Keogh J Scott, Scott Ian A W, Hayes Christine
School of Botany and Zoology, The Australian National University, Canberra, ACT, Australia.
Evolution. 2005 Jan;59(1):226-33.
It is a well-known phenomenon that islands can support populations of gigantic or dwarf forms of mainland conspecifics, but the variety of explanatory hypotheses for this phenomenon have been difficult to disentangle. The highly venomous Australian tiger snakes (genus Notechis) represent a well-known and extreme example of insular body size variation. They are of special interest because there are multiple populations of dwarfs and giants and the age of the islands and thus the age of the tiger snake populations are known from detailed sea level studies. Most are 5000-7000 years old and all are less than 10,000 years old. Here we discriminate between two competing hypotheses with a molecular phylogeography dataset comprising approximately 4800 bp of mtDNA and demonstrate that populations of island dwarfs and giants have evolved five times independently. In each case the closest relatives of the giant or dwarf populations are mainland tiger snakes, and in four of the five cases, the closest relatives are also the most geographically proximate mainland tiger snakes. Moreover, these body size shifts have evolved extremely rapidly and this is reflected in the genetic divergence between island body size variants and mainland snakes. Within south eastern Australia, where populations of island giants, populations of island dwarfs, and mainland tiger snakes all occur, the maximum genetic divergence is only 0.38%. Dwarf tiger snakes are restricted to prey items that are much smaller than the prey items of mainland tiger snakes and giant tiger snakes are restricted to seasonally available prey items that are up three times larger than the prey items of mainland tiger snakes. We support the hypotheses that these body size shifts are due to strong selection imposed by the size of available prey items, rather than shared evolutionary history, and our results are consistent with the notion that adaptive plasticity also has played an important role in body size shifts. We suggest that plasticity displayed early on in the occupation of these new islands provided the flexibility necessary as the island's available prey items became more depauperate, but once the size range of available prey items was reduced, strong natural selection followed by genetic assimilation worked to optimize snake body size. The rate of body size divergence in haldanes is similar for dwarfs (h(g) = 0.0010) and giants (h(g) = 0.0020-0.0025) and is in line with other studies of rapid evolution. Our data provide strong evidence for rapid and repeated morphological divergence in the wild due to similar selective pressures acting in different directions.
岛屿能够维持大陆同种生物的巨型或侏儒形态的种群,这是一个众所周知的现象,但针对这一现象的各种解释性假说却难以厘清。剧毒的澳大利亚虎蛇(澳虎属)就是岛屿体型变异的一个著名且极端的例子。它们特别引人关注,因为存在多个侏儒型和巨型种群,而且根据详细的海平面研究可知岛屿的年龄,进而可知虎蛇种群的年龄。大多数岛屿有5000 - 7000年历史,所有岛屿都不到10000年历史。在这里,我们利用一个包含约4800个碱基对线粒体DNA的分子系统地理学数据集,对两种相互竞争的假说进行了区分,并证明岛屿侏儒型和巨型种群已经独立进化了五次。在每种情况下,巨型或侏儒型种群的近亲都是大陆虎蛇,并且在五次情况中的四次里,近亲也是地理上最接近的大陆虎蛇。此外,这些体型变化进化得极其迅速,这在岛屿体型变异体与大陆蛇之间的遗传差异中得到了体现。在澳大利亚东南部,岛屿巨型种群、岛屿侏儒型种群和大陆虎蛇种群都有分布,最大遗传差异仅为0.38%。侏儒型虎蛇只能捕食比大陆虎蛇的猎物小得多的猎物,而巨型虎蛇则只能捕食季节性可得的、比大陆虎蛇的猎物大三倍的猎物。我们支持这样的假说,即这些体型变化是由于可获得猎物的大小所施加的强烈选择,而非共同的进化历史,并且我们的结果与适应性可塑性在体型变化中也起到重要作用的观点一致。我们认为,在这些新岛屿被占据初期所展现出的可塑性,在岛屿可获得猎物变得更加匮乏时提供了必要的灵活性,但一旦可获得猎物的大小范围缩小,强大的自然选择随后通过遗传同化作用来优化蛇的体型。侏儒型(h(g) = 0.0010)和巨型(h(g) = 0.0020 - 0.0025)虎蛇在霍尔丹法则下的体型分化速率相似,并且与其他快速进化的研究结果一致。我们的数据为由于不同方向的类似选择压力导致的野外快速且反复的形态分化提供了有力证据。