首页|古生物学中形态空间分析的原理与应用

古生物学中形态空间分析的原理与应用

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生物特征由基因型和表型共同定义.对于由化石主导的古生物学而言,基因信息通常难以获得,因此表型(通常指生物个体的形态)对探讨物种的定义、分类和演化显得尤为重要.早期对形态特征的量化多为直观的线性距离、角度或者面积,随着以界标点和半界标点表示轮廓线的连续性变量和以特征矩阵构建的离散变量等相关量化指标出现,以及针对这些指标所使用的多变量分析方法逐渐发展而形成的几何形态测量学,使得古生物学工作者对形态的认识和研究更为全面.几何形态测量学对生物个体的形态进行量化,随后将降维后的形态学信息投影在二维或三维的空间中形成形态空间,从而研究生物的形态特征以及个体或群落之间的聚集和离散程度.此类分析对生物的演化历程提供了非常重要的信息,尤其可以着重探讨地质历史时期大灭绝事件对生物类群宏演化的影响.本文重点介绍了形态空间的研究历史及其原理与方法,不仅包括对形态的量化、降维的步骤,还涉及差异分析和系统发育比较方法的应用与讨论.此外,本文以贯穿整个显生宙时期舌形贝腕足动物的形态空间分析为例,探讨了形态空间方法在古生物学中的应用和存在的问题,以及其前景与展望.
Principle and application of morphospace in paleontology
Biological information is quintessentially encapsulated by the intricate relationship between genotype—the genetic makeup of an organism—and phenotype,which encompasses the tangible attributes and morphology of the organism.In the realm of paleontology,the genetic information is often difficult to obtain,so the phenotypic traits,specifically the morphology or anatomical shape of organisms,serve as the cornerstones for explicating taxonomic classifications,evolutionary relationships,and functional adaptations of extinct animals.Initial attempts at morphological quantification in paleontology were predominantly qualitative,hinging on non-numeric descriptors of form and structure.When quantitative approaches were applied,they were rudimentary at best,commonly relying on simplistic measurements such as linear distances,angles,or area calculations to provide a semblance of the organism's morphology.These simplistic metrics often fell short of capturing the full complexity of biological shapes,potentially glossing over the subtle variations and nuances that are critically informative about an organism's ecology and evolutionary history.In recent decades,the field of paleontology has witnessed a transformative shiftin the way researchers study and quantify organismal form with the advent of the morphometrics revolution.Beginning in earnest in the 1980s,this innovation in analytical techniques infused the discipline with new methodologies for more precise and nuanced morphological quantification.The inception of geometric morphometrics marked a significant milestone in this trajectory.The appearance of these quantitative indicators,such as(semi)landmarks for continuous variables,and character states for discrete variables,alongside the evolution of multivariate analytical techniques—like principal component analysis(PCA),principal coordinate analysis(PCoA),and nonmetric multidimensional scaling(NMDS)has led to the development of geometric morphometrics,which provides more comprehensive and systematic study of morphology.This method quantifies the morphology of an individual organism and projects it in two-or three-dimensional space to conduct morphospace in order to understand the morphological characteristics and differences between individuals,as well as the similarity and disparity between individuals or communities.Such analyses provide important information on the evolutionary history of organisms during the geological period,such as ammonoids,conodonts and dinosaurs.More importantly,it can explore the impact of mass extinction events on organisms,investigating whether these events exhibit selectivity toward specific morphologies or traits.Given the lengthy history of the morphospace concept and its extensive application in paleontology in recent years to understand the evolutionary trajectories of animal groups,this paper provides a detailed history of relevant studies.It also outlines the principles and basic applications of morphospace,including the processes of quantification and ordination.Moreover,it introduces the fundamental concepts of disparity analysis,which quantifies the similarity among different organisms and groups,as well as discusses various phylogenetic comparative methods employed to investigate the evolution of biological traits,phylogenetic signals,evolutionary patterns and rates,speciation,and interspecies interactions.Additionally,by using lingulid brachiopods as an exemplar—a group of special marine invertebrates that have persisted throughout the entire Phanerozoic yet exhibit limited extant generic and phenotypic diversity,branding them as"living fossils"since Darwin's"On the Origin of Species"in 1859—the results of morphospace quantification,disparity,and phylogenetic comparative analysis suggest that the restricted morphological and ecological variety of modem lingulids likely reflects a disproportionate winnowing of morphospace occupation and ecological disparity due to extinction.Last but not least,this paper critically examines the applications and existing challenges of morphospace,providing insights into its future prospects and potential developments in paleontology.

paleontologymorphospacedisparity analysisphylogenetic comparative methodsbrachiopods

梁悦、张志飞

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西北大学地质学系,大陆动力学国家重点实验室,陕西省早期生命与环境重点实验室,西安 710069

古生物学 形态空间 差异分析 系统发育比较方法 腕足动物

国家重点研发计划国家自然科学基金中国博士后科学基金高等学校学科创新引智计划(111计划)陕西省创新人才推进计划

2023YFF0803601423020102023M732836D170132022TD-11

2024

科学通报
中国科学院国家自然科学基金委员会

科学通报

CSTPCD北大核心
影响因子:1.269
ISSN:0023-074X
年,卷(期):2024.69(12)
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