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物种水平趋势的系统发育:四足动物的物种遗传性及伯格曼法则可能的深层起源

The phylogeny of a species-level tendency: species heritability and possible deep origins of Bergmann's rule in tetrapods.

作者信息

De Queiroz Alan, Ashton Kyle G

机构信息

Archbold Biological Station, Venus, Florida 33960, USA.

出版信息

Evolution. 2004 Aug;58(8):1674-84. doi: 10.1111/j.0014-3820.2004.tb00453.x.

Abstract

One of the most widely recognized generalizations in biology is Bergmann's rule, the observation that, within species of birds and mammals, body size tends to be inversely related to ambient temperature. Recent studies indicate that turtles and salamanders also tend to follow Bergmann's rule, which hints that this species-level tendency originated early in tetrapod history. Furthermore, exceptions to Bergmann's rule are concentrated within squamate reptiles (lizards and snakes), suggesting that the tendency to express a Bergmann's rule cline may be heritable at the species level. We evaluated species-level heritability and early origination of Bergmann's rule by mapping size-latitude relationships for 352 species onto a tetrapod phylogeny. When the largest available dataset is used, Bergmann's rule shows significant phylogenetic signal, indicating species-level heritability. This represents one of the few demonstrations of heritability for an emergent species-level property and the first for an ecogeographic rule. When species are discretely coded as showing either Bergmann's rule or its converse, parsimony reconstructions suggest that: (1) the tendency to follow Bergmann's rule is ancestral for tetrapods, and (2) most extant species that express the rule have retained this tendency from that ancient ancestor. The first inference also generally holds when the discrete data or size-latitude correlation coefficients are analyzed using maximum likelihood, although the results are only statistically significant for some versions of the discrete analyses. The best estimates of ancestral states suggest that the traditional adaptive explanation for Bergmann's rule-conservation of metabolic heat-was not involved in the origin of the trait since that origin predates the evolution of endothermy. A more general thermoregulatory hypothesis could apply to endotherms and some ectotherms, but fails to explain why salamanders have retained Bergmann's rule. Thus, if thermoregulation underlies the origin of a Bergmann's rule tendency, this trait may have been continuously maintained while its cause changed. Alternatively, thermoregulation may not underlie Bergmann's rule in any tetrapod group. The results also suggest that many extinct groups not included in our analyses followed Bergmann's rule.

摘要

生物学中最广为人知的普遍规律之一是伯格曼法则,即观察到在鸟类和哺乳动物物种中,体型往往与环境温度呈负相关。最近的研究表明,龟类和蝾螈也倾向于遵循伯格曼法则,这暗示这种物种水平的趋势在四足动物历史早期就已出现。此外,伯格曼法则的例外情况集中在有鳞目爬行动物(蜥蜴和蛇)中,这表明表现出伯格曼法则渐变群的趋势在物种水平上可能是可遗传的。我们通过将352个物种的体型-纬度关系映射到四足动物系统发育树上,评估了伯格曼法则在物种水平上的遗传性和早期起源。当使用最大可用数据集时,伯格曼法则显示出显著的系统发育信号,表明在物种水平上具有遗传性。这是对一种新兴的物种水平特性遗传性的少数证明之一,也是对生态地理规则的首次证明。当物种被离散编码为显示伯格曼法则或其相反情况时,简约重建表明:(1)遵循伯格曼法则的趋势是四足动物的祖先特征,(2)大多数表现出该法则的现存物种从那个古老祖先那里保留了这种趋势。当使用最大似然法分析离散数据或体型-纬度相关系数时,第一个推断通常也成立,尽管结果仅在某些版本的离散分析中具有统计学意义。对祖先状态的最佳估计表明,对伯格曼法则的传统适应性解释——代谢热的保存——与该特征的起源无关,因为该起源早于恒温动物的进化。一个更普遍的体温调节假说是可以应用于恒温动物和一些变温动物的,但无法解释为什么蝾螈保留了伯格曼法则。因此,如果体温调节是伯格曼法则趋势起源的基础,那么这个特征可能在其原因发生变化的同时一直保持着。或者,体温调节可能在任何四足动物群体中都不是伯格曼法则的基础。结果还表明,许多未纳入我们分析的已灭绝类群遵循伯格曼法则。

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