L’Explosion cambrienne ou l’émergence des écosystèmes modernes

Marine ecosystems with complex trophic structure and dominated by animals started to build up in the Early Cambrian. Fossil evidence from exceptional fossil localities such as the Chengjiang Lagerstätte from South China indicate a high level of biological interactivity (e.g. prey–predator relationships) and the colonization of a wide range of pelagic and benthic niches by predators, scavengers, and detritus and suspension feeders. Swimmers are numerous, but there is no evidence for the extensive occupation of the water column by the Early Cambrian. On the contrary, animal life may have concentrated in hyperbenthic environments, close to the sea bottom. This would have been the initial step towards the colonization of the whole pelagic realm and the building-up of pelagic food chains. A chain of biotic innovations and events seems to have catalyzed both the animal diversification and the build-up of a completely new type of ecosytem, with: (1) the achievement of complex nervous systems, visual organs and motor functions; (2) the introduction of new selective pressure (e.g. predation and feed-back effects); and (3) the colonization of new niches. The role of environmental factors (e.g. oxygen, water chemistry, climate) may have been important in the early stages of metazoan evolution, but was probably negligible in the ecological turnover itself that takes place in the Early Cambrian. Close resemblances between the trophic structure of present-day ecosystems and that of Cambrian ones are confirmed by fossil data and recent mathematical models. This unprecedented increase of interdependence between animal species and trophic levels probably increased the general stability of marine ecosystems, but made them for the first time in their history, highly vulnerable to environmental perturbations. This will largely influence the post-Cambrian evolution of the marine world.     

The Early Cambrian colonization of pelagic niches exemplified by Isoxys (Arthropoda)

The anatomy of the bivalved arthropod Isoxys (Early and Middle Cambrian) is reconstructed, based on new evidence from soft parts and exoskeletal design and on a critical review of previous work. Isoxys had a long segmented body flanked with a pair of short antennules, followed by a series of 14 biramous appendages provided with long paddle-like exopods concealed under a widely open bivalved carapace folded dorsally and bearing long cardinal spines. The close resemblance between Isoxys and Recent pelagic crustaceans (halocyprid ostracods, larval stages of malacostracans) indicates that Isoxys was probably an active epipelagic swimmer (evidence from soft parts, carapace design and distributional pattern). Some species (e.g. I. auritus and I. paradoxus from the Maotianshan Shale biota; Early Cambrian) may have lived in the vicinity of the bottom either permanently or temporarily, whereas others may have had ecological preferences for more open-marine settings. The spinosity of Isoxys had a possible role in predatorial deterrence rather than in buoyancy control or in retarding sinking within the water column. The presence of Isoxys in the Maotianshan Shale of S. China indicates that arthropods had already colonized midwater niches by the Early Cambrian. The midwater communities of the Maotianshan Shale comprised numerous other invertebrates, such as abundant medusiform eldonids, vetulicolids, chordates and possibly early vertebrates. This contradicts the opinion that pelagic communities remained poorly developed until late Cambrian/Ordovician times and that the occupation of the midwater niches largely post-dates the initial diversification of the benthic faunas.     

Epibionts on the lingulate brachiopod Diandongia from the Early Cambrian Chengjiang Lagerst?tte,South China

The classic Chengjiang Lagerstätte (Lower Cambrian, Atdabanian stage: Yu''anshan Formation) Yunnan, southwestern China, has yielded, besides the exceptional and often controversial soft-bodied fossils, a fauna of primitive/early lingulid brachiopods. Diandongia pista (Rong 1974) is one of the commonest and most strongly mineralized of the phosphatic brachiopods from the Lagerstätte. The shells of this species have been found to commonly serve as a basibiont host. Epibionts comprise the coeval brachiopod Longtancunella chengjiangensis and the cone-shaped cnidarian-related Archotuba conoidalis, as well as rounded smaller-sized epizoans (lesser than 2 mm). A principle morphological analysis demonstrates that the ovoid and rounded organisms that often occur along the commissure of D. pista resemble small juvenile or immature brachiopods. Epibiont-bearing shells of D. pista with soft-tissue preservation demonstrate that the host brachiopods were overgrown while alive, and provide an argument for D. pista having a semi-infaunal life style with only the slim pedicle embedded in sediment. The epibiotic association sheds direct light on the ecology of Cambrian brachiopods in soft-substrate marine environments. The Chengjiang fossils demonstrate that the Early Cambrian brachiopods, as compared with recent lingulids, occupied different and a wider spectrum of ecological niches and tiers of space.     

Community palaeoecology of the early Cambrian Maotianshan Shale biota: Ecological dominance of priapulid worms

The early Cambrian soft-bodied Maotianshan Shale (Chengjiang) biota of Yunnan Province, China provides a critical glimpse of animal life during the heart of the Cambrian radiation. The Shankou biota is the focus of this current work, and 9963 specimens from it have been examined and tallied. This collection of fossils represents a time-averaged assemblage of unknown duration from a 10-meter thick sequence. The predominantly benthic community recovered from this section is presumably autochthonous. This community was buried through several hundred millimeter to centimeter thick burial events. Only specimens interpreted as buried alive, based on either soft-part preservation or fully articulated skeletons, were counted. Algae were too fragmented to be counted. To avoid including synonymous species, diversity was evaluated at the genus level. A total of 57 genera from 9 phyla, encompassing 14 ecological groups, were found in this assemblage. The results reveal that the three most abundant genera comprise 43.2% of all specimens: the tubiform priapulid worm Paraselkirkia (16.0%), the diminutive priapulid worm Sicyophorus (14.3%), and the brachiopod Heliomedusa (12.9%). No other genera total more than 9% of specimens. At the phylum level, there is an interesting dichotomy between taxonomic diversity and ecological dominance. The arthropods are the most diverse phylum (37% of genera) and rank second in relative abundance of specimens (26.3%). The priapulids, however, with only 17.5% of genera is the most abundant group in the assemblage (43.2%). All other phyla, excluding brachiopods (19.6%), represent only 10.9% of the assemblage. Because of their sheer numerical abundance, this study indicates that priapulid worms may have exerted more influence on energy flow and community structure than other phyla in this particular trophic web. This result contrasts strongly with both traditional views of Maotianshan Shale biota palaeoecology, which often claimed arthropod dominance based solely on taxonomic diversity, and the palaeoecology of modern priapulids, which are relegated to marginal marine settings. This result also demonstrates the importance of collecting quantitative relative abundance data when performing palaeoecological investigations.     

Gut Contents as Direct Indicators for Trophic Relationships in the Cambrian Marine Ecosystem

Present-day ecosystems host a huge variety of organisms that interact and transfer mass and energy via a cascade of trophic levels. When and how this complex machinery was established remains largely unknown. Although exceptionally preserved biotas clearly show that Early Cambrian animals had already acquired functionalities that enabled them to exploit a wide range of food resources, there is scant direct evidence concerning their diet and exact trophic relationships. Here I describe the gut contents of Ottoia prolifica, an abundant priapulid worm from the middle Cambrian (Stage 5) Burgess Shale biota. I identify the undigested exoskeletal remains of a wide range of small invertebrates that lived at or near the water sediment interface such as hyolithids, brachiopods, different types of arthropods, polychaetes and wiwaxiids. This set of direct fossil evidence allows the first detailed reconstruction of the diet of a 505-million-year-old animal. Ottoia was a dietary generalist and had no strict feeding regime. It fed on both living individuals and decaying organic matter present in its habitat. The feeding behavior of Ottoia was remarkably simple, reduced to the transit of food through an eversible pharynx and a tubular gut with limited physical breakdown and no storage. The recognition of generalist feeding strategies, exemplified by Ottoia, reveals key-aspects of modern-style trophic complexity in the immediate aftermath of the Cambrian explosion. It also shows that the middle Cambrian ecosystem was already too complex to be understood in terms of simple linear dynamics and unique pathways.     

节肢动物吐卓虫咬痕化石在云南昆明附近寒武纪早期关山生物群的发现及意义*

本文报道了一块来自云南省昆明市附近寒武纪第二世第四期关山生物群具咬痕的双瓣壳节肢动物吐卓虫 (Tuzoia)化石, 系吐卓虫咬痕化石在全球范围的首次发现。该发现证实作为主动捕食者的吐卓虫很可能被大型捕食者(如奇虾类或莱德利基虫类三叶虫等)捕食, 说明寒武纪早期的海洋生物群落已经具有高度复杂化的食物链, 为进一步了解吐卓虫的生态位及深入探讨寒武纪大爆发时期海洋生态系统食物网结构提供了新的信息。     

Protracted oxygenation across the Cambrian–Ordovician transition: A key initiator of the Great Ordovician Biodiversification Event?

Fluctuations in marine oxygen concentrations have been invoked as a primary driver for changes in biodiversity throughout Earth history. Expansions in reducing marine conditions are commonly invoked as key causal mechanisms for mass extinctions, while increases in marine oxygenation are becoming an increasingly common causal mechanism invoked for biodiversification events. Here we utilize a multiproxy approach to constrain local and global marine paleoredox conditions throughout the late Cambrian–Early Ordovician from two drill core successions in Baltoscandia. Local paleoredox proxies such as manganese concentrations and iron speciation reveal that both sites in the Baltic paleobasin had persistently anoxic and predominantly euxinic (anoxic and sulfidic) bottom water conditions throughout the study interval. Corresponding trace metal datasets indicate nuanced contraction and expansion of global anoxic and euxinic conditions along continental margins during the late Cambrian–Early Ordovician. Lastly, thallium isotope data from these locally reducing sections suggest a global expansion of oxygenated shelf and deeper marine environments from the late Cambrian into the Early Ordovician. This evidence for increasingly oxic marine environments coincides with increases in burrowing depth and tiering in marine animals, as well as diversification of body fossils throughout this ~8-million-year interval. The collective geochemical datasets provide some of the first direct paleoredox evidence for an increase in marine oxygen concentrations as a key mechanism for the Ordovician radiation of marine life.     

节肢动物吐卓虫咬痕化石在云南昆明附近寒武纪早期关山生物群的发现及意义*

本文报道了一块来自云南省昆明市附近寒武纪第二世第四期关山生物群具咬痕的双瓣壳节肢动物吐卓虫 (Tuzoia)化石, 系吐卓虫咬痕化石在全球范围的首次发现。该发现证实作为主动捕食者的吐卓虫很可能被大型捕食者(如奇虾类或莱德利基虫类三叶虫等)捕食, 说明寒武纪早期的海洋生物群落已经具有高度复杂化的食物链, 为进一步了解吐卓虫的生态位及深入探讨寒武纪大爆发时期海洋生态系统食物网结构提供了新的信息。     

寒武纪早期澄江生物群中受创伤的中间型始莱德利基三叶虫(Eoredlichia intermedia)

三叶虫是寒武纪海洋中的常见动物,其受伤愈合现象的化石记录可以反映生态系统内物种之间竞争关系。中国华南寒武纪早期澄江化石生物群因保存软躯体构造化石而成为研究寒武纪大爆发时期古海洋生态群落的绝佳窗口,然而,澄江生物群中三叶虫受伤愈合现象的化石至今未见报道。中间型始莱德利基虫(Eoredlichia intermedia Lu,1940)是澄江生物群的常见物种,也是我国寒武系第二个三叶虫化石带的标准带化石。在2 000余枚中间型始莱德利基虫标本中,发现两枚成体标本的头甲侧边缘、颊刺和胸甲肋节保存了不同程度的残缺。据此推测,Eoredlichia intermedia在生命活动过程中曾受到碎壳型捕食动物的攻击或其他伤害而形成伤口。伤口部位边缘加厚且光滑,显示了明显的愈合痕迹。这些特征表明始莱德利基虫在受到非致死的伤害后,具备自身修复损伤的能力。这是迄今所知最为古老的,也是在澄江生物群中首次发现的三叶虫受伤并且愈合的标本,反映了寒武纪第二世第三期底栖生态系统中碎壳型捕食者和三叶虫猎物之间的军备竞赛生态关系。     

埃迪卡拉纪至寒武纪苗岭世刺细胞动物化石研究现状与展望

刺细胞动物是一类具有刺细胞的水生无脊椎动物,分布在世界各地的海洋和淡水中.作为后生动物最早分化出的一支,刺细胞动物对研究后生动物的起源和早期演化具有极其重要的意义,也为研究后生动物系统发育、地层对比和古地理恢复等方面提供了重要的科研线索.本文简要介绍了刺细胞动物早期(埃迪卡拉纪至寒武纪苗岭世)的化石记录和研究现状,将刺...     

Cyanobacterial macrophytes in an Early Silurian (Llandovery) continental biota: Passage Creek, lower Massanutten Sandstone, Virginia, USA

A compression macrofossil with structure consisting of mineral-replaced filaments embedded in an amorphous carbonaceous matrix is described as a macrophytic cyanobacterial colony from continental assemblages of the Early Silurian (Llandovery) Passage Creek biota, in the lower Massanutten Sandstone (Virginia, USA). Filaments are predominantly multiseriate and consist of spheroidal crystalline aggregates representing early pyrite (subsequently replaced by iron hydroxides) precipitated preferentially inside cells. Interpretation of the fossils as cyanobacteria is based on close similarities to modern organisms in terms of overall morphology and production of copious extracellular investments, filament and cell sizes, and continental epigeal (freshwater or terrestrial) habitat. This interpretation incorporates data on cyanobacterial taphonomy and mechanisms of diagenetic mineral precipitation. These fossils are part of the oldest macrofossil assemblages documenting well-developed and diverse communities on continents. They provide the earliest direct evidence for cyanobacteria in strictly continental habitats, corroborating the commonly held but poorly documented view that cyanobacteria were among the initial colonizers of continents.     

Trace fossils from lower Cambrian Hongjingshao Formation,Yunnan, China: Taxonomy,palaeoecology, palaeoenvironment

The Hongjingshao Formation (Cambrian Series 2, Stage 3) yields abundant, complex trace fossils that have not been systematically investigated up till now. Our detailed ichnological study of the traces from this formation in Malong area, Yunnan, China, recognizes four groups: (1) simple horizontal or sub-horizontal burrows; (2) complex branched burrow systems; (3) arthropod trails and; (4) simple vertical burrows belonging to the Cruziana ichnofacies. A trophic web is reconstructed for the community in Hongjingshao Formation based on both trace and body fossils. The ichnological and sedimentological features of the formation indicate an intertidal setting. The trace fossil assemblage demonstrates that Early Cambrian organisms were able to colonize very shallow marine environments, which further supports the landward expansion of the Early Cambrian ecosystems.     

贵州麻江地区寒武纪杷榔动物群的群落组成和生态空间分析*

寒武纪特异埋藏化石库保存了海洋生物群落最原始的生态结构, 为群落生态研究提供了宝贵材料。贵州寒武纪斜坡相区的杷榔动物群(第4期)具有较高的生物多样性, 但定量生态学研究相对薄弱。本研究对贵州麻江地区杷榔动物群的新产地平定剖面进行化石采集和分类统计, 并运用多元数理统计方法开展定量化分析。鉴定出腕足动物、海绵动物、刺胞动物、节肢动物、软舌螺动物、古虫动物、开腔骨动物和曳鳃动物8个化石门类, 计31属33种, 划分出12种生态类型。其中, 腕足动物具有最高的个体丰度(占比61.5%), 节肢动物物种分异度最高(占比60%), 群落以底栖固着生态类型为主, 游泳捕食为辅。对比分析结果显示, 寒武纪海洋群落具有空间异质性, 且节肢动物在影响群落组成和生态空间利用上起决定性作用。随水体深度增加, 海洋生态群落中的表栖类群出现明显扩张, 也体现了环境对群落面貌的塑造作用。     

Interpopulation trophic niches and ontogenetic shifts of a mangrove fish predator

Fish trophic niches reflect important ecological interactions and provide insight into the structure of mangrove food webs. Few studies have been conducted in mangrove fish predators to investigate interpopulation trophic niches and ontogenetic shifts. Using stable isotope analysis and two complementary approaches, the authors investigated trophic niche patterns within and between two ontogenetic groups (juveniles and sub-adults) of a generalist predator (Acentrogobius viridipunctatus) in four mangroves with heterogeneous environmental conditions (e.g., tidal regimes, salinity fluctuations and mangrove tree community). The authors hypothesized that the trophic niche between populations would vary regionally and trophic position would increase consistently from juvenile to sub-adult stages. The results revealed that both δ¹³C and δ¹⁵N values varied greatly across populations and between ontogenetic groups, and complex spatio-ontogenetic variations were expressed by Layman's metrics. They also found some niche separation in space, which is most likely related to resource availability in spatially diverse ecosystems. In addition, trophic niche position increased consistently from juveniles to sub-adults, indicating ontogenetic feeding shifts. The isotopic plasticity index and Fulton's condition index also showed significant spatial-ontogenetic variation, which is consistent with optimal foraging theory. The findings highlight that trophic plasticity has a high adaptive value for mangrove fish predators in dynamic ecosystems.     

Non-lichenized Antarctic fungi: transient visitors or members of a cryptic ecosystem?

To date over 1 000 non-lichenized fungal species have been recorded by collection or isolation from Antarctica, and additional taxa are now being identified by molecular studies. The number and variety of species recorded so far suggest that the fungi may be the most diverse biota in the Antarctic, and the additional taxa identified by molecular surveys suggest that the true diversity may be far greater than is currently estimated. Fungi occupy many different ecological niches in the Antarctic, and their significance in these niches is only poorly understood. The majority of species described from the region have been identified as members of broadly cosmopolitan groups, but there is some evidence for both endemic strains and populations. This review brings together the current broad systematic and ecological findings for the non-lichenized Antarctic fungi.     

Characterizing niche differentiation among marine consumers with amino acid δ13C fingerprinting

Marine food webs are highly compartmentalized, and characterizing the trophic niches among consumers is important for predicting how impact from human activities affects the structuring and functioning of marine food webs. Biomarkers such as bulk stable isotopes have proven to be powerful tools to elucidate trophic niches, but they may lack in resolution, particularly when spatiotemporal variability in a system is high. To close this gap, we investigated whether carbon isotope (δ¹³C) patterns of essential amino acids (EAAs), also termed δ¹³C_AA fingerprints, can characterize niche differentiation in a highly dynamic marine system. Specifically, we tested the ability of δ¹³C_AA fingerprints to differentiate trophic niches among six functional groups and ten individual species in the Baltic Sea. We also tested whether fingerprints of the common zooplanktivorous fishes, herring and sprat, differ among four Baltic Sea regions with different biochemical conditions and phytoplankton assemblages. Additionally, we investigated how these results compared to bulk C and N isotope data for the same sample set. We found significantly different δ¹³C_AA fingerprints among all six functional groups. Species differentiation was in comparison less distinct, due to partial convergence of the species' fingerprints within functional groups. Herring and sprat displayed region‐specific δ¹³C_AA fingerprints indicating that this approach could be used as a migratory marker. Niche metrics analyses showed that bulk isotope data had a lower power to differentiate between trophic niches than δ¹³C_AA fingerprinting. We conclude that δ¹³C_AA fingerprinting has a strong potential to advance our understanding of ecological niches, and trophic linkages from producers to higher trophic levels in dynamic marine systems. Given how management practices of marine resources and habitats are reshaping the structure and function of marine food webs, implementing new and powerful tracer methods are urgently needed to improve the knowledge base for policy makers.     

Quantitative paleoecology of Cambrian (Series 2–Miaolingian) communities from central Sonora,Mexico

This work presents a paleoecological approach to study benthic communities preserved in Cambrian strata exposed at the Sahuaral locality in central Sonora, northwestern Mexico. Based on species richness and abundance, this work analyzed 627 individuals corresponding to 33 species from five distinctive Cambrian formations: the lowermost Proveedora Formation characterized by bioturbation horizons that record the occurrence of a benthic-infaunal biota from coastal shallow seas, followed by the Buelna Formation which contains reef deposits dominated by autotrophic epibenthic communities. The Cerro Prieto Formation was deposited in a high-energy shallow marine environment, dominated by primary consumers-epifaunal and suspensions feeders. The uppermost unit, the El Gavilán Formation, is an intercalation of shale and fossiliferous limestone characterized by the fossil remains of suspension feeders, detritus feeders and nektobenthic organisms of low energy deep water environments. The paleobiota consist of ichnofossils, microbial oncolites, chancellorids, brachiopods, hyolithids, and trilobites in a sequence of increasing diversity and complexity. Paleoecological indices (Simpson Dominance, Shannon-Weaver Index, Evenness Index, and Total Richness) were determined and compared in the study area. In addition, the effective number of species and the true diversities were obtained, which are similar between each station distributed in each formation. However, it is possible that the high degree of dominance is a factor to indicate an inequality between the true diversity and the species richness identified. These results provide new analytical and paleoecological criteria for better understanding depositional environments and fossil associations where the spatial distribution of benthic organisms, richness and abundance provide a valued window to Cambrian biotas. The structure of the marine community of the El Sahuaral area has a great affinity with other Cambrian communities, such as those from the Stephen Formation (Burgess Shale, British Columbia) and the Chengjiang biota of the Yangtze Shelf deposits in South China.     

Ecological innovations in the Cambrian and the origins of the crown group phyla

Simulation studies of the early origins of the modern phyla in the fossil record, and the rapid diversification that led to them, show that these are inevitable outcomes of rapid and long-lasting radiations. Recent advances in Cambrian stratigraphy have revealed a more precise picture of the early bilaterian radiation taking place during the earliest Terreneuvian Series, although several ambiguities remain. The early period is dominated by various tubes and a moderately diverse trace fossil record, with the classical ‘Tommotian’ small shelly biota beginning to appear some millions of years after the base of the Cambrian at ca 541 Ma. The body fossil record of the earliest period contains a few representatives of known groups, but most of the record is of uncertain affinity. Early trace fossils can be assigned to ecdysozoans, but deuterostome and even spiralian trace and body fossils are less clearly represented. One way of explaining the relative lack of clear spiralian fossils until about 536 Ma is to assign the various lowest Cambrian tubes to various stem-group lophotrochozoans, with the implication that the groundplan of the lophotrochozoans included a U-shaped gut and a sessile habit. The implication of this view would be that the vagrant lifestyle of annelids, nemerteans and molluscs would be independently derived from such a sessile ancestor, with potentially important implications for the homology of their sensory and nervous systems.     

Stable isotope analyses reveal dense trophic species packing and clear niche differentiation in a malagasy primate community

Understanding the mechanisms maintaining local species richness is a major topic in tropical ecology. In ecological communities of Madagascar, primates represent a major part of mammalian diversity and, thus, are a suitable taxon to study these mechanisms. Previous research suggested that ecological niche differentiation facilitates the coexistence of lemurs. However, detailed data on all species making up diverse local primate assemblages is rarely available, hampering community‐wide tests of niche differentiation among Malagasy mammals. Here, we took an indirect approach and used stable isotopes as long‐term indicators of individuals' diets to answer the question of whether trophic patterns and food‐related mechanisms stabilize coexistence in a species‐rich lemur community. We analyzed stable carbon and nitrogen isotopes in hair collected from eight syntopic lemurs in Kirindy Forest. We found that lemur species were well separated into trophic niches and ranged over two trophic levels. Furthermore, species were densely packed in isotopic space suggesting that past competitive interactions between species are a major structuring force of this dry forest lemur community. Results of other comparative studies on primates and our findings underline that—in contrast to communities worldwide—the structure and composition of lemur communities follow predictions of ecological niche theory. Patterns of competitive interactions might be more clearly revealed in Malagasy primate communities than elsewhere because lemurs represent a large fraction of ecologically interacting species in these communities. The pronounced trophic niche differentiation among lemurs is most likely due to intense competition in the past as is characteristic for adaptive radiations. Am J Phys Anthropol 153:249–259, 2014. © 2013 Wiley Periodicals, Inc.     

The Luoping biota: exceptional preservation, and new evidence on the Triassic recovery from end-Permian mass extinction

The timing and nature of biotic recovery from the devastating end-Permian mass extinction (252 Ma) are much debated. New studies in South China suggest that complex marine ecosystems did not become re-established until the middle–late Anisian (Middle Triassic), much later than had been proposed by some. The recently discovered exceptionally preserved Luoping biota from the Anisian Stage of the Middle Triassic, Yunnan Province and southwest China shows this final stage of community assembly on the continental shelf. The fossil assemblage is a mixture of marine animals, including abundant lightly sclerotized arthropods, associated with fishes, marine reptiles, bivalves, gastropods, belemnoids, ammonoids, echinoderms, brachiopods, conodonts and foraminifers, as well as plants and rare arthropods from nearby land. In some ways, the Luoping biota rebuilt the framework of the pre-extinction latest Permian marine ecosystem, but it differed too in profound ways. New trophic levels were introduced, most notably among top predators in the form of the diverse marine reptiles that had no evident analogues in the Late Permian. The Luoping biota is one of the most diverse Triassic marine fossil Lagerstätten in the world, providing a new and early window on recovery and radiation of Triassic marine ecosystems some 10 Myr after the end-Permian mass extinction.