警戒作用

警戒作用（英语：Aposematism）是爱德华·巴格诺尔·波尔顿为阿尔弗雷德·拉塞尔·华莱士的警戒色（英语：warning coloration）的概念创造的术语。它是反捕食者适应一类，警告潜在捕食者捕食物无利可图的信号。^[1]无利可图可包括任何使猎物难吃的防御，例如毒性、恶臭气味或强烈反抗的天性。警戒信号有益于捕食者和猎物，因为双方皆可避免潜在伤害，从而代代遗传下来。比如在外在特征方面，警戒作用总涉及显著的动物色、声音、气味或其它可感知特点的广告信号。

警戒作用利用了苗勒氏拟态，防御强的物种演化以和其它物种相似。每种物种可以通过模仿类似颜色的物种更低成本更迅速地学习，共享对于捕食者的警告信号。

真正的警戒作用是物种实际有化学或物理防御，即警戒作用不是威慑捕食者的唯一方法。在贝氏拟态中，模仿物种和一个警戒作用典范紧密相似以分享保护，同时物种多有可以惊吓掠食者足够长时间的虚张声势的威胁，不设防的猎物趁机逃脱。

防御机制

警戒作用的功能是通过警告潜在捕食者猎物又例如难吃或有毒的防御，来预防攻击。易检测到的警告是主要防御机制，不可见防御其次。^[2]警戒信号主要是视觉的，用明亮颜色和高对比度图案，例如条纹。警告信号是有毒猎物的诚实征兆，显著性随毒性而演化。^[3]因此，生物越亮而显眼，通常越有毒。^[4]这和威胁相反，威胁是以危险的外观试图吓走捕食者，但是虚张声势，没有任何强大防御支撑。^[5]

最常见而有效的颜色是红、黄、白。^[6]这些颜色和绿色植物对比强、抗光影中的改变、高色差，而且提供取决于距离的伪装.^[6]

流行

在陆地生态系统

警戒作用普遍存在于昆虫中，但在脊椎动物中较少，大多局限于少数爬行动物、两栖动物、鱼类、一些恶臭或有侵略性的哺乳动物。林鵙鹟属，羽毛和皮肤有毒的红黑色鸟，显然其毒性来自它们摄取的有毒甲虫,可以包括在内。^[7]

在海洋生态系统

警戒作用在海洋生态系统的存在有争议。特别是在珊瑚礁的海洋生物，色彩明亮或有图案的很多，包括海绵、珊瑚、软体动物和鱼类，但很少或没有和化学或物理防御有联系。虽然加勒比海珊瑚礁海绵色彩缤纷，很多又有化学防御毒性，但这两个因素之间没有明显关系。^[8]

明亮的蓝圈章鱼有毒。它们大部分时间隐藏在裂缝中，同时用皮肤的色素细胞显示有效的伪装图案。然而，如果刺激了它们，它们迅速改变颜色，在三分之一秒内变为有50-60环闪亮虹彩蓝的亮黄色。^[9]常有人说这是一种警戒色，^[10]^[11]^[12]^[13]该假设很少被验证过。^[14]

理论的起源

华莱士（1867年）

波尔顿（1890年）

华莱士于1877年的一篇关于动物色的文章中创造了术语“警戒色”。^[15]1890年爱德华·巴格诺尔·波尔顿于其书《动物之色》中重命名该概念为“警戒作用”（aposematism）。^[16]他如下描述该术语的来历：

第二头（语义上的颜色）包括警戒色和识别标记：前者警告天敌，因此称之为警告作用的[希腊语，apo，来自，和sema，信号]

演化

在演化过程中警戒作用是矛盾的，因为它使个体在捕食者前显眼，它们可能会被杀死，在捕食者学会避免前特征消失。如果警戒色最初在有这样强大缺点的少数个体上，那它将永远不能在该物种上持续很长时间以成为有利特征。^[17]

支持的解释

备选假设

其它皆是是可能的。捕食者可能天生害怕陌生的形态（新奇恐怖症），足够长时间它们会适应，但通常是暂时的。^[18]

拟态

警戒色是十分成功的策略，在有警戒色和无警戒色物种的进化上影响重大。

参阅

障碍原则
伪装
迷彩
拟态
警戒色

注释

[1]
Santos, J.C.; Coloma, Luis A.; Cannatella, D.C. Multiple, recurring origins of aposematism and diet specialization in poison frogs. PNAS October 28, 2003. 2003. doi:10.1073/pnas.100.22.12792.
[2]
Ruxton, G. D.; Sherratt, T. N.; Speed, M. P. Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry. Oxford University Press. 2004. ISBN 0-19-852859-0.
[3]
Maan, M. E.; Cummings, M. E. Poison frog colors are honest signals of toxicity, particularly for bird predators. American Naturalist. 2012, 179 (1): E1–E14. JSTOR 663197. doi:10.1086/663197.
[4]
Blount, Jonathan D.; Speed, Michael P.; Ruxton, Graeme D.; Stephens, Philip A. Warning displays may function as honest signals of toxicity (PDF). Proceedings of the Royal Society B. 2009, 276: 871–877 [2018-03-09]. PMC 2664363 . PMID 19019790. doi:10.1098/rspb.2008.1407. （原始内容存档 (PDF)于2018-07-20）.
[5]
Sendova-Franks, Ana; Scott, Michelle Pellissier. Featured Articles in This Month's Animal Behaviour. Animal Behaviour. 2015, 100: iii–v. doi:10.1016/j.anbehav.2014.12.013.
[6]
Stevens, M.; Ruxton, G. D. Linking the evolution and form of warning colouration in nature. Proceedings of the Royal Society B: Biological Sciences. 2012, 279: 417–426. PMC 3234570 . PMID 22113031. doi:10.1098/rspb.2011.1932.
[7]
Dumbacher, J. P.; Beehler, B. M.; Spande, T. F.; Garraffo, H. M.; Daly, J. W. Batrachotoxin in the genus Pitohui: chemical defense in birds?. Science. 1992, 258 (5083): 799–801. PMID 1439786. doi:10.1126/science.1439786.
[8]
Pawlik, JR; et al. Defenses of Caribbean sponges against predatory reef fish: I. Chemical deterrency. Marine Ecology Progress Series. 1995, 127: 183–194. doi:10.3354/meps127183.
[9]
Mäthger, L.M., Bell, G.R., Kuzirian, A.M., Allen, J.J. and Hanlon, R.T. How does the blue-ringed octopus (Hapalochlaena lunulata) flash its blue rings?. The Journal of Experimental Biology. 2012, 215 (21): 3752–3757 [2018-03-09]. PMID 23053367. doi:10.1242/jeb.076869. （原始内容存档于2018-01-24）.
[10]
Williams, B.L., Lovenburg, V., Huffard, C.L. and Caldwell, R.L. Chemical defense in pelagic octopus paralarvae: Tetrodotoxin alone does not protect individual paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) from common reef predators. Chemoecology. 2011, 21 (3): 131–141. doi:10.1007/s00049-011-0075-5.
[11]
Huffard, C.L., Saarman, N., Hamilton, H. and Simison, W.B. The evolution of conspicuous facultative mimicry in octopuses: an example of secondary adaptation?. Biological Journal of the Linnean Society. 2010, 101 (1): 68–77. doi:10.1111/j.1095-8312.2010.01484.x.
[12]
Lambert, W.A. A Review of Blue-ringed Octopus Conservation (Masters thesis). Prescott College. 2011.
[13]
Hanlon, R.T.; Messenger, J.B. Cephalopod Behaviour. Cambridge University Press. 1998.
[14]
Umbers, K.D. On the perception, production and function of blue colouration in animals. Journal of Zoology. 2013, 289 (4): 229–242. doi:10.1111/jzo.12001.
[15]
Wallace, Alfred Russel. The Colours of Animals and Plants. I.—The Colours of Animals. Macmillan's Magazine. 1877, 36 (215): 384–408 [2018-03-09]. （原始内容存档于2019-05-14）.
[16]
Marek, Paul. Aposematism. Apheloria. [2012-11-24]. （原始内容存档于2017-07-08）.
[17]
Mappes, Johanna; Marples, Nicola; Endler, John A. The complex business of survival by aposematism. Trends in Ecology and Evolution. 2005, 20 (11): 598–603. doi:10.1016/j.tree.2005.07.011.
[18]
Exernova, Alice; Stys, Pavel; Fucikova, Eva; Vesela, Silvie; Svadova, Katerina; Prokopova, Milena; Jarosik, Vojtech; Fuchs, Roman; Landova, Eva. Avoidance of aposematic prey in European tits (Paridae): learned or innate?. Behavioral Ecology. 2007, 18 (1): 148–156. doi:10.1093/beheco/arl061.

参考文献

Capinera, John L. (编). Encyclopedia of Entomology 2nd. Springer. 2008 [2018-03-09]. ISBN 978-1-4020-6242-1. （原始内容存档于2016-06-24）.
Edmunds, M. Defence in Animals. Longman. 1974. ISBN 0-582-44132-3.
Poulton, Edward Bagnall. The Colours of Animals, their meaning and use, especially considered in the case of insects. London: Kegan Paul, Trench & Trübner. 1890 [2018-03-09]. （原始内容存档于2018-03-18）.
Ruxton, G. D.; Speed, M. P.; Sherratt, T. N. (2004). Avoiding Attack. The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry. Oxford University Press. ISBN 0-19-852860-4

外部链接

[1] [1]
Santos, J.C.; Coloma, Luis A.; Cannatella, D.C. Multiple, recurring origins of aposematism and diet specialization in poison frogs. PNAS October 28, 2003. 2003. doi:10.1073/pnas.100.22.12792.

[2] [2]
Ruxton, G. D.; Sherratt, T. N.; Speed, M. P. Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry. Oxford University Press. 2004. ISBN 0-19-852859-0.

[E._Maan,_M._E_2012-3] [3]
Maan, M. E.; Cummings, M. E. Poison frog colors are honest signals of toxicity, particularly for bird predators. American Naturalist. 2012, 179 (1): E1–E14. JSTOR 663197. doi:10.1086/663197.

[4] [4]
Blount, Jonathan D.; Speed, Michael P.; Ruxton, Graeme D.; Stephens, Philip A. Warning displays may function as honest signals of toxicity (PDF). Proceedings of the Royal Society B. 2009, 276: 871–877 [2018-03-09]. PMC 2664363 . PMID 19019790. doi:10.1098/rspb.2008.1407. （原始内容存档 (PDF)于2018-07-20）.

[Sendova-FranksScott2015-5] [5]
Sendova-Franks, Ana; Scott, Michelle Pellissier. Featured Articles in This Month's Animal Behaviour. Animal Behaviour. 2015, 100: iii–v. doi:10.1016/j.anbehav.2014.12.013.

[M._Stevens,_G._D_2012-6] [6]
Stevens, M.; Ruxton, G. D. Linking the evolution and form of warning colouration in nature. Proceedings of the Royal Society B: Biological Sciences. 2012, 279: 417–426. PMC 3234570 . PMID 22113031. doi:10.1098/rspb.2011.1932.

[7] [7]
Dumbacher, J. P.; Beehler, B. M.; Spande, T. F.; Garraffo, H. M.; Daly, J. W. Batrachotoxin in the genus Pitohui: chemical defense in birds?. Science. 1992, 258 (5083): 799–801. PMID 1439786. doi:10.1126/science.1439786.

[8] [8]
Pawlik, JR; et al. Defenses of Caribbean sponges against predatory reef fish: I. Chemical deterrency. Marine Ecology Progress Series. 1995, 127: 183–194. doi:10.3354/meps127183.

[Mathger2012-9] [9]
Mäthger, L.M., Bell, G.R., Kuzirian, A.M., Allen, J.J. and Hanlon, R.T. How does the blue-ringed octopus (Hapalochlaena lunulata) flash its blue rings?. The Journal of Experimental Biology. 2012, 215 (21): 3752–3757 [2018-03-09]. PMID 23053367. doi:10.1242/jeb.076869. （原始内容存档于2018-01-24）.

[Williams2011-10] [10]
Williams, B.L., Lovenburg, V., Huffard, C.L. and Caldwell, R.L. Chemical defense in pelagic octopus paralarvae: Tetrodotoxin alone does not protect individual paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) from common reef predators. Chemoecology. 2011, 21 (3): 131–141. doi:10.1007/s00049-011-0075-5.

[Huffard2010-11] [11]
Huffard, C.L., Saarman, N., Hamilton, H. and Simison, W.B. The evolution of conspicuous facultative mimicry in octopuses: an example of secondary adaptation?. Biological Journal of the Linnean Society. 2010, 101 (1): 68–77. doi:10.1111/j.1095-8312.2010.01484.x.

[Lambert2011-12] [12]
Lambert, W.A. A Review of Blue-ringed Octopus Conservation (Masters thesis). Prescott College. 2011.

[Hanlon1998-13] [13]
Hanlon, R.T.; Messenger, J.B. Cephalopod Behaviour. Cambridge University Press. 1998.

[Umbers2013-14] [14]
Umbers, K.D. On the perception, production and function of blue colouration in animals. Journal of Zoology. 2013, 289 (4): 229–242. doi:10.1111/jzo.12001.

[:1-15] [15]
Wallace, Alfred Russel. The Colours of Animals and Plants. I.—The Colours of Animals. Macmillan's Magazine. 1877, 36 (215): 384–408 [2018-03-09]. （原始内容存档于2019-05-14）.

[Marek-16] [16]
Marek, Paul. Aposematism. Apheloria. [2012-11-24]. （原始内容存档于2017-07-08）.

[mappes2005-17] [17]
Mappes, Johanna; Marples, Nicola; Endler, John A. The complex business of survival by aposematism. Trends in Ecology and Evolution. 2005, 20 (11): 598–603. doi:10.1016/j.tree.2005.07.011.

[18] [18]
Exernova, Alice; Stys, Pavel; Fucikova, Eva; Vesela, Silvie; Svadova, Katerina; Prokopova, Milena; Jarosik, Vojtech; Fuchs, Roman; Landova, Eva. Avoidance of aposematic prey in European tits (Paridae): learned or innate?. Behavioral Ecology. 2007, 18 (1): 148–156. doi:10.1093/beheco/arl061.

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