早期生命形态演化研究中的定量方法

定量古生物学是现代古生物学的一个分支,提倡用定量的手段来研究地质历史时期生命的演化过程。我国从事定量古生物研究的群体较小,特别是对前寒武纪早期生命演化的定量研究还没有系统地展开。这篇文章将主要介绍如何利用定量手段来研究前寒武纪化石的形态演化。对于前寒武纪化石,由于大部分化石分类属性的不确定性,通常使用几何性状对化石的最基本形态结构进行分析,并用存在/缺失(1/0)这种离散变量对每个性状进行量化。非参数多维标量分析方法[Non-parametric multidimensional scaling analysis(MDS)]可以将高维度的离散数据投影到二维或者三维的形态空间上,进而探讨生物群在形态空间中所占有的范围;由离散变量计算得出的生物群的表形分异度(morphological disparity)可以用MDS方差或者平均差异参数[Mean dissimilarity coeffi-cient(MDC)]来计算。形态空间的范围(morphospace range)和表形分异度是相互联系的,如果形态空间范围是固定的,那么表形分异度实际上代表了生物群在形态空间中的分布密度。在解释数据之前,需要对可能存在的样本效应进行测试。常用的方法包括稀释法(rarefaction)、随机取样法(randomization)和自举法(bootstrapping)等。为了帮助读者进一步了解这些方法的使用,文中列举了三个实例:伊迪卡拉生物的形态演化,元古代宏观藻类的形态演化和元古代及寒武纪疑源类的演化。     

A new type of Precambrian megascopic fossils: the Jinxian biota from northeastern China

Precambrian fossils are crucial for our understanding of the evolution of early organisms. Megascopic body fossils are more important because they potentially represent macroorganisms. However, the Precambrian fossil record is sparse and dominated by microfossils and microbial structures. Here we show a new type of megascopic fossils recovered from the Xingmincun Formation (probably Neoproterozoic age), northeastern China. The specimens are flat, flexible (easily corrugated) and discoidal in outline. Concentric or spiral ridges are preserved on both sides. Petrographical thin section examination indicates that the specimen consist of a thin layer of microcrystalline quartz grains (about 20–30 μm thick) wrapped by an outer sheath, composed primarily of chlorites. Field Emission Scanning Electron Microscopy (FE-SEM) coupled with an x-ray energy dispersive spectrometer system (EDX) analysis shows microstructures and relative element abundance of the fossils, but contributes little in solving their biological affinities. The fossils have previously been linked to discoidal impressions of the Ediacara biota. Close examination on new materials indicates that they are radically different from either the Ediacara impressions or any other Precambrian megascopic remains. Concentric or spiral ridges may result from rhythmic growth and the presence of twin specimens may suggest that the organisms undergo asexual reproduction or inhibition of growth in one direction. Referring them to any known fossil or living group has proved to be difficult. We conclude that they represent a distinct group of Precambrian megascopic organisms regardless of their affinities remaining problematic.     

The rise of bilaterians: a few closing comments

Freeman (2009, 2010) argue that Ediacara “faunas” should be analysed using “promorphologies” of extant animals, that rangeomorphs and erniettomorphs are colonial animals, and that Vernanimalcula is a bilaterian animal. Recent studies of Ediacara fossils have provided multiple lines of evidence that these fossils represent a sample of phylogenetically diverse marine organisms that included crown-group animals as well as stem-group animals and non-animals. Thus, it is inappropriate to analyse Ediacara fossils using “promorphologies” of extant animals. The interpretation of rangeomorphs and erniettomorphs as colonial animals is inconsistent with the functional morphologies of these Ediacara fossils. Although Vernanimalcula is a fossil, the purported germ layers of Vernanimalcula are of diagenetic origin and there is no morphological evidence in support of its bilaterian affinity.     

The origin of the animals and a ‘Savannah’ hypothesis for early bilaterian evolution

The earliest evolution of the animals remains a taxing biological problem, as all extant clades are highly derived and the fossil record is not usually considered to be helpful. The rise of the bilaterian animals recorded in the fossil record, commonly known as the ‘Cambrian explosion’, is one of the most significant moments in evolutionary history, and was an event that transformed first marine and then terrestrial environments. We review the phylogeny of early animals and other opisthokonts, and the affinities of the earliest large complex fossils, the so‐called ‘Ediacaran’ taxa. We conclude, based on a variety of lines of evidence, that their affinities most likely lie in various stem groups to large metazoan groupings; a new grouping, the Apoikozoa, is erected to encompass Metazoa and Choanoflagellata. The earliest reasonable fossil evidence for total‐group bilaterians comes from undisputed complex trace fossils that are younger than about 560 Ma, and these diversify greatly as the Ediacaran–Cambrian boundary is crossed a few million years later. It is generally considered that as the bilaterians diversified after this time, their burrowing behaviour destroyed the cyanobacterial mat‐dominated substrates that the enigmatic Ediacaran taxa were associated with, the so‐called ‘Cambrian substrate revolution’, leading to the loss of almost all Ediacara‐aspect diversity in the Cambrian. Why, though, did the energetically expensive and functionally complex burrowing mode of life so typical of later bilaterians arise? Here we propose a much more positive relationship between late‐Ediacaran ecologies and the rise of the bilaterians, with the largely static Ediacaran taxa acting as points of concentration of organic matter both above and below the sediment surface. The breaking of the uniformity of organic carbon availability would have signalled a decisive shift away from the essentially static and monotonous earlier Ediacaran world into the dynamic and burrowing world of the Cambrian. The Ediacaran biota thus played an enabling role in bilaterian evolution similar to that proposed for the Savannah environment for human evolution and bipedality. Rather than being obliterated by the rise of the bilaterians, the subtle remnants of Ediacara‐style taxa within the Cambrian suggest that they remained significant components of Phanerozoic communities, even though at some point their enabling role for bilaterian evolution was presumably taken over by bilaterians or other metazoans. Bilaterian evolution was thus an essentially benthic event that only later impacted the planktonic environment and the style of organic export to the sea floor.     

Das Präkambrium. Die Wiege des Lebens

The Precambrian is the cradle of life. With a time span of about 4 Billion years it represents the largest part of earth history. Life changed the planet during the Precambrian by a lot of interactions with plate tectonics and raised into better qualities. A special milestone was the release of free oxygen by the stromatolithes at about 2.5 Billion years. An extreme bottleneck for the evolution of life was the Snowball Earth representing the freezing of the entire earth surface and the covering by an ice sheet. Plate tectonic processes were responsible for the melting of the ice sheet. In the aftermath of that glaciation the rapid radiation of the first complex higher life forms begun. These were represented by the so‐called Ediacara Biota, which occurred in the time span of about 630 and 543 Million years before today. The Ediacara Biota are unique in the evolution of life and existed in a close interaction with a leather‐like biomat at the sea‐floor which provided stability, hide and food. Among the Ediacara Biota the first primitive arthropods, the molluscs and the anthozoans occurred. In addition, in the fossil record are reported a lot of mystic life forms without a good or any classification. The Ediacara Biota represent the critical evolutionary step to pave the way for the explosion‐like radiation of life during the Cambrian that started at 542 Million years before present.     

Downsized Dinosaurs: The Evolutionary Transition to Modern Birds

Living birds are the most diverse land vertebrates and the heirs of a rich chapter in the evolution of life. The origin of modern birds from animals similar to Tyrannosaurus rex is among the most remarkable examples of an evolutionary transition. A wealth of recently discovered fossils has finally settled the century-old controversy about the origin of birds and it has made the evolutionary saga toward modern birds one of the best documented transitions in the history of life. This paper reviews the evidence in support of the origin of birds from meat-eating dinosaurs, and it highlights the array of fossils that connect these fearsome animals with those that fly all around us.     

GEOMETRIC MORPHOMETRICS OF THE SKULL ROOF OF STEREOSPONDYLS (AMPHIBIA: TEMNOSPONDYLI)

Abstract:  Geometric morphometric analysis using relative warps is applied to the skull roof of 62 species of stereospondyls and their closest outgroups (i.e. basal archegosauriforms) from among temnospondyl amphibians. Twenty-one landmarks and five taxonomic groups are used for comparisons. Their skull evolution is quantified in a morphospace defined by two relative warps axes. The majority of groups show poor concordance between morphological and phylogenetic distances. The only exception is represented by Yates and Warren's study of stereospondyl relationships, in which concordance is high. Only basal archegosauriforms and rhinesuchids show significant overlap in morphospace, although this might be due to low sample sizes. Regression of estimated mean disparity against taxon sample size shows that species within both the trematosauroid and the rhytidostean groups are more widely dispersed in morphospace than species belonging to any of the remaining stereospondyl groups. Stereospondyl skull evolution was characterized by divergence between major clades and convergence within those clades. Changes in patterns of morphospace occupation through time agree with the hypothesis of an 'explosive' radiation in the early Early Triassic, after the extinction of basal archegosauriforms at the end of the Permian.     

寒武纪早期岩家河生物群:研究进展和展望*

埃迪卡拉纪-寒武纪转换时期动物的起源、演化和"寒武纪大爆发"一直是国际古生物学界研究的热点问题,其中寒武纪早期小壳化石群与埃迪卡拉纪化石群和寒武纪早期澄江化石库之间的内在关系是古生物学界研究的难题,其主要原因是寒武纪早期与小壳化石群伴生的宏体动、植物化石的缺乏。发现于峡东地区的寒武纪早期岩家河生物群填补了这一缺失环节,该生物群包含宏体动物、宏观藻类、小壳化石、球形化石(可能的胚胎化石)、微古植物和蓝菌类等化石,部分宏体化石显示了从埃迪卡拉纪向寒武纪过渡色彩。化石保存方式有碳质膜、黄铁矿化、磷酸盐化、硅化。因此对岩家河生物群生物多样性和埋藏学进行综合研究,将可提供纽芬兰世(梅树村期)碳酸盐台地—碳酸盐台地内部的局部凹陷盆地相的一个较完整的生物景观图,对探索"寒武纪大爆发主幕"前夕生物的辐射、演化模式及保存机制具有重要的科学意义。     

Evolution of Cranial Shape in Caecilians (Amphibia: Gymnophiona)

Insights into morphological diversification can be obtained from the ways the species of a clade occupy morphospace. Projecting a phylogeny into morphospace provides estimates of evolutionary trajectories as lineages diversified information that can be used to infer the dynamics of evolutionary processes that produced patterns of morphospace occupation. We present here a large-scale investigation into evolution of morphological variation in the skull of caecilian amphibians, a major clade of vertebrates. Because caecilians are limbless, predominantly fossorial animals, diversification of their skull has occurred within a framework imposed by the functional demands of head-first burrowing. We examined cranial shape in 141 species, over half of known species, using X-ray computed tomography and geometric morphometrics. Mapping an existing phylogeny into the cranial morphospace to estimate the history of morphological change (phylomorphospace), we find a striking pattern: most species occupy distinct clusters in cranial morphospace that closely correspond to the main caecilian clades, and each cluster is separated by unoccupied morphospace. The empty spaces in shape space are unlikely to be caused entirely by extinction or incomplete sampling. The main caecilian clades have different amounts of morphological disparity, but neither clade age nor number of species account for this variation. Cranial shape variation is clearly linked to phyletic divergence, but there is also homoplasy, which is attributed to extrinsic factors associated with head-first digging: features of caecilian crania that have been previously argued to correlate with differential microhabitat use and burrowing ability, such as subterminal and terminal mouths, degree of temporal fenestration (stegokrotaphy/zygokrotaphy), and eyes covered by bone, have evolved and many combinations occur in modern species. We find evidence of morphological convergence in cranial shape, among species that have eyes covered by bone, resulting in a narrow bullet-shaped head. These results reveal a complex history, including early expansion of morphospace and both divergent and convergent evolution resulting in the diversity we observe today.     

Global marine redox changes drove the rise and fall of the Ediacara biota

The role of O₂ in the evolution of early animals, as represented by some members of the Ediacara biota, has been heavily debated because current geochemical evidence paints a conflicting picture regarding global marine O₂ levels during key intervals of the rise and fall of the Ediacara biota. Fossil evidence indicates that the diversification the Ediacara biota occurred during or shortly after the Ediacaran Shuram negative C‐isotope Excursion (SE), which is often interpreted to reflect ocean oxygenation. However, there is conflicting evidence regarding ocean oxygen levels during the SE and the middle Ediacaran Period. To help resolve this debate, we examined U isotope variations (δ²³⁸U) in three carbonate sections from South China, Siberia, and USA that record the SE. The δ²³⁸U data from all three sections are in excellent agreement and reveal the largest positive shift in δ²³⁸U ever reported in the geologic record (from ~ ?0.74‰ to ~ ?0.26‰). Quantitative modeling of these data suggests that the global ocean switched from a largely anoxic state (26%–100% of the seafloor overlain by anoxic waters) to near‐modern levels of ocean oxygenation during the SE. This episode of ocean oxygenation is broadly coincident with the rise of the Ediacara biota. Following this initial radiation, the Ediacara biota persisted until the terminal Ediacaran period, when recently published U isotope data indicate a return to more widespread ocean anoxia. Taken together, it appears that global marine redox changes drove the rise and fall of the Ediacara biota.     

Convergence and divergence in the evolution of cat skulls: temporal and spatial patterns of morphological diversity

Background

Studies of biological shape evolution are greatly enhanced when framed in a phylogenetic perspective. Inclusion of fossils amplifies the scope of macroevolutionary research, offers a deep-time perspective on tempo and mode of radiations, and elucidates life-trait changes. We explore the evolution of skull shape in felids (cats) through morphometric analyses of linear variables, phylogenetic comparative methods, and a new cladistic study of saber-toothed cats.

Methodology/Principal Findings

A new phylogenetic analysis supports the monophyly of saber-toothed cats (Machairodontinae) exclusive of Felinae and some basal felids, but does not support the monophyly of various saber-toothed tribes and genera. We quantified skull shape variation in 34 extant and 18 extinct species using size-adjusted linear variables. These distinguish taxonomic group membership with high accuracy. Patterns of morphospace occupation are consistent with previous analyses, for example, in showing a size gradient along the primary axis of shape variation and a separation between large and small-medium cats. By combining the new phylogeny with a molecular tree of extant Felinae, we built a chronophylomorphospace (a phylogeny superimposed onto a two-dimensional morphospace through time). The evolutionary history of cats was characterized by two major episodes of morphological divergence, one marking the separation between saber-toothed and modern cats, the other marking the split between large and small-medium cats.

Conclusions/Significance

Ancestors of large cats in the ‘Panthera’ lineage tend to occupy, at a much later stage, morphospace regions previously occupied by saber-toothed cats. The latter radiated out into new morphospace regions peripheral to those of extant large cats. The separation between large and small-medium cats was marked by considerable morphologically divergent trajectories early in feline evolution. A chronophylomorphospace has wider applications in reconstructing temporal transitions across two-dimensional trait spaces, can be used in ecophenotypical and functional diversity studies, and may reveal novel patterns of morphospace occupation.     

Evolutionary Transitions in the Fossil Record of Terrestrial Hoofed Mammals

In the past few decades, many new discoveries have provided numerous transitional fossils that show the evolution of hoofed mammals from their primitive ancestors. We can now document the origin of the odd-toed perissodactyls, their early evolution when horses, brontotheres, rhinoceroses, and tapirs can barely be distinguished, and the subsequent evolution and radiation of these groups into distinctive lineages with many different species and interesting evolutionary transformations through time. Similarly, we can document the evolution of the even-toed artiodactyls from their earliest roots and their great radiation into pigs, peccaries, hippos, camels, and ruminants. We can trace the complex family histories in the camels and giraffes, whose earliest ancestors did not have humps or long necks and looked nothing like the modern descendants. Even the Proboscidea and Sirenia show many transitional fossils linking them to ancient ancestors that look nothing like modern elephants or manatees. All these facts show that creationist attacks on the fossil record of horses and other hoofed mammals are completely erroneous and deceptive. Their critiques of the evidence of hoofed mammal evolution are based entirely on reading trade books and quoting them out of context, not on any firsthand knowledge or training in paleontology or looking at the actual fossils.     

湖北宜昌乔家坪村寒武系水井沱组海绵动物化石群

海绵动物是最原始的动物门类之一,其演化史对于研究多细胞动物的起源和早期演化至关重要.现有的化石记录显示,水体较深的贫氧环境可能是寒武纪早期海绵动物演化的重要场所之一.前人在贵州、湖南、安徽等多地的斜坡至盆地相黑色页岩中已经报道了数个海绵化石群.在这样的环境中,由于水动力作用较弱,海绵骨架结构容易得到原位保存.湖北宜昌秭...     

Skeletal Nets of the Ediacaran Fronds

The Ediacara fauna is traditionally regarded as the first complex, diverse and widespread macroscopic life. The uncertainty of systematic position of its members has led to very different views on the early evolution of metazoans. In part, this may be due to a lack of data on sclerotization: a hard skeleton is a part of the archetype of the most taxa known from the fossil record, whereas the Ediacara fauna as a whole is most often considered soft-bodied, which complicates comparison. Here we report the Late Precambrian frond-like fossils (Petalonamae) from the Vendian assemblage of the Southeastern White Sea area (∼555.3 Ma), which show evidence of elaborate skeleton composed of a regular meshwork reinforced by dense longitudinal and circular bands. Judging from the nature of preservation and the dynamics of the environment Ediacaran fronds secreted a relatively rigid but flexible skeleton. The fact that frond-like Petalonamae had a supporting structure similar to that of sponges and cnidarians seems to be a powerful argument in favor of their metazoan affinity. The new observations indicated also that the widespread skeletonization had occurred long before the “Cambrian skeletal revolution”.     

The evolution of cell types in animals: emerging principles from molecular studies

Cell types are fundamental units of multicellular life but their evolution is obscure. How did the first cell types emerge and become distinct in animal evolution? What were the sets of cell types that existed at important evolutionary nodes that represent eumetazoan or bilaterian ancestors? How did these ancient cell types diversify further during the evolution of organ systems in the descending evolutionary lines? The recent advent of cell type molecular fingerprinting has yielded initial insights into the evolutionary interrelationships of cell types between remote animal phyla and has allowed us to define some first principles of cell type diversification in animal evolution.     

Horseshoe crab phylogeny and independent colonizations of fresh water: ecological invasion as a driver for morphological innovation

Xiphosurids are an archaic group of aquatic chelicerate arthropods, generally known by the colloquial misnomer of ‘horseshoe crabs’. Known from marine environments as far back as the early Ordovician, horseshoe crabs are generally considered ‘living fossils’ – descendants of a bradytelic lineage exhibiting little morphological or ecological variation throughout geological time. However, xiphosurids are known from freshwater sediments in the Palaeozoic and Mesozoic; furthermore, the contention that xiphosurids show little morphological variation has never been tested empirically. Attempts to test this are hampered by the lack of a modern phylogenetic framework with which to explore different evolutionary scenarios. Here, I present a phylogenetic analysis of Xiphosurida and explore patterns of morphospace and environmental occupation of the group throughout the Phanerozoic. Xiphosurids are shown to have invaded non‐marine environments independently at least five times throughout their evolutionary history, twice resulting in the radiation of major clades – bellinurines and austrolimulids – that occupied novel regions of morphospace. These clades show a convergent ecological pattern of differentiation, speciation and subsequent extinction. Horseshoe crabs are shown to have a more dynamic and complex evolutionary history than previously supposed, with the extant species representing only a fraction of the group's past ecological and morphological diversity.     

Slime travelers: Early evidence of animal mobility and feeding in an organic mat world

Mobility represents a key innovation in the evolution of complex animal life. The ability to move allows for the exploration of new food sources, escapes from unfavorable environmental conditions, enhanced ability to exchange genetic material, and is one of the major reasons for the diversity and success of animal life today. The oldest widely accepted trace fossils of animal mobility are found in Ediacaran‐aged rocks (635–539 Ma). The earliest definitive evidence for movement associated with exploitation of resources for feeding occurs in the White Sea assemblage of the Ediacara Biota—macroscopic, soft‐bodied fossils of Ediacaran age. Here, we evaluate potential support for mobility in dickinsoniomorphs, presenting new data regarding abundant Dickinsonia and associated trace fossils from the Ediacara Member, South Australia. Results quantitatively demonstrate that Dickinsonia was capable of mobility on relatively short, ecological timescales. This organism was bilaterally symmetrical, likely moved via muscular peristalsis, and left trace fossils due to active removal of the organic mat related to feeding. Analogous structures associated with Yorgia indicate that it was also mobile and fed in a similar manner. Morphological evidence suggests that two other modular taxa, Andiva and Spriggina, were able to move but did not feed in a manner that impacted the organic mat. Together, these data suggest that mobility was present in multiple disparate bilaterally symmetrical Ediacaran taxa.     

Resolving the first steps to multicellularity

Multicellular life has evolved many times, yet each origin requires free cells to integrate unselfishly into a higher-level individual. How can such transitions evolve? In a new paper, Herron and Michod investigate the recent origins of multicellularity in colonial algae. Their phylogenetic reconstructions provide a striking dissection of early steps, and altruistic traits are at the crux of it. Key evolutionary reversals are also revealed, where cellular selfishness might have thwarted multicellular integration.     

Cnidarian taphonomy and affinities of the Ediacara biota

Plaster impressions and sand casts of extant medusae, a chondrophoran, and a pennatulid share basic structural characteristics with fossils in the Upper Proterozoic Ediacara assemblage. Impressions of extant medusae and Proterozoic circular impressions show general similarities in arrangement and position of radial and concentric structures and a central raised boss. However, annular rings and radial grooves are more numerous in the Proterozoic fossils and strongly folded or deformed fossils are rare as compared with impressions of modem medusae. Recent pennatulids yield impressions that are more deformed and irregular than the Proterozoic genus Charniodiscus. The greater frequency of deformation of most simulated fossil medusoids relative to Precambrian circular impressions implies that Proterozoic medu-soids were substantially stiffer than many modern taxa of comparable sizes. Many fossils with abundant circular rings have no constructional counterparts among the extant forms studied here and their medusoid affinities should remain in doubt. The structural simplicity of impressions of Ediacara organisms and extant cnidarians suggests that their mutual similarities may be due to convergence. However, there is no compelling morphological reason to reject the claim that some Proterozoic fossils may share affinities with living cnidarians. □ Taphonomy. Ediacara biota, cnidarians, phylogenetic relationships.     

The puzzle of the Krebs citric acid cycle: Assembling the pieces of chemically feasible reactions,and opportunism in the design of metabolic pathways during evolution

The evolutionary origin of the Krebs citric acid cycle has been for a long time a model case in the understanding of the origin and evolution of metabolic pathways: How can the emergence of such a complex pathway be explained? A number of speculative studies have been carried out that have reached the conclusion that the Krebs cycle evolved from pathways for amino acid biosynthesis, but many important questions remain open: Why and how did the full pathway emerge from there? Are other alternative routes for the same purpose possible? Are they better or worse? Have they had any opportunity to be developed in cellular metabolism evolution? We have analyzed the Krebs cycle as a problem of chemical design to oxidize acetate yielding reduction equivalents to the respiratory chain to make ATP. Our analysis demonstrates that although there are several different chemical solutions to this problem, the design of this metabolic pathway as it occurs in living cells is the best chemical solution: It has the least possible number of steps and it also has the greatest ATP yielding. Study of the evolutionary possibilities of each one-taking the available material to build new pathways-demonstrates that the emergence of the Krebs cycle has been a typical case of opportunism in molecular evolution. Our analysis proves, therefore, that the role of opportunism in evolution has converted a problem of several possible chemical solutions into asingle-solution problem, with the actual Krebs cycle demonstrated to be the best possible chemical design. Our results also allow us to derive the rules under which metabolic pathways emerged during the origin of life.