Stabbing Spines: A review of the Biomechanics and Evolution of Defensive Spines.

Spines are ubiquitous in both plants and animals, and while most spines were likely originally used for defense, over time many have been modified in a variety of ways. Here we take an integrative approach to review the form, function, and evolution of spines as a defensive strategy in order to make new connections between physical mechanisms and functional behavior. While this review focuses on spines in mammals, we reference and draw ideas from the literature on spines in other taxa, including plants. We begin by exploring the biomechanics of defensive spines, their varied functions, and nondefensive modifications. We pay particular attention to the mechanics involved in passive puncture and the ways organisms have overcome limitations associated with the low energy input. We then focus on the ecological, physiological, and behavioral factors that promote the evolution of spiny defenses, including predator- and habitat-mediated hypotheses. While there is considerable evidence to support both, studies have generally found that (1) defensive spines are usually effective against one class of attacker (e.g., larger predators) but ineffective against or even facilitate predation by others and (2) species that are more visible or exposed to predators are under much stronger selection to evolve defensive spines or some other robust defense. What type of defensive morphology that evolves, however, is less predictable and probably strongly dependent on both the dominant source of predation and the habitat structure of the organism (e.g., arboreal, terrestrial, and fossorial). We then explore traits that often are correlated with defensive spines and armor, potentially forming armor syndromes, suites of traits that evolve together with body armor in a correlated fashion. In mammals, these include aposematic warning coloration, locomotion style, diet, metabolic rate, and relative brain size. Finally, we encourage integration of mechanistic, behavioral, and evolutionary studies of defensive spines and suggest future avenues of research in the biomechanics, evolution, and behavior of spines and spiny organisms.