Pheromones serve key functions in the biology of swarming locusts. However, research has focused largely on the mass-swarming desert locust, Schistocerca gregaria. We extended these investigations to the pheromonal profile of the rarely swarming American bird grasshopper, S. americana (Drury). The headspace of mature gregarious S. americana males contained three characteristic electroantennogram-active components: (Z)-3-nonen-1-ol, (Z)-2-octen-1-ol, and nonanal. These substances were accompanied by aromatics such as phenol that are also released by females and immatures. Male-specific pheromone components were released independently from epidermal gland cells, with the highest emission rate being for (Z)-3-nonen-1-ol from the abdomen and legs. The emission of the major compound, (Z)-3-nonen-1-ol, is stress-sensitive, and coincides with sexual maturity and crowding. The emission pattern strongly supports a role of (Z)-3-nonen-1-ol in the reproductive biology of S. americana. The pheromone is involved in courtship-inhibition and is used as mate assessment pheromone in cryptic female choice. In double mating experiments, females choose sperm of males with high (Z)-3-nonen-1-ol emission. Furthermore the pheromone accelerated maturation of immature adults and supports synchronization of sexual development.
Living in high-density groups of animals has advantages and disadvantages for mating. The advantage of facilitated mate finding is compromised by difficulties in protecting a suitable partner from competitors. Thus, males regularly are faced with increased competition for sperm, and females with harassment by males at high population densities. To cope with these problems, mating tactics and mate choice mechanisms have to be adjusted. An adaptation to gregarious condition observed in locusts includes the use of male-emitted pheromones. Males of the Central American locust, Schistocerca piceifrons, release sex-specific volatiles, which were identified as phenethyl alcohol (synonym: phenyl-ethyl-alcohol, 2-phenyl-1-ethanol, 2-phenylethanol, PEA), (Z)-3-nonen-1-ol (3-Nol), and (Z)-2-octen-1-ol (2-Ool). The emission of the two major compounds, PEA and 3-Nol, was restricted to crowded conditions. Furthermore, the release of both volatiles was coupled to males reaching sexual maturity, indicating a function in reproductive behavior. However, neither the single substances nor their mixtures were attractive or repellent to the locusts. Instead, females prefer the sperm of high pheromone-emitting males to fertilize their ova. In this way, the male-specific volatiles act as mate assessment pheromones utilized in a context of cryptic female choice. This function is well supported by the highly variable but individual-specific emission rates of the three compounds. Schistocerca piceifrons males release a virtually unique personal pheromone signature, a prerequisite for mate assessment pheromones.
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