Ruxton Graeme D, Speed Michael P
Division of Environmental & Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Graham Kerr Building, Glasgow G12 8QQ, UK.
Proc Biol Sci. 2006 Feb 7;273(1584):373-8. doi: 10.1098/rspb.2005.3238.
It is common for species that possess toxins or other defences to advertise these defences to potential predators using aposematic ("warning") signals. There is increasing evidence that within such species, there are individuals that have reduced or non-existent levels of defence but still signal. This phenomenon (generally called automimicry) has been a challenge to evolutionary biologists because of the need to explain why undefended automimics do not gain such as a fitness advantage by saving the physiological costs of defence that they increase in prevalence within the population, hence making the aposematic signal unreliable. The leading theory is that aposematic signals do not stop all predatory attacks but rather encourage predators to attack cautiously until they have identified the defence level of a specific individual. They can then reject defended individuals and consume the undefended. This theory has recently received strong empirical support, demonstrating that high-accuracy discrimination appears possible. However, this raises a new evolutionary problem: if predators can perfectly discriminate the defended from the undefended and preferentially consume the latter, then how can automimicry persist? Here, we present four different mechanisms that can allow non-trivial levels of automimics to be retained within a population, even in the extreme case where predators can differentiate defended from undefended individuals with 100% accuracy. These involve opportunity costs to the predator of sampling carefully, temporal fluctuation in predation pressure, predation pressure being correlated with the prevalence of automimicry, or developmental or evolutionary constraints on the availability of defence. These mechanisms generate predictions as to the conditions where we would expect aposematically signalling populations to feature automimicry and those where we would not.
拥有毒素或其他防御机制的物种通过警戒(“警告”)信号向潜在捕食者宣传这些防御机制是很常见的。越来越多的证据表明,在这类物种中,有些个体的防御水平降低或不存在,但仍会发出信号。这种现象(通常称为自拟态)一直是进化生物学家面临的一个挑战,因为需要解释为什么无防御能力的自拟态个体不会通过节省防御的生理成本而获得诸如适应性优势,从而导致它们在种群中的比例增加,进而使警戒信号变得不可靠。主要理论是,警戒信号并不能阻止所有的捕食攻击,而是鼓励捕食者谨慎攻击,直到它们确定特定个体的防御水平。然后它们可以拒绝有防御能力的个体,捕食无防御能力的个体。这一理论最近得到了有力的实证支持,表明高精度的辨别似乎是可能的。然而,这又引发了一个新的进化问题:如果捕食者能够完美地区分有防御能力和无防御能力的个体,并优先捕食后者,那么自拟态是如何持续存在的呢?在这里,我们提出了四种不同的机制,即使在捕食者能够100%准确区分有防御能力和无防御能力个体的极端情况下,这些机制也能使种群中保留一定数量的自拟态个体。这些机制包括捕食者仔细取样的机会成本、捕食压力的时间波动、捕食压力与自拟态流行程度的相关性,或者防御可用性的发育或进化限制。这些机制对我们预期会出现自拟态的警戒信号种群的条件以及不会出现自拟态的条件产生了预测。