14Insects regulate their body temperature mostly behaviorally, by changing posture or 15 microhabitat. These strategies may be ineffective in some habitats, for example on carrion. 16 Carrion beetles create a biofilm-like matrix by applying to cadaver surface anal or oral 17 exudates. We tested the hypothesis that biofilm formed by communally breeding Necrodes 18 littoralis L. beetles (Silphidae) produces heat, that enhances beetle fitness. We demonstrated 19 that heat produced in the biofilm is larger than in meat decomposing without insects. Beetles 20 regularly warmed up in the biofilm. Moreover, we provide evidence that biofilm formation by 21 adult beetles has deferred thermal effects for larval microhabitat. We found an increase in heat 22 production of a biofilm and a decrease in development time and mortality of larvae, after 23 adult beetles applied their exudates on meat. Behavioral strategy revealed here for N. littoralis 24 is basically a new form of thermoregulation and parental care in insects. 25 26 Keywords 27 Insect thermoregulation; Animal behavior; Parental care; Carrion ecology; Insect-microbe 28 interactions; Resource competition 29 30 31Temperature is a key component of animal environment. Insects usually use external 32 heat to regulate their body temperature (1, 2). By changing body orientation or selecting 33 microhabitat with specific thermal characteristics, insects may maintain their body 34 temperature within thermal optima (3, 4). These mechanisms may be ineffective at certain 35 developmental stages (e.g. larvae) and in some microhabitats (e.g. carrion or dung), indicating 36 that other thermal options may be important for some insect ectotherms.
37Carrion is an example of a "bonanza" resource, i.e. very rich but at the same time 38 scattered and ephemeral (5). There is severe competition between microbes, insects, and 39 vertebrates over carrion resources (6)(7)(8)(9)(10)(11)(12). Insects, e.g. blow flies (Calliphoridae) or carrion 40 beetles (Silphidae) use carrion for breeding and their larvae are main carrion reducers in 41 terrestrial environments (11, 13, 14). Necrophagous larvae usually feed in aggregations (15).
42Larval aggregates on carrion may have much higher inner temperature than ambient air (by 43 10-30°C), an effect originally discovered in aggregations of larval blow flies (4, 16-20). Heat 44 in these aggregates was hypothesized to derive from microbial activity (21, 22), larval 45 exothermic digestive processes (19) or larval frenetic movements (20, 23). However, the 46 specific mechanisms involved are not known. 47 Carrion beetles, in particular burying beetles (Silphidae: Nicrophorus), create a 48 biofilm-like matrix on cadavers by applying to its surface anal and oral exudates (24). In case 49 of Nicrophorus the behavior was hypothesized to moisturize carrion (25), facilitate digestion 50 (25-27), suppress microbial competitors (24, 28-31), deter insect competitors by reducing 51 carrion-originating attractants (25, 32, 33), support larval aggregation (25) or...