早期节肢动物化石对演化发育生物学的启示

基于特定遗传机制而发生的生物演化过程是生命本质所在,是一个能引起争论和激励思维的研究领域。我国帽天山页岩动物群保存的“中间环节”化石不仅为揭示节肢动物和脊椎动物等重大生物类群起源和早期演化之谜提供可供追溯的实证,而且可能成为探索基因调控机制演化与形态发生的模型动物。生物发育调控基因及其调控机制的变化是导致形态变异的根本原因。因此新的研究领域——演化发育生物学的产生使地质学和生物学联合研究动物起源和演化机制以及某些调控基因的起源和演化成为可能。本研究依据古生物学新的化石发现和研究成果,其中包括最新提出的关于节肢动物头、眼柄、前附肢的原型及其演化模式,分析其对演化发育生物学的影响和启示,并尝试为发育生物学提出新的前瞻性研究课题。因此,早期节肢动物化石的研究不仅将古生物学、演化生物学和发育生物学研究融为一体,而且为发育生物学基因层面研究和古生物学演化层面研究的融合找到切入点。     

动物的起源和寒武纪大爆发:来自中国的化石证据

近30年来,中国埃迪卡拉纪—寒武纪发现了包括瓮安生物群和澄江动物群在内一系列重要化石群,为揭示动物的起源和寒武纪大爆发过程提供了全球独一无二的古生物学证据。文中全面而简洁地介绍发现于华南埃迪卡拉纪—寒武纪的所有化石群和重要动物化石发现,总结其中特异化石群保存类型和埋藏环境。其中,陡山沱型磷酸盐化胚胎化石群、庙河型宏体碳质膜化石群和牛蹄塘型海绵动物群,以及陡山沱组硅化的动物休眠胚胎化石和宽川铺动物群中"rsten"型化石均是全球这一时期华南独有的特异埋藏化石群。从时空两方面对化石群进行综合分析认为,动物可靠的最早化石记录发现于埃迪卡拉纪早期,成冰纪及其以前的动物化石均存在疑问。动物的起源和寒武纪大爆发经历了从浅水逐步向深水迁移,以及阶段性辐射和灭绝的过程。首先动物经历了在埃迪卡拉纪早期的隐形微体动物早期演化阶段,之后经过以埃迪卡拉型生物为代表的大型复杂化演化阶段,在寒武纪初期发生以梅树村动物群为代表的两侧对称动物和生物骨骼矿化的大爆发,并随后又被以澄江动物群为代表的寒武纪动物群所替代,在筇竹寺期达到了动物造型和生态系统复杂化的高峰。     

动物的起源和寒武纪大爆发：来自中国的化石证据北大核心

近30年来,中国埃迪卡拉纪—寒武纪发现了包括瓮安生物群和澄江动物群在内一系列重要化石群,为揭示动物的起源和寒武纪大爆发过程提供了全球独一无二的古生物学证据。文中全面而简洁地介绍发现于华南埃迪卡拉纪—寒武纪的所有化石群和重要动物化石发现,总结其中特异化石群保存类型和埋藏环境。其中,陡山沱型磷酸盐化胚胎化石群、庙河型宏体碳质膜化石群和牛蹄塘型海绵动物群,以及陡山沱组硅化的动物休眠胚胎化石和宽川铺动物群中"rsten"型化石均是全球这一时期华南独有的特异埋藏化石群。从时空两方面对化石群进行综合分析认为,动物可靠的最早化石记录发现于埃迪卡拉纪早期,成冰纪及其以前的动物化石均存在疑问。动物的起源和寒武纪大爆发经历了从浅水逐步向深水迁移,以及阶段性辐射和灭绝的过程。首先动物经历了在埃迪卡拉纪早期的隐形微体动物早期演化阶段,之后经过以埃迪卡拉型生物为代表的大型复杂化演化阶段,在寒武纪初期发生以梅树村动物群为代表的两侧对称动物和生物骨骼矿化的大爆发,并随后又被以澄江动物群为代表的寒武纪动物群所替代,在筇竹寺期达到了动物造型和生态系统复杂化的高峰。     

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。     

化石记录的再评估

分子生物学和分支系统学的发展以及化石记录自身的局限性使得古生物地层资料在生物演化研究中的地位遭到质疑。连续的化石记录的发现肯定了化石分析的意义,数据库的建立与定量分析方法的应用为古生物学的研究增添了新生力量。     

古生物学研究中的采样问题及多样性估计方法——以华夏正形贝动物群为例

传统的古生物学采样方法通常很少对同一化石层面展开进行,由此得到的多样性信息往往难以反映化石组合的真实情况。通过观察稀疏化曲线平缓程度来判断化石采样是否充分的经验方法并不严谨。本文在介绍古生物化石采样方法的基础上,提出采样是否充分的本质是采样获得的多样性与所属化石群落潜在多样性之间的接近程度。借鉴现代生物学中的多样性估计方法,以华夏正形贝腕足动物群为例,本研究采用EstimateS软件,定量估计4个化石点的潜在多样性;探讨客观判断采样充分的方法,同时通过曲线拟合法预测达到特定多样性可能需要的样本数量,为二次研究提供重要信息。该多样性估计方法是古生物研究中一个较新的领域,在多样性对比及特定化石群的多样性评估中均有较好应用前景。为得到更全面的多样性信息,文章建议相关研究在使用稀疏化的同时辅以多样性估计方法。     

地点覆盖度、集合群落、地质时期动物群:联接古生物学和现代种群生物学的概念

在古生物群落中新近出现的地点覆盖度或发生率的概念,以及对生物集合种群的空间动态学的深入了解,使古生物学和现代生物学的联系更加紧密。集合群落的概念以及地质时期动物群的古生物学认识显示,将古生物学和生态学结合在一起是大有希望的。评述了目前对这些概念的理解,并以欧亚大陆大型食草动物群落的发育作为范例,介绍了运用这些理论来研究化石记录的方法。研究结果表明,化石记录中种群覆盖度的研究可以在以化石为基础的进化研究和以现生有机体为基础的进化研究之间架起一座桥梁。地点覆盖度最大的类群,像现代生态学中发生率最大的类群一样,往往是地理分布最广泛、在当地最丰富的类群。     

沙河街组介形虫化石群定量分析方法及其地质意义

本文通过对山东济阳坳陷沙河街组含介形虫化石岩心样品进行散样方法试验,表明水煮法、汽油法是第三纪地层微古分析行之有效的样品处理方法,利用这种方法可以获得大量的数据,从而可以进行介形虫等微体化石群的定量分析。文中对沙河街组介形虫化石群的一个组合和三个亚组合进行了定量分析。分析结果表明,较之以前定性描述,定量分析更能完整、准确地反映化石群面貌,并可进行种群结构、群落结构研究,从而为进行沙河街组沉积时期的     

江西玉山上奥陶统凯迪阶三衢山组的丛花海绵(Corymbospongia)*

串管海绵在中–晚奥陶世经历了的首次辐射演化事件, 在北美、新南威尔士、哈萨克斯坦、西伯利亚和中国西北等地的上奥陶统地层中皆有报道, 但研究程度较低。本文系华南奥陶系串管海绵首次系统古生物学的报道, 详细描述江西玉山上奥陶统凯迪阶三衢山组的Corymbospongia (丛花海绵属)化石, 并基于C. amplia Rigby, Karl, Blodgett & Baichtal, 2005和C. mica Rigby & Potter, 1986的标本展示该属化石外壁的微孔等特征性结构, 这2个种具有跨板块的生物地理分布。综合串管海绵产出层的碳酸盐岩微相以及伴生钙藻和珊瑚化石等群落分子的指相参照, 识别C. amplia和C. mica均具有明显的生态专属性: C. amplia为造礁生物, 在小型微生物岩中出现; C. mica则代表适应于更浅水生境的平地群落代表。本研究将为从底栖生物群落角度开展扬子区生物地层学及古地理重建提供进一步的依据。     

澄江动物群研究与进展

介绍云南澄江动物化石群的发现过程、生物组成及埋藏模式,并分析化石动物多样化的原因。综述自1984年发现192属228种生物化石以来取得的一些研究新进展。     

Influence of transportation and time-averaging in fossil assemblages from the Pennsylvanian of Oklahoma

The relative effects of shell destruction, transportation, and time-averaging have been assessed in seventeen soft-bottom, benthic fossil collections from the Pennsylvanian Wewoka Formation in Hughes County, Oklahoma. Invertebrates were identified to species level, and each was subject to a taphonomic evaluation to determine whether it was in place. Species that were not deemed parautochthonous (living at the location of collection shortly before final burial) were eliminated from the faunal lists to reduce the effects of time-averaging and transportation in the fossil assemblages. Both reciprocal averaging and non-metric multidimensional scaling ordinations revealed two biofacies. These were distinguished using both presence/absence data and numerical counts of species despite an overwhelming number of taxa occurring in both biofacies. Comparison between ordinations based on entire faunal lists with those based on parautochthonous faunal lists indicates that the two biofacies faithfully reproduce differences in the living distributions of member taxa but that the two biofacies underwent different degrees of time-averaging and transportation. Time-averaging leads to accumulation of taxa over time, thereby making presence/absence composition of collections more similar. Transportation, on the other hand, homogenizes originally patchy population distributions and makes relative abundances more uniform. Based on differences in abundances and presence/absence patterns, the stratigraphically lower biofacies underwent more transportation and more time-averaging than the upper biofacies.     

云南早寒武世澄江化石库中两种埋藏相的化石定量分析

通过昆明海口马房村同一个采坑的化石连续采集和统计,对澄江化石库中两种埋藏相(以事件层泥岩和背景层泥岩所代表)内化石保存和组合特征进行定量研究,结果表明:(1)事件层内化石量并不随单一事件层厚度变化而变化。物种在剖面上不同层位的事件层内分布不均匀,表现为单层富集现象;而背景层内的化石含量与背景层的沉积厚度有关,背景层的沉积厚度越厚,化石数量就越多。(2)事件层与背景层内化石组合和结构存在很大差异。事件层内化石组合与背景层内化石组合相比具有更多的物种组成和更复杂的古群落结构;而背景层内化石组合面貌单调,在剖面上保持稳定。(3)事件层内化石生物体腐烂发生在埋藏之后,绝大多数化石生物是被活埋后保存下来的;而背景层内化石主要表现为外壳和弱矿化有机质生物表皮在海床表面长时间停滞而堆积形成,明显受到时间均衡作用(time-average)的影响。(4)事件层内化石在埋藏前无搬运作用,或发生近原栖息地的快速搬运,埋藏后无生物扰动。背景层内大量壳体相连的双瓣壳节肢动物化石和完整海绵体化石的存在则表明,背景层代表一个低氧、低能并具有有限生物扰动的沉积环境。     

Significance of time-averaging for palaeocommunity analysis

Fürsich, F. T. & Aberhan, M. 1990 04 15: Significance of time-averaging for palaeo community analysis. Lethaia , Vol. 23, pp. 143–152. Oslo. ISSN 0024–1164.
Time-averaging caused by sedimentological, biological, or digenetic factors is widespread in marine environments. It can be recognized using stratigraphic, ecological, taphonomic, or sedimcntologic criteria. Community parameters such as diversity, relative abundance of species, taxonomic and, to a lesser extent, trophic and life habit composition may seriously differ between thanatocoenosis (death assemblage) and time-averaged taphocoenosis, depending on the degree of community stability. Time-averaged community relics have a quality different from that of Recent communities and any comparison between the two has to take this into account. Whereas allochthonous time-averaging, which is strongly related to the energy level, is particularly common in very shallow environments, the importance of autochthonous time-averaging, related in addition to the rate of net sedimentation, increases with distance from the shore and with depth. Short-term time-averaging (in the order of up to several thousand years) prevails in near shore shallow environments, whilst long-term time-averaging (in the order of 10⁴ to 10⁵ years) becomes more important towards lower shelf and deep sea environments.     

An ecological reassessment of the southern African carnivore guild: a case study from Member 4, Sterkfontein, South Africa

The southern African late Pliocene to early Pleistocene carnivore guild was much larger than that of the present day. Understanding how this guild may have functioned is important for the reconstruction of carnivore-hominin interactions and to assess the potential for hominin scavenging in southern Africa. In modern ecosystems, the coexistence of larger carnivore species is constrained by several factors, which include high levels of interspecific competition. Here, the composition of the fossil carnivore guild is examined using Sterkfontein Member 4 (Cradle of Humankind, South Africa) as a case study. Sterkfontein Member 4 contains 10 larger carnivore taxa (body mass > 21.5 kg) and may also contain two Australopithecus species. Two possible causes of higher numbers of carnivore species in the South African fossil record are initially considered. First, that there is a bias introduced through comparing assemblages of differing sizes; second, carnivore biodiversity may have been artificially inflated due to previous taxonomic splitting of carnivore species, such as Crocuta. These possibilities are rejected and modern ecological data are used to construct a simple spatial model to determine how many carnivores could have co-existed. Although the resulting model indicates that the carnivore taxa present in Member 4 could have co-occurred, modern ecological studies indicate that it is highly unlikely that they would have co-existed simultaneously. Considering the complex depositional processes that operate in the southern African cave sites, it is proposed that the larger carnivore guild observed in the Sterkfontein Member 4 fossil assemblage is a palimpsest created by time-averaging. In light of this, we suggest that sites which have a large number of carnivore taxa should be examined for time-averaging, while those sites which have relatively few species may be a better reflection of carnivore communities.     

Shell survival and time-averaging in nearshore and shelf environments: estimates from the radiocarbon literature

Radiocarbon dates provide a means for estimating the time a shell may persist in active sedimentary environments and the actual temporal extent of time-averaging in marine deposits. Information compiled from the published literature on the radiocarbon age of marine shells gave information on a total of 734 radiocarbon dates from 276 localities from nearshore (< 10 m depth) and shelf (> 10 m depth) habitats. The median age of 128 nearshore shells is 2,465 years; that of 158 shells from the shelf is 8,870 years. The distribution of shell ages in both nearshore and shelf environments is strongly skewed: most dates are in the 0–3,000 range, and the number of shells in older age-classes falls off rapidly. The maximum age of a shell in an active sedimentary environment is a measure of time-averaging, because it estimates the amount of time represented in the deposit. The median duration of time-averaging in 63 nearshore deposits is 1,250 years; the median duration of time-averaging in 129 shelf deposits is 9,190 years. Radiocarbon-dated shells from fossil deposits confirm our estimate of time-averaging in nearshore environments: the median difference between maximum and minimum ages in 49 inactive beach ridges is 1,390 years; the median difference in other, predominantly nearshore deposits, is 830 years. Greater shell survival and longer durations of time-averaging in shelf settings may result from lower rates of sedimentation, lower rates of taphonomic destruction, greater rates of bioturbation, the history of post-glacial sea level, sample bias, or some combination of these factors. Our results may estimate the actual magnitude of time-averaging of shelly deposits forming under conditions of low net sediment accumulation. Such extensive time-averaging may confound attempts at detailed paleoecological and paleoenvironmental reconstruction. □ Time-averaging, benthic assemblages, taphonomy, radiocarbon, paleoecology.     

Taphonomie des plantes et interprétations paléoécologiques. Une synthèse

The taphonomy is a powerful and requisite tool for environmental reconstructions of ancient plant communities. Necrobiotic processes, which lead to the production of plant fragments, inform us on fossil plant physiology. Among the processes that can be drawn from necrobiotic studies is the retention of leaf organs on plants, the relative quantity of pollen grains produced by different fossil species or the significance of wildfire dynamics in ancient plant communities. Biostratinomy examination is a fundamental tool for elucidating fossil plant habitats. Numerous experimental data allow paleobotanists for evaluating the role of transport in the origin of fossil assemblages. Autochthonous plant assemblages, which are characterised by the preservation of fossil rooting structures, are relatively rare in the nature. In consequence, the search for palaeoecological information from parautochthonous to allochthonous assemblages has been a priority in taphonomy. As a result, taphonomic models have been elaborated in well-known sedimentological contexts, such as small lacustrine deltas, which allow for the distinction between riparian or perideltaic plant remains. Lithospheric processes modify plant debris after burial. The differences in the degrees of transformations (or alterations) during the diagenesis provide for information about the original morphology and biochemical composition of the plant tissues, which are also paleoecologically useful. Thus, amber diagenesis modifies resin biochemistry into new molecules that are still informative from the chemotaxonomical point of view.     

Variability within the 10-Year Pollen Rain of a Seasonal Neotropical Forest and Its Implications for Paleoenvironmental and Phenological Research

Tropical paleoecologists use a combination of mud-water interface and modern pollen rain samples (local samples of airborne pollen) to interpret compositional changes within fossil pollen records. Taxonomic similarities between the composition of modern assemblages and fossil samples are the basis of reconstructing paleoclimates and paleoenvironments. Surface sediment samples reflect a time-averaged accumulation of pollen spanning several years or more. Due to experimental constraints, modern pollen rain samples are generally collected over shorter timeframes (1–3 years) and are therefore less likely to capture the full range of natural variability in pollen rain composition and abundance. This potentially biases paleoenvironmental interpretations based on modern pollen rain transfer functions. To determine the degree to which short-term environmental change affects the composition of the aerial pollen flux of Neotropical forests, we sampled ten years of the seasonal pollen rain from Barro Colorado Island, Panama and compared it to climatic and environmental data over the same ten-year span. We establish that the pollen rain effectively captured the strong seasonality and stratification of pollen flow within the forest canopy and that individual taxa had variable sensitivity to seasonal and annual changes in environmental conditions, manifested as changes in pollen productivity. We conclude that modern pollen rain samples capture the reproductive response of moist tropical plants to short-term environmental change, but that consequently, pollen rain-based calibrations need to include longer sampling periods (≥7 years) to reflect the full range of natural variability in the pollen output of a forest and simulate the time-averaging present in sediment samples. Our results also demonstrate that over the long-term, pollen traps placed in the forest understory are representative samples of the pollen output of both canopy and understory vegetation. Aerial pollen traps, therefore, also represent an underutilized means of monitoring the pollen productivity and reproductive behavior of moist tropical forests.     

Iterative evolution of sympatric seacow (Dugongidae, Sirenia) assemblages during the past ~26 million years

Extant sirenians show allopatric distributions throughout most of their range. However, their fossil record shows evidence of multispecies communities throughout most of the past ～26 million years, in different oceanic basins. Morphological differences among co-occurring sirenian taxa suggest that resource partitioning played a role in structuring these communities. We examined body size and ecomorphological differences (e.g., rostral deflection and tusk morphology) among sirenian assemblages from the late Oligocene of Florida, early Miocene of India and early Pliocene of Mexico; each with three species of the family Dugongidae. Although overlapping in several ecomorphological traits, each assemblage showed at least one dominant trait in which coexisting species differed. Fossil sirenian occurrences occasionally are monotypic, but the assemblages analyzed herein show iterative evolution of multispecies communities, a phenomenon unparalleled in extant sirenian ecology. As primary consumers of seagrasses, these communities likely had a strong impact on past seagrass ecology and diversity, although the sparse fossil record of seagrasses limits direct comparisons. Nonetheless, our results provide robust support for previous suggestions that some sirenians in these extinct assemblages served as keystone species, controlling the dominance of climax seagrass species, permitting more taxonomically diverse seagrass beds (and sirenian communities) than many of those observed today.     

On using time-averaging restraints in molecular dynamics simulation

Introducing experimental values as restraints into molecular dynamics (MD) simulations to bias the values of particular molecular properties, such as nuclear Overhauser effect intensities or distances, 3J coupling constants, chemical shifts or crystallographic structure factors, towards experimental values is a widely used structure refinement method. To account for the averaging of experimentally derived quantities inherent in the experimental techniques, time-averaging restraining methods may be used. In the case of structure refinement using 3J coupling constants from NMR experiments, time-averaging methods previously proposed can suffer from large artificially induced structural fluctuations. A modified time-averaged restraining potential energy function is proposed which overcomes this problem. The different possible approaches are compared using stochastic dynamics simulations of antamanide, a cyclic peptide of ten residues.     

Fidelity of molluscan life and death assemblages on sublittoral hard substrata around granitic islands of the Seychelles

This study quantified the degree of coincidence between living and dead molluscan faunas in a shallow-water coral reef environment in the Indian Ocean. The results were compared with those from a similar life:death study in the northern Red Sea, and with those published for reef corals and soft substrata molluscs. The proportions of quantitatively important taxa are robust to sampling intensity, but fidelity indices and rank-order correlations are strongly influenced by quantitatively unimportant taxa. Distinct differences between life and death assemblages were recognized, which are due to distinct biases in the death assemblage. Bivalves that lived in close contact with living corals are preferentially overgrown after death and should provide considerable temporal and ecological information in a potential fossil record, as they will be preserved within a rapidly growing reef framework. Some gastropod taxa are preferentially transported into surrounding soft substrata postmortem. Here they will be affected by time-averaging and taphonomic disintegration typically occurring in sediments resulting in the associated loss of much temporal information. Most gastropod shells, however, are inhabited by hermit crabs postmortem, which may strongly alter the fossil gastropod community structure. The results are similar to a comparable study in the northern Red Sea, with a major exception being the strong dominance of hermit crab-inhabited gastropod shells in the death assemblage of the Seychelles. Comparison of life:death assemblages between hard and soft substrata, in keeping with the northern Red Sea study, showed the strong dominance of dead shells in the soft substrata with the converse on the hard substrata. This results from different accumulation conditions for dead shells in soft substrata. Fidelity indices are well suited to demonstrate that sedimentary death assemblages are typically remarkably robust reflections of local community composition but they do not record the strong biases in death assemblages of coral reef associated hard substrata molluscs and are therefore unsuitable for comparisons of life and death assemblages in reef environments.