Bauer Thomas
Zoologisches Institut der Universität Wien, Wien, Österreich.
Oecologia. 1974 Mar;14(1-2):139-196. doi: 10.1007/BF00344904.
The study of two species of stenotopic riparian carabides exemplifies the living conditions and behaviour of the visually hunting carnivorous ground-beetles. 1. E. riparius lives in barren riparian regions exposed to the sun, whereasE. cupreus is found on shady banks covered with vegetation. As far as preying is concerned, the behaviour of these two species does not differ from that of other visually hunting insects. Territorial behaviour can sometimes be observed when they defend a den; it occurs regularly when they defend prey. They react to aggression either by resorting to action or by becoming motionless. In the first case, escape behaviour can be observed, whereas in the latter case they resort to a state of akinesia on bieng touched. When severely menaced, they tend to stridulate and to empty their pygidial glands. InE. cupreus, flying capacity serves for the extention of the habitat only, whereas withE. riparius it is a means of escape (flying speed in the latter: 5.9 km/h=3.7 mph). Copulation behaviour consists of a chain of activities, the three links of which-approach, prehension, and copulation-are released by visual, olfactory, and tactile stimuli. The olfactory stimulus they peceive before mounting implies the interspecific copulation barrier. The oviposition behaviour is adapted to the specific biotopic conditions. On average, the development from egg to imago takes 28.3 days inE. riparius, 37.5 days inE. cupreus; it takes place in summer. The maturation of the imagines occurs without obligatory dormancy within one to two months. Thus, the species ofElaphrus represent an original type of annual rhythmicity. In order to hibernate, the beetles bury themselves about 5 cm below the surface of the ground. Their life span amounts to app. 12 months, one hibernation included. 2. The imagines of both species are active by day. The diurnal pattern of activity is closely related to changes in illumination and temperature. The males are more active. The nocturnal activities of the larvae are thought to represent an isolating mechanisms against the imagines. 3. Surface, number of ommatidia and the anatomy of the compound eyes show that the eyes ofE. riparius are less sensitive to light than those ofE. cupreus. This is accompanied by changes of the inner proportions of the eyes in the usual manner. The acuteness of vision of the two species is similar; the field of vision ofE. riparius extends horizontally, that ofE. cupreus dorsally. Individual parts of the eye display a specialization typical of visually hunting insects. During the optomotoric experiment,E. cupreus achieves its optimal acuteness of vision at a lower light intensity thanE. riparius. 4. In light and infrared gradientsE. riparius prefers higher intensities thanE. cupreus. 5. The beetles enlarge hiding-places on the ground by some instinctive activity. Radiation and descent of moisture in the air enhance the appetence for the fixed pattern by raising body temperature and increasing transpiration. Thus the fixed pattern must be considered to be a mechanism which regulates the behaviour of the animals by radiation.
对两种狭域分布的河岸步甲的研究例证了视觉捕食性食肉地甲虫的生存条件和行为。1. 河岸埃氏步甲生活在阳光直射的贫瘠河岸地区,而铜色埃氏步甲则见于植被覆盖的阴凉河岸。就捕食而言,这两个物种的行为与其他视觉捕食性昆虫并无不同。当它们保卫巢穴时,有时会观察到领地行为;当它们保卫猎物时,这种行为则经常发生。它们对攻击的反应要么是采取行动,要么是静止不动。在前一种情况下,可以观察到逃避行为,而在后一种情况下,它们在被触碰时会进入静止不动状态。受到严重威胁时,它们往往会发出摩擦声并排空臀腺。在铜色埃氏步甲中,飞行能力仅用于扩大栖息地范围,而在河岸埃氏步甲中,飞行是一种逃避手段(后者的飞行速度为5.9千米/小时 = 3.7英里/小时)。交配行为由一系列活动组成,其中接近、抓握和交配这三个环节分别由视觉、嗅觉和触觉刺激引发。它们在爬上之前察觉到的嗅觉刺激构成了种间交配障碍。产卵行为适应特定的生物栖息地条件。平均而言,河岸埃氏步甲从卵发育到成虫需要28.3天,铜色埃氏步甲需要37.5天;发育过程在夏季进行。成虫的成熟在一到两个月内完成,无需强制性休眠。因此,埃氏步甲属的物种代表了一种原始的年度节律类型。为了冬眠,甲虫会在地面以下约5厘米处挖掘洞穴。它们的寿命约为12个月,包括一次冬眠。2. 这两个物种的成虫在白天活动。昼夜活动模式与光照和温度的变化密切相关。雄性更为活跃。幼虫的夜间活动被认为是一种与成虫隔离的机制。3. 复眼的表面积、小眼数量和解剖结构表明,河岸埃氏步甲的眼睛对光的敏感度低于铜色埃氏步甲。这伴随着眼睛内部比例以通常方式发生变化。这两个物种的视觉敏锐度相似;河岸埃氏步甲的视野水平扩展,铜色埃氏步甲的视野则向背侧扩展。眼睛的各个部分表现出视觉捕食性昆虫特有的特化。在视动实验中,铜色埃氏步甲在比河岸埃氏步甲更低的光照强度下达到其最佳视觉敏锐度。4. 在光和红外梯度中,河岸埃氏步甲比铜色埃氏步甲更喜欢更高的强度。5. 甲虫通过一些本能活动扩大地面上的藏身之处。空气中水分的辐射和下降通过提高体温和增加蒸腾作用增强了对固定模式的偏好。因此,必须将固定模式视为一种通过辐射调节动物行为的机制。