现生鸟类颈椎形态测定分析及其对中生代鸟类饮食生态的启示

doi:10.19615/j.cnki.2096-9899.240305

摘要/Abstract

摘要：

中生代鸟类的食性推断往往依赖于传统的形态对比和直接的化石证据，但不完整的化石记录和特异埋藏的稀缺性使得我们需要寻找更多有用特征和方法来对化石鸟类进行食性评估。鸟类颈部结构是一个高度模块化、形态和功能密切关联的系统；在恐龙向鸟类演化的过程中，前肢由于飞行适应独立出来，使得颈部需要更多地帮助鸟类实现捕食及其他生态功能。因此，颈部有可能成为鸟类食性推断的显著特征之一。利用形态测定和统计分析，建立起现生鸟类和中生代灭绝鸟类的颈椎形态和食性模式之间的量化关系。基于现生鸟类构建了形态-功能框架，评估了早白垩世热河生物群中发现的5种鸟类的食性模式。结果表明，颈椎形态分异与鸟类取食多样性量化相关，3种反鸟表现出了食虫或食肉鸟类的颈椎形态特征，两种今鸟型类则表现出杂食性或植食性，以及原始的水生适应形态特征。新的分析结果与之前发现的化石直接证据以及其他相关的形态研究基本一致，因此，颈椎作为与取食功能密切相关的骨骼系统，可以为中生代鸟类取食生态的推断提供一定信息。

关键词: 中生代鸟类, 颈椎形态, 食性, 生态分异

Abstract:

The inference of Mesozoic avialan bird diets previously relied on traditional methods such as morphological comparisons among taxa and direct evidence such as identifiable stomach contents. However, the application of these approaches has been limited because of uncommon preservation of relevant fossil evidence. We searched for additional informative characteristics to help develop new methods to assess the diet of fossil birds. In particular, the morphology of the avialan neck is highly modularized and plays roles in multiple functions including food acquisition. The structure of and variation among the cervical vertebrae likely reflects the demands of feeding ecology in fossil and extant birds because the avialan neck evolved to, at least in part, replace the forelimbs by assisting with activities such as cranioinertial feeding and other ecological functions. Here, we utilize morphometric and statistical analyses to establish an initial quantitative relationship between cervical morphology and dietary modes in both extant and extinct birds. This morphometric framework derived from the cervical morphology of living birds is used as a basis to estimate the diet categories of five taxa of Mesozoic birds. The results indicate that there is a quantitative correlation between cervical morphology differentiation and their interrelated feeding modes. The enantiornithine taxa examined exhibit cervical morphologies similar to extant insectivorous or carnivorous birds. The ornithurine species show cervical morphologies that are more aligned with generalist or herbivorous birds, and exhibit preliminary morphological features tied to aquatic adaptions. These findings are consistent in part with other direct fossil evidence, as well as hypotheses developed from other skeletal comparisons. Therefore, the cervical vertebral series, as a skeletal system closely linked to food acquisition, can serve as one of the valuable metrics to provide information for inferring the diet of long extinct Mesozoic birds.

Key words: Mesozoic birds, cervical morphology, diet, ecological variation

中图分类号:

Q915.865

刘碧莹, Thomas A. STIDHAM, 王小平, 李志恒, 周忠和. 现生鸟类颈椎形态测定分析及其对中生代鸟类饮食生态的启示. 古脊椎动物学报, 2024, 62(2): 99-119.

LIU Bi-Ying, Thomas A. STIDHAM, WANG Xiao-Ping, LI Zhi-Heng, ZHOU Zhong-He. Morphometric analysis of the cervical vertebral series in extant birds with implications for Mesozoic avialan feeding ecology. Vertebrata Palasiatica, 2024, 62(2): 99-119.

图/表 6

Fig. 1 Measurements acquired for individual cervical vertebrae The fifth cervical vertebra of Falco subbuteo was used as an exemplar here as showing in ventral (A), dorsal (B), and dorsal-caudal (C) views with measurement axes and angles Abbreviations: CL. centrum length; CW. centrum width; ZA. zygapophyseal angle;ZP. zygapophoseal protrusion

Fig. 2 Comparison of raw measurements from sampled birds The data covers cervical vertebra from C3 to C8. Box plots showing the comparison of cervical change in length (A), width (B), ratio of centrum length to width (C), zygapophyseal angle (D), and zygapophoseal protrusion (E). The black line in the box represents the median value of the measurements for given group, and the endpoint of the upper and lower whiskers represent the maximum and minimum values, respectively Abbreviations: Car. carnivory (red); Ena. enantiornithine (black); Gen. generalist (yellow);Her. herbivore (green); Ins. insectivore (purple); Orn. ornithurine (gray); Pis. piscivore (blue)

Fig. 3 Significant phylogenetic signal detected in cervical vertebral traits using Blomberg’s K Traits with K-value over or equal one indicates strong phylogenetic signal as following browning motion and are phylogenetically conserved. Traits with a K-value smaller than one indicate a departure from Brownian motion. Different categories of measurements are labeled with different colors Abbreviations: A. zygapophyseal angle; L. centrum length; R. ratio of centrum length to width;W. centrum width; ZP. zygapophoseal protrusion

Fig. 4 PCA of the cervical measurements for extant birds only, C3 to C12 covered Abbreviations see Fig. 2. See online supplementary file 1 for details on loadings and percentages of variance explained (Fig. S2)

Fig. 5 PCA results with both extant and extinct birds, C3 to C8 covered Abbreviations see Fig. 2. See online supplementary file 1 for details on loadings and percent variance explained (Fig. S3)

Fig. 6 Phylogenetic PCA for extant and extinct birds with data for vertebrae C3 to C8 Variables in Fig. 3 were involved with this pPCA analysis. Abbreviations see Fig. 2

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