Comparative Studies of the Behaviour of Gulls (Laridae): a Progress Report1)
Comparative Studies of the Behaviour of Gulls (Laridae): a Progress Report1)
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海鸥（鸥科）行为的比较研究：进展报告1)
DOI：
10.1163/156853960x00098
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发表时间：
1960
期刊：
影响因子：
--
通讯作者：
N. Tinbergen
中科院分区：
文献类型：
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作者：
N. Tinbergen
本文描述了鸥科（Laridae）的一些鸟类，特别是银鸥群、冠鸥和三趾鸥，并讨论了它们的功能、成因、进化起源和作为信号的进一步进化。在简述了比较行为研究的基本原理之后（第二章），描述了最常见的单一展示和展示序列（第三章）。它们在整个家族中相当相似，尽管存在许多物种特异性差异。行为相似性加强了海鸥是一个单系群的结论。第四章给出了显示器具有信号功能的证据。在距离增加和距离减小显示之间进行区分。发生的原因不止一个，但不超过约六个距离增加的姿态进行了讨论，有人认为，每个显示器可能是适应于处理一个不同类别的对手：意外入侵者需要另一个驱避剂比故意持续入侵者，并在每一类实际的入侵者遇到的另一种方式比潜在的入侵者。此外，一个特别响亮的电话是一个典型的长途威胁。这种叫声通常在双重意义上起着广告的作用：它击退竞争对手，吸引未交配的雌性。一些减少距离（或安抚）的姿势用于对抗的情况下，更经常地在配对或见面仪式结束时使用。需要和显示器的精确功能的更系统的研究的可能性被强调。第五章论述了陈列动机分析的目的和方法。强调“自然实验”的价值。三个独立的方法的应用导致的结论是，对抗显示是矛盾的，即结果的同时唤醒的倾向，攻击和逃跑的倾向：波动之间的关系，这些倾向和显示检查。在这些展示的动机和它们传递给其他个体的信息之间有着惊人的相关性。安抚姿态总是包含恐惧的成分;这种倾向与停留的倾向相冲突，这可能是，但通常不是攻击倾向的一部分;它可能是性吸引，吸引到巢穴，或吸引到食物提供者。强调需要对一种姿势可能发生的许多不同形式进行动机分析。不同物种的配对仪式中动机的相似性远大于展示序列的形式相似性，验证了求爱的冲突理论。显示器（第六章）的起源各不相同。有些显然是作为由情境直接引起的模式的预备性或意向性运动（“自主性”运动）而出现的;其中，有些被重定向到无生命的物体。其他人来自运动属于功能模式不直接引起的情况下（“位移活动”），他们的各种起源进行了讨论，它表明，他们是第二个组成部分的双重运动，其中第一个组成部分是“自动？在上述意义上，”当地“是一个概念，并为流离失所活动提供了便利。在第七章中，一些进化变化的最终原因进行了讨论，并提出了一个初步的功能分类的所谓的变化显示。有人认为，变化已被增强了至少四种不同类型的选择压力：（i）朝着改善信号功能（显着性）;（2）朝着增加种内的不模糊性;（3）朝着增加种间的不模糊性;（4）作为一个correlary的选择压力在其他功能系统。在某些海鸥中，（4）可能是造成性隔离的主要原因。一个初步的分类给出了假设的行为机制的进化变化。被认为是分类使用的行为特征的价值。讨论后的自适应和非自适应的差异LORLNz的相对有效性的强调系统发育的保守性的显示器被重新考虑，并证明收敛的相似性的发生。
This paper describes a number of displays of various gulls (Laridae) with special reference to the Herring Gull group, the hooded gulls, and the Kittiwake, and discusses their functions, causation, evolutionary origin and further evolution as signals. After a sketch of the rationale of comparative behaviour studies (chapter II), the most common single displays and display sequences are described (chapter III). They are rather similar through the family, though many species-specific differences exist. The behaviour similarities strengthen the conclusion that the gulls are a monophyletic group. Evidence is presented in chapter IV to show that the displays have signal function. A distinction is made between distance-increasing and distance-reducing displays. The reasons for the occurence of more than one, yet no more than about six distance- increasing postures are discussed ; it is argued that each display may well be adapted to deal with a distinct category of opponent : accidental trespassers require another repellent than intentional persistent intruders; and within each category actual trespassers are met in another way than potential trespassers. In addition, one particularly loud call is a typical long-distance threat. This call usually acts as advertisement in a double sense: it repels competitors and attracts unmated females. Some of the distance-reducing (or appeasement) postures are used in agonistic situations, and still more regularly at the end of the pair formation or meeting ceremony. The need for and the possibility of more systematic studies of the precise functions of displays is stressed. Chapter V discusses aims and methods of analysis of the motivation of the displays. The value of "natural experiments" is stressed. The application of three independent methods leads to the conclusion that agonistic displays are ambivalent, i.e. the outcome of the simultaneous arousal of a tendency to attack and a tendency to flee: the relation between fluctuations in these tendencies and the displays shown is examined. There is a striking correlation between the motivation of these displays and the information they pass on to other individuals. Appeasement gestures always contain an element of fear; this tendency is in conflict with a tendency to stay, which can be, but usually is not part of the tendency to attack; it may be sexual attraction, attraction to at nest site, or attraction to a provider of food. The need for motivational analyses of the many different forms in which one posture can occur is stressed. The similarity of motivation in the pair formation ceremonies in the diff.erent species is much greater than the formal similarity of the display sequences; the conflict theory of courtship is tested. The origin of the displays (chapter VI) is varied. Some have clearly arisen as preparatory or intention movements of the patterns directly aroused by the situation ("aut- ochthonous" movements) ; of these, some are redirected to inanimate objects. Others are derived from movements belonging to functional patterns not directly aroused by the situation ("displacement activities") ; their various origins are discussed, and it is shown that they are second components of a dual movement, of which the first component is "auto?hthonous" in the above sense and facilitates the displacement activity. In chapter VII some ultimate causes of evolutionary change are discussed, and a preliminary functional classification of alleged changes in displays is presented. It is argued that change has been enhanced by at least four different types of selection pressure: (i) towards improvement of signal function (conspicuousness) ; (2) towards increase in intraspecific unambiguity; (3) towards increased interspecific unambiguity; and (4) as a corrollary of selection pressure in other functional systems. In some gulls, (4) may have made the major contribution to sexual isolation. A preliminary classification is given of the postulated evolutionary changes in behaviour mechanisms. The value of behaviour characters for taxonomic use is considered. After a discussion of adaptive and non-adaptive differences the relative validity of LORLNz's emphasis on the phylogenetic conservatism of displays is reconsidered, and the occurrence of convergent similarities is demonstrated.