Allocation of more reproductive resource to egg size rather than clutch size of gregarious desert locust (Schistocerca gregaria) through increasing oogenesis period and oosorption rate.

The desert locust, Schistocerca gregaria, shows a density-dependent reproductive trade-off by laying fewer but larger eggs in crowded conditions (gregarious phase) than in isolated conditions (solitarious phase). However, the physiological mechanisms controlling reproductive resource allocation remain unclear. We examined how egg production processes, including ovulation timing (i.e., oogenesis period), oocyte and ovarian growth rates, and oosorption rate (resorbing developing terminal oocytes), regulate reproductive outputs (egg biomass per clutch, egg size, and clutch size) during a reproductive cycle in S. gregaria by rearing them either under isolated or crowded conditions. We observed a common density-dependent negative correlation between egg size and clutch size, with no significant difference in egg biomass between the two rearing conditions. Dissection of female locusts after different days of oviposition revealed that the daily oocyte growth rate was almost similar between the two rearing conditions, but crowd-reared females ovulated later than isolated-reared ones, resulting in further oocyte growth in the former. Terminal oocytes were renewed by previous penultimate oocytes at the onset of a new reproductive cycle, and oosorption mainly occurred at an early stage in both rearing conditions; however, crowd-reared locusts displayed higher levels of oosorption compared to their isolated-reared counterparts. Crowding induced a high oosorption rate, resulting in a reduced clutch size and a prolonged oogenesis period, which in turn allowed oocytes to intake more yolk, which was probably recycled via oosorption. These results suggest that the length of the oogenesis period and oosorption rate are manipulated by maternal density, and these physiological modifications interactively regulate reproductive trade-off in S. gregaria.

Density-dependent mating behaviors reduce male mating harassment in locusts.

Male mating harassment may occur when females and males do not have the same mating objectives. Communal animals need to manage the costs of male mating harassment. Here, we demonstrate how desert locusts in dense populations reduce such conflicts through behaviors. In transient populations (of solitarious morphology but gregarious behavior), we found that nongravid females occupied separate sites far from males and were not mating, whereas males aggregated on open ground (leks), waiting for gravid females to enter the lekking sites. Once a male mounted a gravid female, no other males attacked the pair; mating pairs were thereby protected during the vulnerable time of oviposition. In comparison, solitarious locusts displayed a balanced sex ratio in low-density populations, and females mated irrespective of their ovarian state. Our results indicate that the mating behaviors of desert locusts are density dependent and that sex-biased behavioral group separation may minimize the costs of male mating harassment and competition.

The desert locust, Schistocerca gregaria, plastically manipulates egg size by regulating both egg numbers and production rate according to population density.

Egg-size adjustment is one of the important plastic life-history traits for animals living in heterogeneous environments. The adaptive investment hypothesis predicts that mothers should increase progeny size according to certain cues predicting adverse future conditions of their offspring. However, reproductive resources are limited, and females have to simultaneously reduce egg number to allocate more resources to increase size. It remains unclear how single individuals alter egg size and number according to temporally heterogeneous environments. In the present study, we examined how desert locusts, Schistocerca gregaria, plastically alter egg size and number according to population density. We also investigated the trans-generational maternal effects on progeny characteristics as well as their own maternal physiological response (oviposition interval). Females kept in crowded conditions laid significantly larger and heavier eggs by reducing clutch size (number of eggs per egg pod) compared to isolated females, suggesting the existence of a reproductive trade-off between the two traits. The crowding-forced isolated females induced concerted changes not only in egg size but also in egg number tending towards those characteristics of gregarious control, implying that single individuals showed trade-off when egg size was increased. Double-blind testing confirmed the rapid crowding effects on egg size. Females also responded to crowding by extending the oviposition interval. As the oviposition interval extended, egg size increased, but clutch size decreased. Eggs from crowding-forced isolated females began to produce gregarious-phase type hatchlings (large and black) instead of solitarious-phase type ones (small and green). These results suggested that S. gregaria plastically manipulate egg size by regulating egg numbers and egg production rate, and indicated the presence of trans-generational maternal effects on progeny phase.