Behavioural Biology of the Collared Lemming [Dicrostonyx groenlandicus (Trail)]: An Analysis of Acoustic Communication

Ronald J. Brooks , Edwin M. Banks
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引用次数: 64

Abstract

The acoustic behaviour of collared lemmings was investigated in a laboratory study. Observations were conducted on behaviour of lemmings in a variety of test situations including: paired intraspecific male-male, male-oestrous female and male-dioestrous female encounters in an arena unfamiliar to both animals; paired intraspecific male-oestrous female encounters in the female's home cage; groups of lemmings in large enclosures; isolated, adult lemmings exposed to several stimulus situations; isolated pups subjected to a series of tests during development and mother-neonate interactions.

A repertoire of the acoustic behaviour of lemmings was established and catalogued according to both physical and auditive properties of the several calls described, as well as the behavioural context in which the calls occurred. Physical analysis was accomplished using spectrographic and oscillographic measurements of calls recorded on tape. Ontogeny of the vocalizations was traced from birth to maturity. Sexual and individual variation in the sounds was also investigated.

An attempt was made to determine the communicative function of each sound by correlating its occurrence with twenty previously defined non-vocal acts and postures and by observing responses of other lemmings to the calls. Detailed quantitative analysis was applied to the data, but interpretation of results of this analysis was limited by lack of knowledge concerning the perceptual capabilities of lemmings.

The following results were obtained:

  • 1.

    (1) Six distinct types of sounds were recognized; ultrasonic chirp, ultrasonic mating twitter, huh-huh call, tooth-chatter, squeal-squawkgrind complex and a group of less defined sounds classified into four sub-types: peeps, whines, mechanically produced sounds and an assortment of snorts, chuckles, pops and snarls.

  • 2.

    (2) The first three types listed above were associated with relatively specific behavioural contexts. Ultrasonic chirps were given exclusively by pups less than 15 days old in response to cold stress and brief, non-painful tactile stimuli. This vocalization apparently aided the dam in noticing and locating pups which had fallen out of the nest.

  • 3.

    (3) The ultrasonic mating twitter was emitted only by adults performing male-like sexual behaviour such as mounting, chasing and allogrooming. This call indicated that the caller was sexually aroused.

  • 4.

    (4) Huh-huh calls were given by adult lemmings when they detected the presence of a predator. The call accompanied an erratic, explosive leaping behaviour which apparently served to distract the predator and allow the lemming more time to escape.

  • 5.

    (5) Tooth-chatter was associated with threat behaviour and increased excitation.

  • 6.

    (6) Squeals, squawks and grinds were distinct calls but intergraded along a continuum for frequency, duration, intensity, harmonic structure and various other physical parameters. These calls were associated with agonistic behaviour, specifically fear motivated actions. The graded system for these calls apparently was based upon small variations in motivation and the existence of this system suggests there is some selective advantage in communicating these variations. As a lemming became more frightened, calls of this type became shorter, louder and shriller (squeal). These calls were given by subordinates, in male-male encounters and by females in male-oestrous female encounters.

  • 7.

    (7) Squeals were the only calls given at all ages. It was suggested that such sounds when given by very young pups inhibited the dam from injuring them.

  • 8.

    (8) All calls displayed considerable variation in physical properties.

  • 9.

    (9) The interaction of behavioural and ecological factors was discussed in terms of the evolution of age-related changes in various properties of the ultrasonic chirp.

  • 10.

    (10) It was concluded that the communicative role of the calls could only be understood in the context of simultaneous non-vocal behaviour. Meaning of each signal or message probably varies with this context and the precise configuration of the total display will need to be known fully to appreciate the communicative significance of each call. Emission of a particular call reflected the underlying physiological state of the animal. When this relationship becomes better established vocal behaviour may be a useful tool in the study of motivation and in assessing physiological state and various thresholds in the vocalizing animal.

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项圈旅鼠的行为生物学[dirostonyx groenlandicus (Trail)]:声学通信分析
在一项实验室研究中调查了带项圈旅鼠的声学行为。研究人员对旅鼠在多种测试情境下的行为进行了观察,包括:在两种动物都不熟悉的环境中成对的种内雄性-雄性、雄性-发情的雌性和雄性-发情的雌性相遇;在雌性家笼中成对的种内雄性-雌性交配;成群的旅鼠在大围栏里;孤立的成年旅鼠暴露在几种刺激情况下;孤立的幼崽在发育和母幼互动期间接受一系列测试。根据所描述的几种叫声的物理和听觉特性,以及叫声发生的行为背景,建立了旅鼠声学行为的曲目并进行了分类。物理分析是通过对记录在磁带上的通话进行光谱和示波器测量来完成的。发声的个体发生可以追溯到从出生到成熟。研究人员还调查了声音的性别和个体差异。通过将每种声音的出现与20种先前定义的非声音行为和姿势相关联,并通过观察其他旅鼠对这些声音的反应,试图确定每种声音的交流功能。对数据进行了详细的定量分析,但由于缺乏对旅鼠感知能力的了解,对分析结果的解释受到限制。结果表明:1.(1)识别出6种不同类型的声音;超声波唧唧声、超声波交配推特声、哼哼声、牙颤声、吱吱声和一组不太明确的声音分为四类:吱吱声、呜咽声、机械产生的声音以及各种各样的鼻息声、咯咯声、砰砰声和咆哮声。(2)上面列出的前三种类型与相对特定的行为环境有关。未满15天的幼犬在受到冷应激和短暂的无痛触觉刺激时,只能发出超声波鸣叫。这种叫声显然有助于雄鼠注意和定位从巢中掉下来的幼崽。(3)只有成年雄鼠在进行类似雄性的性行为时才会发出这种超声波交配鸣叫,比如攀爬、追逐和梳理毛发。当成年旅鼠发现有捕食者时,就会发出“Huh-huh”的叫声。这种叫声伴随着一种不稳定的、爆炸式的跳跃行为,这显然是为了分散捕食者的注意力,让旅鼠有更多的时间逃跑。(5)牙颤与威胁行为和增加的兴奋有关。(6)尖叫、嘎嘎声和碾磨声是不同的叫声,但在频率、持续时间、强度、和声结构和各种其他物理参数上都是连续统合的。这些呼叫与敌对行为有关,特别是恐惧动机的行为。这些呼叫的分级系统显然是基于动机的微小变化,这个系统的存在表明在交流这些变化时有一些选择性优势。当旅鼠变得更害怕时,这种叫声变得更短、更响、更尖(尖叫)。这些叫声是由下属发出的,在雄性与雄性相遇时发出的,而雌性在雄性与雌性发生情色时发出的。(8)所有的叫声在物理性质上都表现出相当大的变化。(9)从超声啁啾各种性质的年龄相关变化的演变角度,讨论了行为和生态因素的相互作用。(10)结论是,只有在同时发生非声音行为的背景下,才能理解叫声的交流作用。每个信号或信息的含义可能会随着上下文而变化,并且需要完全了解整个显示器的精确配置,以欣赏每个呼叫的交流意义。发出一种特殊的叫声反映了动物潜在的生理状态。当这种关系得到更好的建立时,发声行为可能成为研究动机和评估发声动物的生理状态和各种阈值的有用工具。
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