Facial protective reflexes are usually well developed in flat-faced mammalian predators like cats and tarsiers, whose eyes and ears are especially exposed to injury by prey. The reflexes also are exaggerated in social species for use in communication; thus ear flattening in horse and dog displays has a counterpart in scalp retraction among Old World monkeys. The scalp movements and raised brows are effectively used in communication, despite the greatly reduced mobility of ears among monkeys. Limbs and other appendages (e.g., antennae) are withdrawn or used to protect sensitive areas by both vertebrates and invertebrates. Among mollusks and such groups as sea squirts and barnacles, the whole soft body can be retracted into a protective shell, or carapace; a kind of door (operculum) may be used to stop the entrance (e.g., among snails and barnacles), and trap-door spiders pull the stopper in place behind them. Bone may have evolved in fossil vertebrates as protective armour in jawless ancient fishes (ostracoderms), probably as a result of natural selection in the face of dominant arthropod predators (eurypterids). With the evolution of jaws, the vertebrates themselves gave rise to nearly all later large predators. Evolutionary advantage then apparently came with complex sense organs and behaviour; in most vertebrate lines there is evidence of a progressive reduction in body armour (dermal bone). Thus, although such cartilaginous fishes as sharks and rays do not exhibit such bony skins, they may well have evolved from heavily armoured ancient fishes (placoderms).

Armour nevertheless has evolved repeatedly, particularly among animals incapable of fast locomotion; trunkfish (boxfish), for example, have a body entirely boxed by bony plates; and tortoises and turtles are perhaps the most completely armoured of four-legged animals. The turtles seem to have evolved early from the basal stock of the reptiles; thanks to the shell (carapace) within which they can withdraw head, limbs, and tail, they represent one of the few reptilian orders that have remained consistently successful. The turtle’s dorsal carapace appears to consist of newly evolved plates of dermal bones, but the belly plates (ventral plastron) may well be retained in part from fish ancestors. Reptilian land vegetarians usually tended to evolve armour, as in the fossil dinosaurs such as stegosaurs and ankylosaurs.

South American toothless animals (edentates) such as anteaters are probably survivors of a comparable early development in mammals. The armour of armadillos and the presence of bony plates in the skin of the extinct sloths suggest that the whole group may derive from an armoured ancestor. The appearance of hair in the mammal line seems to have led to the evolution of a light, spiny type of armour. Such modern mammals as hedgehogs, echidnas, insect-eating tenrecs, and some rodents and their relatives (lagomorphs) all possess defensive spines that are commonly erectile and are often able to roll into a ball like an armadillo.

Chemical means of defense may be widely distributed in the body, making the animal distasteful to predators. Some of these chemical compounds may be derived from plants eaten or synthesized by the animal itself (e.g., bufotoxin in toads). Although the animal attacked may be killed and thus not benefit, his fellows do, since they are likely to be avoided by the predator. Poison or distasteful substances may also be ejected from a bodily reservoir and squirted at the enemy (e.g., the skunk, some ants) or inserted into a puncture made by a spine (e.g., triggerfish) or teeth (e.g., certain snakes). Many poisons act to paralyze muscles by blocking nerve transmission at the neuromuscular junction (e.g., cobra venom).

Warning behaviour

Mobbing behaviour apparently advertises the presence of a predator that is potentially but not immediately dangerous; thus, mammalian nest predators can be safely mobbed by flying birds, as can owls in the daytime. From the safety of trees, monkeys and squirrels mob predators on the ground. Mobbing calls are typically easy to locate, the calls being short and staccato, and they provide excellent cues of distance and direction. Conspicuous movements, such as tail flicks among small birds and squirrels, accompany the calls.

More urgently, intense warning behaviour is given in response to sources of immediate danger. Warning calls are usually long whistles that make location difficult because of their gradual onset and termination and their narrow ranges of pitch. The evolution of warning behaviour that puts the displaying animal in danger (such as these intense warning calls) seems likely to come about only if the benefit to offspring and other members of the species is great. Indeed, it has been calculated that, if an individual loses his life as the result of his warning behaviour, increased transmission of his family’s genes will result only if the reproductive rate of relatives (as close as sisters) is doubled as a result of his sacrifice.