Pandolfi John M, Lovelock Catherine E, Budd Ann F
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0121, USA.
Evolution. 2002 Mar;56(3):479-501. doi: 10.1111/j.0014-3820.2002.tb01360.x.
The Pleistocene extinction of the widespread organ-pipe Montastraea coral had measurable morphological and ecological effects on surviving lineages of the Montastraea "annularis" species complex. Extinction of the organ-pipe Montastraea occurred after more than 500,000 years of dominance in the shallow-water reef habitat of Barbados. Extinction resulted in a morphological shift of the columnar Montastraea lineage from thick to thin columns in modern reef environments. Pleistocene colonies of the columnar morphotype sympatric with organ-pipe Montastraea showed greater column widths than those in allopatry. We subjected our data to a number of criteria for interpreting the morphological shift as character release following lifting of competitive pressure after extinction. The morphological differences do not appear to be due either to chance or to physical properties of the marine environment. Differential local extinction and recolonization of four members of the species complex did not occur on Barbados, so that the species coexisted and appear to have coevolved between more than 600,000 and 82,000 years ago. The morphological shift is related to coral growth form and growth rate, and thus reflects the acquisition of a primary resource in corals--light. Character release occurred at the same oceanic Caribbean island (Barbados) where environments have fluctuated with similar variance throughout the period of coexistence. Not only has competition among living members of the Montastraea "annularis" species complex been convincingly demonstrated, but trends in relative abundance among fossil members of the species complex strongly suggest that a competitive hierarchy was operating during their Pleistocene coexistence on Barbados. We also observed an ecological analogue to character release on another Caribbean island. Curaçao. The distribution and abundance of living columnar M. annularis s.s. and massive M. faveolata from the leeward reef crest in Curaçao is greater now than in the Pleistocene, when organ-pipe Montastraea dominated this shallow-water reef habitat. Extinction of the faster growing, shallow-water organ-pipe Montastraea resulted in higher abundance of the columnar Montastraea lineage in shallow-water habitats, where it shifted its morphology to one adapted to high light levels. The species extinction released surviving lineages from a competitive network that had resulted in lower rank abundance in the Pleistocene community and enhanced abundance of both columnar M. annularis s.s. and M. faveolata in modern communities. Full validation of our interpretation of character release must await experiments that demonstrate whether phenotypic differences between populations have a genetic basis. However, we believe the results of this study point to the important, yet heretofore neglected, role that biological interactions have played in the evolution of closely related reef coral species.
分布广泛的管风琴形蒙大拿石芝珊瑚在更新世灭绝,这对蒙大拿石芝“环纹”物种复合体的幸存谱系产生了可测量的形态学和生态学影响。管风琴形蒙大拿石芝在巴巴多斯浅海礁石栖息地占据主导地位超过50万年之后灭绝。灭绝导致柱状蒙大拿石芝谱系在现代礁石环境中从粗柱状向细柱状发生形态转变。与管风琴形蒙大拿石芝同域分布的柱状形态型更新世群体的柱宽比异域分布的群体更大。我们依据若干标准对数据进行分析,以解释这种形态转变是灭绝后竞争压力解除后的特征释放。形态差异似乎既不是由于偶然因素,也不是由于海洋环境的物理特性。该物种复合体的四个成员在巴巴多斯并没有出现不同程度的局部灭绝和重新定殖,因此这些物种共存,并且在距今60多万年至8.2万年前似乎共同进化。形态转变与珊瑚的生长形态和生长速率有关,因此反映了珊瑚对一种主要资源——光的获取。特征释放在同一个加勒比海岛屿(巴巴多斯)发生,在共存期间该岛环境以相似的幅度波动。不仅蒙大拿石芝“环纹”物种复合体现存成员之间的竞争已得到令人信服的证明,而且该物种复合体化石成员的相对丰度趋势强烈表明,在更新世它们于巴巴多斯共存期间存在一个竞争等级体系。我们在另一个加勒比海岛屿库拉索岛也观察到了与特征释放类似的生态现象。库拉索岛背风礁顶现存的柱状环纹蒙大拿石芝和块状蜂巢蒙大拿石芝的分布范围和丰度现在比更新世时更大,当时管风琴形蒙大拿石芝占据这个浅海礁石栖息地。生长较快的浅海管风琴形蒙大拿石芝的灭绝导致浅海栖息地柱状蒙大拿石芝谱系的丰度增加,其形态转变为适应高光水平的形态。物种灭绝使幸存谱系从一个竞争网络中解脱出来,这个竞争网络在更新世群落中导致较低的等级丰度,而在现代群落中则提高了柱状环纹蒙大拿石芝和蜂巢蒙大拿石芝的丰度。要完全验证我们对特征释放的解释,必须等待实验来证明种群之间的表型差异是否具有遗传基础。然而,我们认为这项研究的结果表明了生物相互作用在近缘珊瑚礁珊瑚物种进化中所起的重要但迄今被忽视的作用。