Aggregation in an heterospecific population of blowfly larvae: social behaviour is impacted by species-specific thermal requirements and settlement order.

Larvae of several blowfly species grow on carcasses and actively aggregate together. They face harsh developmental conditions resulting in a strong pressure to reduce development time: this is achieved either through thermoregulation or aggregation. We investigate how these two developmental strategies are modulated within heterospecific groups. In a first experiment, larvae of two species with different thermal requirements were deposited simultaneously on a thermal gradient. This resulted in the formation of two monospecific groups, each located at the species-specific thermal preferendum. However, when Calliphora vomitoria (Linnaeus) larvae were placed first, the later arriving Lucilia sericata (Meigen) larvae attracted the whole group to its own thermal preferendum. In the reverse experiment, half of the replicates resulted in single dense heterospecific groups observed at temperatures ranging from C. vomitoria to L. sericata preferendum. The other half of the replicates resulted in loose groups spread out on the thermal gradient. These results highlight the emergence of collective decisions ranging from thermal optimization to heterospecific aggregation at suboptimal temperatures. They demonstrate that species settlement order strongly affects self-organization processes and mixed-species group formation. We conclude that thermal optimization and heterospecific niche construction are two developmental strategies of carrion fly larvae. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

The maggot, the ethologist and the forensic entomologist: Sociality and thermoregulation in necrophagous larvae.

Necrophagous insects are mostly known through forensic entomology. Indeed, experimental data investigating the effect of temperature on larval development underlies post-mortem interval estimations. However, such developmental studies rarely considered the behavior of maggots. In contrast, previous results supposed that calliphoridae larvae use behavioral strategies to optimize their development on carcasses. To test this idea, we analyzed the trade-off between thermal regulation (individual thermal preferences) and social behavior (aggregation) in Lucilia sericata larvae. The first set of experiments analyzed the behavior of third instars in response to thermal changes in their environment. The results demonstrated a clear thermoregulation behavior, supporting the assumption that larvae continuously move to reach a suitable internal temperature. The second set of experiments focused on the trade-off between thermal optimization and aggregation. The results showed a constant search for congeners and an attractiveness of aggregates, sometimes to the detriment of thermal optimization. Together, these results demonstrate a balance between behavioral thermoregulation and social strategies, two significant mechanisms for developmental optimization in necrophagous larvae. In conclusion, these findings highlights unexpected (social) strategies to cope with ephemeral resource and high selection pressure. They also raise important questions for forensic entomology.

Mixed-species aggregations in arthropods.

This review offers the first synthesis of the research on mixed-species groupings of arthropods and highlights the behavioral and evolutionary questions raised by such behavior. Mixed-species groups are commonly found in mammals and birds. Such groups are also observed in a large range of arthropod taxa independent of their level of sociality. Several examples are presented to highlight the mechanisms underlying such groupings, particularly the evidence for phylogenetic proximity between members that promotes cross-species recognition. The advantages offered by such aggregates are described and discussed. These advantages can be attributed to the increase in group size and could be identical to those of nonmixed groupings, but competition-cooperation dynamics might also be involved, and such effects may differ between homo- and heterospecific groups. We discuss three extreme cases of interspecific recognition that are likely involved in mixed-species groups as vectors for cross-species aggregation: tolerance behavior between two social species, one-way mechanism in which one species is attractive to others and two-way mechanism of mutual attraction. As shown in this review, the study of mixed-species groups offers biologists an interesting way to explore the frontiers of cooperation-competition, including the process of sympatric speciation.

To eat or get heat: Behavioral trade-offs between thermoregulation and feeding in gregarious necrophagous larvae.

The thermoregulation behavior of Lucilia sericata larvae (Diptera: Calliphoridae), a necrophagous species that feeds on vertebrate cadavers, was investigated. These larvae require high heat incomes to develop, and can elevate temperatures by forming large aggregates. We hypothesized that L. sericata larvae should continue to feed at temperatures up to 38 °C, which can be reached inside larval masses. Thermal regulation behavior such as movement between a hot food spot and colder areas was also postulated. The hypotheses were tested by tracking for 1 h the activity of single, starved third instar larvae in a Petri dish containing 1 food spot (FS) that was heated to a constant temperature of 25 °C, 34 °C or 38 °C with an ambient temperature of 25 °C. The influence of previous conspecific activity in the food on larval behavior was also tested. The crops of larvae were dissected to monitor food content in the digestive systems. Based on relative crop measurements, larvae fed at all food temperatures, but temperature strongly affected larval behavior and kinematics. The total time spent by larvae in FS and the duration of each stay decreased at high FS temperature. Previous activity of conspecifics in the food slightly increased the time spent by larvae in FS and also decreased the average distance to FS. Therefore, necrophagous L. sericata larvae likely thermoregulate during normal feeding activities by adjusting to local fluctuations in temperature, particularly inside maggot masses. By maintaining a steady internal body temperature, larvae likely reduce their development time.

[Thermoregulation behavior in necrophagous dipteran larvae]. / Les larves de diptères nécrophages en entomologie médico-légale : une histoire de température - Prix médecine/sciences.

The forensic entomology is the use of insects to date the death. The forensic expert assessment is based on the development of necrophagous insects which are growing on the cadaver, to calculate their age and then estimate the Post-Mortem Interval. This development depends on a number of parameters like temperature, species or behavior. The French Forensic Taphonomy unit, the only expert team on the subject in France, works on the biology, physiology and ethology of the necrophagous insects. Their works are focused on thermoregulation behavior and thermal preferendum of maggot masses, aggregation phenomenon and social interaction or on food intake. These works are particularly of interest to understand the pre-social parameters and evolution strategies. More importantly, their aim is to better understand the development of necrophagous insects and, in fine, to improve the forensic expert assessment.

Effects of the environment on bone mass: A human taphonomic study.

An experiment was designed using human ribs placed in different environments to document how moisture and temperature affect the bone mass according to the postmortem interval. The bones were defleshed, weighed and partially buried, with some ribs being left unburied as controls. The ribs were weighed daily, and the mass loss was monitored over a period of 90days. The results showed that significant differences in bone mass loss exist between environments, where the bone mass loss was significantly faster in an environment with low moisture content. This mass loss is thought to be primarily associated with the desiccation of the body and then for a greater part, with the atmospheric moisture content. However, the loss of bone mass can also be explained by early alterations in the organo-mineral matrix which were highlighted by Raman spectroscopy method.

Experimental study of Lucilia sericata (Diptera Calliphoridae) larval development on rat cadavers: Effects of climate and chemical contamination.

Household products such as bleach, gasoline or hydrochloric acid have been used to mask the presence of a cadaver or to prevent the colonization of insects. These types of chemicals affect insect development and alter the forensic entomology analysis. This study was designed to test the effects of six household products (bleach, mosquito repellent, perfume, caustic soda, insecticide and unleaded gasoline) on blowfly (Lucilia sericata, Diptera: Calliphoridae) larval development. Furthermore, the effects of climate (rain or dry conditions) on larval development were analyzed. For each replication, 100 first instars were placed on a rat cadaver on which one household product was spilled. We observed a decrease in the survival rates of the larvae but no significant effect on their development times or the adult size. The same trends were observed under rainy conditions. However, the rain altered the effects of some tested household products, especially gasoline. These results demonstrate for the first time the successful development of necrophagous larvae on chemically contaminated cadavers, and provide evidence for the range of possible effects to expect.

In vitro effects of household products on Calliphoridae larvae development: implication for forensic entomology.

Several parameters can delay the first arrival of flies on a corpse and the subsequent development of the larvae. This study focuses on the development of Lucilia sericata (Diptera: Calliphoridae) (Meigen, 1826) on household chemical-contaminated substrates. bleach, perfume, hydrochloric acid, caustic soda, insecticide, mosquito repellent, and gasoline in quantities consistent with an amount that could possibly be spilled on a corpse were mixed with beef liver to simulate contaminated fleshes. Larvae were bred at 25 °C on these media until emergence. Four developmental parameters were followed: survival rates, development times, sex ratios, and adult sizes. Hydrochloric acid, insecticide, and gasoline killed all larvae. In low quantities, caustic soda and mosquito repellent increased the development time and decreased the adult size. However, high quantities of these chemicals killed all larvae. Lastly, bleach and perfume did not affect the survival rate and barely impacted the development time or adult size. These results demonstrate common household products spilled on a corpse can strongly affect the development of Calliphoridae larvae. The effects of such products should be considered in forensic entomology cases.