Risk of migration: not all gregarious locust nymphs reach appropriate refuges.

Sedentary animals choose appropriate refuges against predators, while migratory ones may not necessarily do so. In ectotherms, refuge selection is critical during low temperatures, because they cannot actively evade predators. To understand how migratory ectotherms alter their defensive behaviors depending on refuge quality in cold temperatures, we evaluated migratory gregarious desert locust nymphs (Schistocerca gregaria) in the Sahara Desert, where daily thermal constraints occur. We recorded how roosting plant type (bush/shrub) and its height influenced two alternative defense behaviors (dropping/stationary) during cold mornings, in response to an approaching simulated ground predator. Most locusts in bushes dropped within the bush and hid irrespective of their height, whereas those roosting > 2 m height in shrubs remained stationary. These defenses are effective and match with refuge plant types because dynamic locomotion is not required. When nymphs roosted on shrubs < 1.5-m height, which was an unsafe position, nearly half showed both defensive behaviors, indicating that escaping decisions become ambiguous when the refuges are inappropriate. These results suggest that locusts display flexible defensive behaviors when finding appropriate refuges and selecting refuge before daily thermal limitations occur could be critical for migratory ectotherms, which is a risk associated with migration.

Thermoregulatory behavior of lekking male desert locusts, Schistocerca gregaria, in the Sahara Desert.

Most terrestrial animals are constrained by extreme heat conditions such as midday desert environments, while a few terrestrial ectothermic insects are active in such ecological niches. Sexually mature males of the desert locust (Schistocerca gregaria) in the Sahara Desert remain on the open ground, despite the ground temperatures exceeding their lethal limit, to form leks and to mate incoming gravid females during the daytime. Lekking male locusts apparently suffer from extreme heat stress and greatly fluctuating thermal conditions. The present study examined the thermoregulatory strategies of the lekking male S. gregaria. Our field observations showed that lekking males changed their body orientation toward the sun depending on the temperature and time of day. In the relatively cool morning, males basked by orienting perpendicular to the sun's rays, maximizing the area of body surface exposed to the sun's rays. In contrast, around midday, when the ground surface temperature exceeded lethal high temperatures, some males tended to shelter inside the plants or remain in the shade. However, the remainder stayed on the ground, stilted (i.e., extending their legs to raise their bodies off the hot ground) and oriented parallel to the sun's rays, which minimized radiative heating. Measurements of body temperature throughout the hot middle period of the day confirmed that the stilting posture prevented overheating. Their critical lethal body temperature was as high as 54.7 °C. In this lekking system, gravid females enter male leks by flying. These incoming females usually landed on open ground, whereupon nearby males immediately approached, mounted, and mated the female, implying that males with greater heat-tolerance can increase mating chance. These results suggest that behavioral thermoregulation and physiologically high heat tolerance of male desert locusts allows them to endure extreme thermal conditions for lekking.

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.

A general model of the thermal constraints on the world's most destructive locust, Schistocerca gregaria.

All terrestrial ectotherms are constrained to some degree by their thermal environment and the extent to which they can behaviorally buffer variable thermal conditions. New biophysical modeling methods (NicheMapR) allow the calculation of the body temperature of thermoregulating animals anywhere in the world from first principles, but require detailed observational data for parameterization and testing. Here we describe the thermoregulatory biology of marching bands of the desert locust, Schistocerca gregaria, in the Sahara Desert of Mauritania where extreme heat and strong diurnal fluctuations are a major constraint on activity and physiological processes. Using a thermal infrared camera in the field, we showed that gregarious nymphs altered the microhabitats they used, as well as postural thermoregulatory behaviors, to maintain relatively high body temperature (nearly 40°C). Field and laboratory experiments demonstrated that the preferred body temperature accelerated digestive rates. Migratory bands frequently left foraging sites with full guts before consuming all vegetation and moved to another habitat before emptying their foregut. Thus, the repertoire for behavioral thermoregulation in the desert locust strongly facilitates foraging and digestion rates, which may accelerate developmental rates and increase survival. We used our data to successfully parameterize a general biophysical model of thermoregulatory behavior that could capture hourly body temperature and activity at our remote site using globally available environmental forcing data. This modeling approach provides a stronger basis for forecasting thermal constraints on locust outbreaks under current and future climates.

Daily microhabitat shifting of solitarious-phase Desert locust adults: implications for meaningful population monitoring.

The Desert locust Schistocerca gregaria is a major world pest that causes substantial agricultural and economic damage. Effective pest control relies on effective monitoring, which requires knowledge of locust microhabitat selection. Yet little is known about microhabitat selection of solitarious adult locusts in the field. We conducted field surveys to investigate fine-scale diel temporal and spatial distributions of solitarious adults in the Sahara Desert in Mauritania, a major breeding and recession area. We found that solitarious adults moved among different, specific microhabitats throughout the 24-h period in a cyclical manner. At night, they roosted in trees, moved to the ground to feed shortly after dawn, sheltered in low vegetation during the hot midday, and returned to the ground in the late afternoon. Hence, they switched microhabitats and plant species throughout each day. These cyclical daily movements among diverse microhabitats and specific plant species were correlated with time of day, light intensity, temperature, humidity, and specific plant species, and may relate to anti-predator defence, thermoregulation, and feeding. The present study suggests that locust monitoring should be adjusted, based on time of day, locust age, phase state and relative abundance of specific plant species. For example, we recommend surveying ground after morning and trees at night, for solitarious adults, when at low density.