Foster S A, Garcia V B, Town M Y
Department of Ecology and Evolution, State University of New York at Stony Brook, 11794, Stony Brook, NY, USA.
Department of Zoology, University of Washington, 98195, Seattle, WA, USA.
Oecologia. 1988 Jan;74(4):577-585. doi: 10.1007/BF00380056.
In Crystal Lake, British Columbia, small fry (≤15 mm SL) of the threespine stickleback (Gasterosteus aculeatus) are concentrated in vegetation while larger fry are not. Because fry in all size classes feed primarily on zooplankton, even when in vegetation, we hypothesized that size-limited predation was responsible for the observed shift in habitat use with size. The major predators on fry in Crystal Lake are adult threespine stickleback, the water scorpion, Ranatra sp. (Hemiptera: Nepidae), backswimmers, Notonecta spp. (Hemiptera: Notonectidae), and dragonfly naiads of the genus Aeshna (Odonata: Aeschnidae). On the basis of distribution and hunting behavior we excluded the insects Ranatra sp., and Notonecta sp. as causal agents for this shift in resource by fry in water >0.25 m deep. Ranatra was found almost exclusively near the shoreline in water <0.25 m deep, and both insects hunted primarily as ambush predators within vegetation. Such predators seemed more likely to drive vulnerable fry from vegetation than to restrict them to it. In contrast, Aeshna naiads and adult stickleback frequently hunted outside of vegetation. In prey preference experiments the naiads did not show the decline in predation efficiency on fry >15 mm SL that would be expected if size-limited predation by this insect was responsible for the observed shift in resource use by fry. Adult stickleback only fed on fry <15 mm SL, and in an experimental situation, consumed fry at a rate 10 times greater than that exhibited by any of the insects. Predation experiments demonstrated that small fry (11-15 mm) spent more time in vegetation in the presence of adult conspecifics than they did in control pools, as would be expected if size-limited cannibalism caused small, vulnerable fry to be restricted to vegetation. Fry >15 mm SL were found outside of vegetation more often than in control treatments. The probable cause of this result is that adults become aggressive toward fry at this size, and often could be seen chasing large fry from vegetation during the experiments. Dragonfly naiads (Aeshna spp.) spent most of their time in vegetation in the experimental pools. Both size classes of fry spent less time in vegetation in the presence of dragonfly naiads than they did in control treatments, an apparent reflection of their similar vulnerabilities to these naiads. The presence of vegetation in pools reduced predation rates by adult stickleback on small fry. Because the experiments presented here indicate that fry are capable of rapidly assessing predation risk and of altering their behavior adaptively, we conclude that small fry occupy vegetation as a refuge from cannibalism. Once fry have reached the size-threshold at which they are no longer vulnerable to adult conspecifics they are able to forage farther from vegetation thereby reducing risk of predation by insects in vegetation and possibly acquiring more abundant food resources.
在不列颠哥伦比亚省的克里斯特尔湖,三刺鱼(学名:Gasterosteus aculeatus)的幼鱼(体长≤15毫米)集中在水生植物区域,而较大的幼鱼则不然。由于所有大小等级的幼鱼主要以浮游动物为食,即便处于水生植物区域时也是如此,因此我们推测,大小受限的捕食行为是观察到的幼鱼栖息地利用随大小变化的原因。克里斯特尔湖幼鱼的主要捕食者有成年三刺鱼、红娘华(学名:Ranatra sp.,半翅目:蝎蝽科)、仰泳蝽(学名:Notonecta spp.,半翅目:仰泳蝽科)以及艾氏属的蜻蜓稚虫(蜻蜓目:蜓科)。基于分布情况和捕食行为,我们排除了红娘华和仰泳蝽作为导致水深超过0.25米处幼鱼资源利用发生这种变化的因素。红娘华几乎只在水深小于0.25米的海岸线附近被发现,而且这两种昆虫主要在水生植物中以伏击式捕食者的方式捕猎。这类捕食者似乎更有可能把易受攻击的幼鱼从水生植物中驱赶出来,而不是将它们限制在其中。相比之下,艾氏蜻蜓稚虫和成年三刺鱼经常在水生植物之外捕猎。在猎物偏好实验中,如果这种昆虫的大小受限捕食行为是观察到的幼鱼资源利用变化的原因,那么蜻蜓稚虫对体长超过15毫米的幼鱼的捕食效率应该会下降,但实验结果并非如此。成年三刺鱼只捕食体长小于15毫米的幼鱼,并且在实验环境中,其捕食幼鱼的速度比任何一种昆虫都快10倍。捕食实验表明,与对照水池相比,在有成年同类存在的情况下,幼小鱼(11 - 15毫米)在水生植物中的停留时间更长,这正如预期的那样,如果大小受限的同类相食导致弱小、易受攻击的幼鱼被限制在水生植物中。体长超过15毫米的幼鱼在水生植物之外被发现的频率高于对照处理组。这一结果的可能原因是,成年三刺鱼对这个大小的幼鱼变得具有攻击性,在实验过程中经常可以看到它们把较大的幼鱼从水生植物中驱赶出来。蜻蜓稚虫(艾氏属)在实验水池中大部分时间都待在水生植物中。与对照处理组相比,两种大小等级的幼鱼在有蜻蜓稚虫存在的情况下在水生植物中的停留时间都减少了,这显然反映出它们对这些蜻蜓稚虫具有相似的易受攻击性。水池中有水生植物降低了成年三刺鱼对幼小鱼的捕食率。由于这里展示的实验表明幼鱼能够快速评估捕食风险并适应性地改变它们的行为,因此我们得出结论,幼小鱼占据水生植物作为躲避同类相食的避难所。一旦幼鱼达到不再易受成年同类攻击的大小阈值,它们就能够在离水生植物更远的地方觅食,从而降低在水生植物中被昆虫捕食的风险,并有可能获取更丰富的食物资源。