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"Aposematism is quite simply the correlation between conspicuous signals, such as bright coloration, and prey unprofitabil-ity," Candy Rowe wrote in 2001. But why should some prey become unprofitable in the first place, while others do not?

Unprofitability is difficult to define, and even more difficult to measure. It is certainly contextually defined, because the propensity of an animal to eat something is highly dependent on its level of hunger and its ability to use the prey for energy once eaten. Palatability (i.e., the predator's perception of prey profitability) greatly determines whether the predator will or will not eat the prey. Predator—prey coevolution led predators in part to rely on proximal perception to gauge the prey profitability. In particular, taste sensitivity may well have evolved in predators as an assessment of food toxicity: indeed, predators usually consider toxic chemicals to be distasteful. Some insects have external defenses such as horns, or spines, many of which cause irritation. Such physical defenses may be coupled to venom, as with the irritant hairs of many caterpillars or hymenopteran stingers. These insects may be otherwise perfectly profitable, and some predators evolve ways around the physical defenses, such as bee-eaters that are able to remove a bee's stinger and venom sac. Other insects have passive chemical defenses that predators discover upon consumption, such as chemicals in the hemolymph or sequestration glands of lubber grasshoppers or monarch butterflies. Such insects usually develop extra signals such as powerful smells, at least when handled, to advertise their toxicity before being consumed.

Toxicity is not the only way an insect can be unprofitable to predators. Difficulty in capturing prey (due to fast escape, erratic flight, breakable wings, etc.), or difficulty in handling prey (due to toughness or a hard cuticle) are other ways that insects can bring no net reward to the predators that spend energy chasing them, even if the chase results in the prey being seized. However, multiple unprofitability traits might be important in the evolution of warning signals.

Predators can have three kinds of response to a prey depending on their perception of prey profitability. If consuming a prey leads the predator to be more likely to attack similar prey in the future, perhaps even using the prey's appearance as a search image, the prey is called palatable. In feeding experiments, this usually leads birds to attack nearly 100% of the palatable prey offered. Of course, the predator may satiate after consuming a number of prey, and consequently the propensity to attack may decrease at high prey densities. In contrast, if experienced predators are less likely to attack similar prey, the prey is called unpalatable. Of course, predators' memorizing capacity, and the strength of the prey unpalatability, may all influence how fast information regarding prey is acquired and how long it is retained. However, a distasteful prey will inevitably lower the predators' instantaneous propensity to attack this prey further, an effect analogous to an immediate satiation. Finally, eating the prey may have no effect on the predator's subsequent behavior, which means that the prey is effectively neutral. This category is mainly derived from theory; there is little evidence that it exists in nature.

Variations in unpalatability among prey species, along what is called the "(un)palatability spectrum," affect the rate at which predators modify their behavior with experience. Predator's perceived toxicity is likely to be a sigmoid function of actual toxin concentration per unit prey mass, meaning that little of the palatability spectrum may fall into intermediate perceptions between "unpalatable" and "fully palatable." Although how predators learn is still under debate, experiments and theory suggest that they respond to a large extent to the (perceived) concentration of nasty chemicals they can tolerate per unit time.

The distastefulness of insects is generally linked to the host plants they utilize. Indeed, many distasteful or defended insects are herbivorous; most defended nonherbivorous insects are Hymenoptera. Some plant families, like the Solanaceae and the Passifloraceae, which are hosts to many chemically defended insects, contain alkaloids and cyanogens, respectively, as secondary metabolites. Some insects, like monarch butterflies (Danausplexippus) that feed on Asclepias plants (milkweeds, Asclepiadaceae), sequester the compounds of such plants and store them; these insects thus avoid the toxic effects of the toxic compounds altogether. In soft-bodied insects (e.g., larvae), toxins are usually stored near the teguments or in special glands, ready to release their contents upon handling. The toxicity of insects that extract and sequester plant chemical compounds is dependent on the concentration of these compounds in the host plant. Sawfly larvae (Hymenoptera: Tenthredinidae), for example, reflex-bleed drops of hemolymph when touched; the unpalatability of such larvae is shown to be directly dependent on the glucosinolate concentration of their crucifer host plant over 24 h before "bleeding."

Other insects, however, synthesize their toxins de novo, like many chrysomelid beetles; they probably use the same enzymatic machinery that serves (or has served, in their ancestors) to detoxify the plant's secondary compounds. Although many of these species still use precursors derived from their food plant, these insects are usually less dependent on the plant's toxicity to develop their own noxious compounds. Some groups like ithomiine or heliconiine butterflies also get toxin precursors in their adult diet.

Whatever route to distastefulness is taken, we observe a general correlation between clades of distasteful insects and toxicity in host plant families. In butterflies, the distasteful Troidinae (Papilionidae) tend to feed on Aristolochiaceae, monarchs (Nymphalidae: Danainae) usually feed on milkweeds (Asclepiadaceae), longwing butterflies (Nymphalidae: Heli-coniinae) feed on Passifloraceae, and clearwings (Nymphalidae: Ithomiinae) mainly on Solanaceae and Apocynaceae. In contrast, butterfly clades feeding on chemical-free mono-cotyledonous plants, like browns (Nymphalidae: Satyrinae) on grasses, or owl-butterflies (Nymphalidae: Brassolinae) on palms or Marantaceae, did not evolve distastefulness. Thus toxicity in insects may frequently have evolved as a mere byproduct of adaptation to utilize new kinds of food, particularly toxic plants. The costs of detoxification or toxin production could be covered by the benefits of invading competition-free hosts, perhaps assisted by the increased survival afforded by chemical protection.

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Bee Keeping

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