicus . GC–MS analysis and bioassays demonstrated that this pheromone consists of 3-octanone, 3-octanol, and palmitic acid and shows a colony-specific, dose-dependent attractant response but not an arrestant response. Furthermore, these pheromone components were most likely emitted from the surface of the body. This suggests that aggregation pheromone composition and function differ significantly among termite species, even within the same genus. This study advances our understanding of the regulatory mechanisms of termite aggregation behavior.

Royal recognition is a central feature of insect societies, allowing them to maintain the reproductive division of labor and regulate colony demography. Queen recognition has been broadly demonstrated and queen recognition pheromones have been identified in social hymenopterans, and in one termite species. Here we describe behaviors that are elicited in workers and soldiers by neotenic queens and kings of the subterranean termite, Reticulitermes flavipes, and demonstrate the chemical basis for the behavior. Workers and soldiers readily perform a lateral or longitudinal shaking behavior upon antennal contact with queens and kings. When royal cuticular chemicals are transferred to live workers or inert glass dummies, they elicit antennation and shaking in a dose-dependent manner. The striking response to reproductives and their cuticular extracts suggests that royal-specific cuticular compounds act as recognition pheromones and that shaking behavior is a clear and measurable queen and king recognition response in this termite species.

The hallmark of social insects is their caste system: reproduction is primarily monopolized by queens, whereas workers specialize in the other tasks required for colony growth and survival. Pheromones produced by reining queens have long been believed to be the prime factor inhibiting the differentiation of new reproductive individuals. However, there has been very little progress in the chemical identification of such inhibitory pheromones. Here we report the identification of a volatile inhibitory pheromone produced by female neotenics (secondary queens) that acts directly on target individuals to suppress the differentiation of new female neotenics and identify n-butyl-n-butyrate and 2-methyl-1-butanol as the active components of the inhibitory pheromone. An artificial pheromone blend consisting of these two compounds had a strong inhibitory effect similar to live neotenics. Surprisingly, the same two volatiles are also emitted by eggs, playing a role both as an attractant to workers and an inhibitor of reproductive differentiation. This dual production of an inhibitory pheromone by female reproductives and eggs probably reflects the recruitment of an attractant pheromone as an inhibitory pheromone and may provide a mechanism ensuring honest signaling of reproductive status with a tight coupling between fertility and inhibitory power. Identification of a volatile pheromone regulating caste differentiation in a termite provides insights into the functioning of social insect colonies and opens important avenues for elucidating the developmental pathways leading to reproductive and nonreproductive castes.

Background Social insects are among the most serious invasive pests in the world, particularly successful at monopolizing environmental resources to outcompete native species and achieve ecological dominance. The invasive success of some social insects is enhanced by their unicolonial structure, under which the presence of numerous queens and the lack of aggression against non-nestmates allow high worker densities, colony growth, and survival while eliminating intra-specific competition. In this study, we investigated the population genetics, colony structure and levels of aggression in the tawny crazy ant, Nylanderia fulva , which was recently introduced into the United States from South America. Results We found that this species experienced a genetic bottleneck during its invasion lowering its genetic diversity by 60%. Our results show that the introduction of N. fulva is associated with a shift in colony structure. This species exhibits a multicolonial organization in its native range, with colonies clearly separated from one another, whereas it displays a unicolonial system with no clear boundaries among nests in its invasive range. We uncovered an absence of genetic differentiation among populations across the entire invasive range, and a lack of aggressive behaviors towards conspecifics from different nests, even ones separated by several hundreds of kilometers. Conclusions Overall, these results suggest that across its entire invasive range in the U.S.A., this species forms a single supercolony spreading more than 2000 km. In each invasive nest, we found several, up to hundreds, of reproductive queens, each being mated with a single male. The many reproductive queens per nests, together with the free movement of individuals between nests, leads to a relatedness coefficient among nestmate workers close to zero in introduced populations, calling into question the stability of this unicolonial system in which indirect fitness benefits to workers is apparently absent.

The inability to distinguish between species can be a serious problem in groups responsible for pathogen transmission. Culicoides biting midges transmit many pathogenic agents infecting wildlife and livestock. In North America, the C. variipennis species complex contains three currently recognized species, only one of which is a known vector, but limited species-specific characters have hindered vector surveillance. Here, genomic data were used to investigate population structure and genetic differentiation within this species complex. Single nucleotide polymorphism data were generated for 206 individuals originating from 17 locations throughout the United States and Canada. Clustering analyses suggest the occurrence of two additional cryptic species within this complex. All five species were significantly differentiated in both sympatry and allopatry. Evidence of hybridization was detected in three different species pairings indicating incomplete reproductive isolation. Additionally, COI sequences were used to identify the hybrid parentage of these individuals, which illuminated discordance between the divergence of the mitochondrial and nuclear datasets.

Knowledge of the relative influence of population- versus species-level genetic variation is important to understand patterns of phenotypic variation and ecological relationships that exist among and within morphologically indistinguishable cryptic species and subspecies. In the case of cryptic species groups that are pests, such knowledge is also essential for devising effective population management strategies. The globally important crop pest Thrips tabaci is a taxonomically difficult group of putatively cryptic species. This study examines population genetic structure of T. tabaci and reproductive isolation among lineages of this species complex using microsatellite markers and mitochondrial COI sequences. Overall, genetic structure supports T. tabaci as a cryptic species complex, although limited interbreeding occurs between different clonal groups from the same lineage as well as between individuals from different lineages. These results also provide evidence that thelytoky and arrhenotoky are not fixed phenotypes among members of different T. tabaci lineages that have been generally associated with either reproductive mode. Possible biological and ecological factors contributing to these observations are discussed.

Native to eastern Asia, the Formosan subterranean termite Coptotermes formosanus (Shiraki) is recognized as one of the 100 worst invasive pests in the world, with established populations in Japan, Hawaii and the southeastern United States. Despite its importance, the native source(s) of C. formosanus introductions and their invasive pathway out of Asia remain elusive. Using ~22,000 SNPs, we retraced the invasion history of this species through approximate Bayesian computation and assessed the consequences of the invasion on its genetic patterns and demography. We show a complex invasion history, where an initial introduction to Hawaii resulted from two distinct introduction events from eastern Asia and the Hong Kong region. The admixed Hawaiian population subsequently served as the source, through a bridgehead, for one introduction to the southeastern US. A separate introduction event from southcentral China subsequently occurred in Florida showing admixture with the first introduction. Overall, these findings further reinforce the pivotal role of bridgeheads in shaping species distributions in the Anthropocene and illustrate that the global distribution of C. formosanus has been shaped by multiple introductions out of China, which may have prevented and possibly reversed the loss of genetic diversity within its invasive range.Blumenfeld and Eyer et al. retrace the invasion history of the termite Coptotermes formosansus from eastern Asia to the US using approximate Bayesian computation. They find a complex invasion history, with multiple introductions originating from eastern Asia, as well as a bridgehead introduction originating from Hawaii, which appears to have mitigated the loss of genetic diversity within the invasive US range.

One of the major goals of ecology is to understand how co-habiting species partition limited resources. In the eastern U.S., at least three species of Reticulitermes subterranean termites often occur in sympatry; however, little is known about how these species divide food resources. In this study, we characterized the foraging activity of Reticulitermes flavipes (Kollar), R. hageni Banks, and R. virginicus (Banks) across seasons to assess the impact of environmental conditions on resource partitioning. A field site consisting of two grids of wooden monitors was sampled monthly for 28 months. Foraging activity in all three species was correlated with the interaction of temperature and moisture. This correlation was influenced by temperature and moisture approximately equally in R. flavipes, whereas temperature contributed more to the correlation in R. hageni, and moisture contributed more in R. virginicus. These differences caused each species to preferentially forage during specific environmental conditions: R. flavipes continued foraging after high moisture events, R. hageni increased foraging under higher soil moisture, and R. virginicus increased foraging under lower soil temperatures. We attempted to explain these patterns by the species’ physiological limits; however, we found no differences in upper lethal limit, desiccation, or submersion limits across species. These results add to the overall understanding of resource partitioning by emphasizing the ability of multiple species to utilize the same resource under different environmental conditions and raise questions regarding the physiological and/or behavioral mechanisms involved.

The production of royal pheromones by reproductives (queens and kings) enables social insect colonies to allocate individuals into reproductive and non-reproductive roles. In many termite species, nestmates can develop into neotenics when the primary king or queen dies, which then inhibit the production of additional reproductives. This suggests that primary reproductives and neotenics produce royal pheromones. The cuticular hydrocarbon heneicosane was identified as a royal pheromone in Reticulitermes flavipes neotenics. Here, we investigated the presence of this and other cuticular hydrocarbons in primary reproductives and neotenics of this species, and the ontogeny of their production in primary reproductives. Our results revealed that heneicosane was produced by most neotenics, raising the question of whether reproductive status may trigger its production. Neotenics produced six additional cuticular hydrocarbons absent from workers and nymphs. Remarkably, heneicosane and four of these compounds were absent in primary reproductives, and the other two compounds were present in lower quantities. Neotenics therefore have a distinct ‘royal’ blend from primary reproductives, and potentially over-signal their reproductive status. Our results suggest that primary reproductives and neotenics may face different social pressures. Future studies of these pressures should provide a more complete understanding of the mechanisms underlying social regulation in termites.