Carbonaceous Compression Fossils Research Articles

Macroscopic carbonaceous compression fossils from the Tonian Liulaobei Formation in the Huainan region of North China

Macroscopic carbonaceous compression fossils are key to understand the early evolution of eukaryotes, explore the development of multicellularity, and strengthen the basis of biostratigraphic correlation during the Tonian Period (ca. 1000–720 Ma). Despite the potential geobiological importance of early macrofossils, the fossil record of Tonian macro-organisms is rarely documented. Here we describe an assemblage of diversified carbonaceous macrofossils from the Tonian Liulaobei Formation in the Huainan region, North China. In this study, we examine a large population of these macrofossils, clarify their taxonomy, and tentatively interpret their phylogenetic affinities. This assemblage consists of thirteen distinct morphotaxa, including eight genera and three new species, showing remarkable taxonomic diversity and domination of coenocytic macroalgae. The new data improve our knowledge about early Neoproterozoic biodiversity, and indicate a Tonian increase in morphological and taxonomic diversity of macrofossils compared to the Mesoproterozoic Era. This study also suggests that macroalgae may have begun to diversify and played a crucial ecological role in Tonian marine ecosystems.

Phosphatized microfossils from the Miaohe Member of South China and their implications for the terminal Ediacaran biodiversity decline

The upper Ediacaran Miaohe Member (∼550 Ma) in South China is well known for exceptionally preserved macroscopic carbonaceous compression fossils (i.e., the Miaohe biota) that are pivotal in understanding the marine ecology and environment during the terminal Ediacaran Period. However, micro-organisms, which are also important biotic components of marine ecosystems, are poorly documented and largely limited to simple leiospheres and filaments in the Miaohe Member and equivalent strata in South China and around the world. Here we report a moderately diverse assemblage of cellularly preserved microfossils from phosphorites and siliceous phosphorite nodules and bands of the Miaohe Member at the Maxi section, Hubei Province, South China. Ten named species of eight genera, four open nomenclatures, and two unnamed taxa have been identified, including several multicellular algal fossils and various coccoidal and filamentous cyanobacteria. Abundant spheroidal fossils are morphologically analogous to Megasphaera, which has previously been interpreted as a putative metazoan embryo. Together with previously described microfossils from equivalent strata of Miaohe Member, these microfossils expand our understanding of the evolution of marine ecosystems prior to the Cambrian explosion. The late Ediacaran Maxi assemblage and the early Ediacaran Weng’an biota, both preserved through phosphatization, share several taxa. Notably, Doushantuo-Pertatataka-type acanthomorphs (DPAs) are absent from the Maxi assemblage but abundant in the Weng’an biota, consistent with a late Ediacaran decline of DPAs, which may have ushered in the terminal Ediacaran decline of soft-bodied macro-organisms of the Ediacara biota during the Kotlin crisis or in the last 10 million years of the Ediacaran Period.

Non-calcified warm-water marine macroalgae from the Ordovician strata of Spiti, Tethys Himalaya, India

The calcareous siltstones within the Ordovician section of the Takche Formation near Takche, Spiti region of Tethyan Himalaya, India, contain numerous specimens of non-calcified marine macroalgae in association with brachiopods, gastropods, tentaculitoids, and few trace fossils. The algal remains, representing five genera, are preserved as black or dark brown carbonaceous compression fossils, interpreted as warm-water marine macroalgae, namely, Inocladus sp., Callisphenus? sp., Algites sp. (siphonous algae), Fisherites sp., and Mastopora sp. (non-siphonous). Callisphenus is characterised by a radially symmetrical short pyriform thallus, with a central axis surrounded by short laterals whereas Inocladus sp. is characterised by an unsegmented simple thallus with internal parallel medullary siphons and cortical tubes. These algal remains, probably transported from a more near-shore living niche, co-occur with Cyclocrinitids, Tentaculitoids tube worms, trace fossils and brachiopods, suggesting that deposition of the studied units took place in low energy hydrodynamic conditions influenced by intermittent storm events. The study represents the first diversified macroalgal records from the Ordovician strata in the Tethyan realm of Indian subcontinent.

New Material of Carbonaceous Compressions from the ∼1.5 Ga Singhora Group, Chhattisgarh Supergroup, India, and their Interpretation as Benthic Algae

The origin, antiquity, and affinity of benthic seaweeds (multicellular algae) in the geological past are shrouded in mystery due to their preservation bias. In this study, we present a new material of well-preserved carbonaceous compression fossils in shale horizons of the Mesoproterozoic (ca. ∼1,500–1,300 Ma) Singhora Group of the Chhattisgarh Supergroup. Eleven distinct taxa, including one new taxon, Palaeoscytosiphon shuklaii, n. gen. et. sp., and one new species, Jiuqunaoella sergeevii, n. sp., are established. Four unidentified morphologies are also reported. Morphologically, the carbonaceous fossils are fan-shaped, palmate, elongated, leaf-like algal thalli with/without holdfast at the base, isolated or dichotomously branched long filaments, along with multicellular reproductive structures. The results of laser Raman spectroscopy and energy dispersive X-ray spectroscopy (EDX) are also presented in support of their biogenicity. Collectively, the preservation mode of the Singhora carbonaceous fossils suggests multicellular algal affinity and adds to a Burgess Shale-type (BST) taphonomic window in the Pre-Ediacaran biosphere.

First record of organic-walled microfossils from the Tonian Shiwangzhuang Formation of the Tumen Group in western Shandong, North China

Organic-walled microfossils offer important information on the biospheric evolution in pre-Cryogenian and provide biostratigraphic implications for many Proterozoic fossiliferous sequences that are poorly age constrained for the lack of reliable radiometric date. Recently, macroscopic carbonaceous compression fossils have been reported for the first time from the Tonian Shiwangzhuang Formation of the Tumen Group in western Shandong, North China. However, organic-walled microfossils have never been discovered from this formation up till now. To improve our knowledge about Proterozoic biodiversity in North China, we conducted a micropaleontological survey on the argillaceous limestone samples of the Shiwangzhuang Formation, which also contain macroscopic carbonaceous compression fossils, from the Baishicun section in Anqiu, western Shandong, North China. Our investigation shows that the Shiwangzhuang microfossil assemblage is dominated by smooth-walled sphaeromorphic acritarchs and cyanobacterium-like filaments and relatively low abundance of other acritarchs, including 16 taxa, such as Polysphaeroides filliformis, Ostiana microcystis, Simia annulare, ?Jacutianema sp., Arctacellularia tetragonala, Pellicularia tenera, Polythrichoides lineatus, and Navifusa actinomorpha. The Shiwangzhuang organic-walled microfossil assemblage, although consisting of long-ranging and not age diagnostic taxa, is consistent with a Tonian age suggested by macroscopic carbonaceous compression fossils, including the Chuaria-Tawuia and Sinosabellidites-Protoarenicola-Pararenicola assemblages, revealed from the same fossiliferous horizon of the Shiwangzhuang Formation and by organic-walled microfossil assemblage, including the late Mesoproterozoic to Tonian index fossil Trachyhystrichosphaera aimika, from the underlying Tongjiazhuang Formation. However, it is also worth noting that a Cryogenian or Ediacaran age cannot be completely excluded based just on the Shiwangzhuang microfossils because of their limited biostratigraphic utility.

An assemblage of macroscopic and diversified carbonaceous compression fossils from the Tonian Shiwangzhuang Formation in western Shandong, North China

Macroalgae play a crucial ecological and biogeochemical role in modern marine ecosystems. Although eukaryotic algae have a deep history within the Precambrian, most pre-Cryogenian alga fossils are microscopic and macroalga fossils are poorly documented during this period. Here we report a well-preserved assemblage of macroscopic and diversified carbonaceous compression fossils from the Tonian Shiwangzhuang Formation in western Shandong, North China. This assemblage consists of seventeen distinct morphotaxa, including eleven macroalgae, showing remarkable taxonomic diversity that is more diverse than many other Tonian macrofossil assemblages all over the world. Three taxa from the Shiwangzhuang assemblage show evidence of simple multicellularity with two of them consisting of giant cells and comparable to giant-celled green algae, and seven taxa with diagnostic transverse annulations are likely coenocytic algae. The Shiwangzhuang assemblage indicates that some eukaryotic clades had achieved macroscopic growth through simple multicellularity or coenocytism, paving the way, either ecologically or phylogenetically, for the eventual appearance of complex multicellularity. In addition, there are some biostratigraphically significant genera, including the Chuaria-Tawuia and Sinosabellidites-Protoarenicola-Pararenicola assemblages, indicating a Tonian age for the Shiwangzhuang Formation. This study also suggests that the Tonian diversity of multicellular or coenocytic macroalgae is probably underestimated, pending for a systematic and thorough investigation of Tonian macroscopic carbonaceous compressions. Macroalgae may have an initial ecological expansion, at least locally if not globally, in the Tonian Period.

Comment on: Tang et al. [2019]: A problematic animal fossil from the early Cambrian Hetang Formation, South China

AbstractTang et al. (2019) described new specimens of carbonaceous compression fossils from the early Cambrian Hetang Formation in South China, for which they established the new taxon Cambrowania ovata Tang and Xiao in Tang et al., 2019. Tang et al. (2019) interpreted these fossils as the remains of metazoans, representing either the carapaces of bivalve arthropods, or early life-cycle stages of sponges. We contest the animal affinity of these fossils; instead, we propose that the specimens described as Cambrowania ovata are actually large Leiosphaeridia—in other words, collapsed hollow organic spheroidal acritarchs. The features described by Tang et al. (2019) all fall into the morphology of carbonaceous compressions of Leiosphaeridia with pyritized/baritized folds and compaction wrinkles. Such Leiosphaeridia are a common component of Cambrian (and older) siliciclastic deposits, and frequently exhibit such a pattern of pyritization, baritization, and encrustation with other diagenetic minerals.

A systematic description of new macrofossil material from the upper Ediacaran Miaohe Member in South China

The upper Ediacaran Miaohe Member (∼551 Ma) in South China contains exceptionally preserved macroscopic carbonaceous compression fossils that provide a crucial view of the marine ecosystem shortly before the Cambrian explosion. However, these fossils were previously known only from one locality near the Miaohe Village on the northern bank of the Yangtze River. It was puzzling that equivalent strata a short distance away contained no fossils, raising questions about the environmental, ecological and taphonomic restriction of these fossils. Here, we describe macroscopic fossils from the Miaohe Member at three new localities to the north and north-west of Miaohe Village. These fossils are taxonomically similar to those from Miaohe Village, and include mostly benthic multicellular algae, a few putative metazoans and several morphologically simple problematic taxa. Twenty-seven species are described, including one new genus (Maxiphyton gen. nov.) and six new species (Baculiphyca brevistipitata sp. nov., Doushantuophyton? laticladus sp. nov., Enteromorphites magnus sp. nov., Konglingiphyton? laterale sp. nov., Maxiphyton stipitatum sp. nov. and Sinocylindra linearis sp. nov.). Insofar as the new localities represent shallower palaeoenvironments than Miaohe Village in the late Ediacaran, the new fossils expand the palaeoenvironmental distribution of macrofossils in the Miaohe Member. Most of the new fossils are preserved as carbonaceous compressions through kerogenization, but pyritization and aluminosilicification also played an important role. This confirms the taphonomic model that there is a continuous preservational gradient among kerogenization, pyritization and aluminosilicification. This taphonomic window provides a glimpse of the late Ediacaran marine ecosystem and complements the cast-mould preservation that accounts for the much-celebrated Ediacara-type soft-bodied macrofossils.

Stable carbon isotopes of sedimentary kerogens and carbonaceous macrofossils from the Ediacaran Miaohe Member in South China: Implications for stratigraphic correlation and sources of sedimentary organic carbon

Ediacaran stratigraphic correlation depends on the integration of paleontological and δ13Ccarb data. However, these data often come from different lithologies (e.g., shales vs. carbonates), making it difficult to directly link biostratigraphic and chemostratigraphic markers. Ediacaran successions in South China consist of mixed carbonates and shales, offering an opportunity for integrated biostratigraphic and chemostratigraphic investigations. Black shales of the Ediacaran Miaohe Member in the Yangtze Gorges area contain macroalgae and Ediacara-type macrofossils, both preserved as carbonaceous compressions. The Miaohe Member has been traditionally regarded as an equivalent to Member IV of the uppermost Doushantuo Formation that records part of the Shuram negative δ13Ccarb excursion in South China. Thus, the Miaohe Member may have the potential to clarify the stratigraphic relationship between Ediacara-type fossils and the Shuram excursion. Recently, however, the Miaohe Member has been partially correlated with the Shibantan Member of the Dengying Formation (the “A” correlation) or partially with Member IV (the “Z” correlation). These correlations have different implications for the relationship between the Miaohe Member and the Shuram excursion. Here we test these two correlations using δ13C data of sedimentary kerogen or total organic carbon (δ13CorgTOC), taking advantage of the distinct δ13CorgTOC values between the Shibantan Member and Member IV. Our data show that δ13CorgTOC of the Miaohe Member is more similar to that of Member IV than to Shibantan Member, providing tentative support for the “Z” correlation. In addition, we also analyzed δ13Corg of individual carbonaceous compression fossils (δ13Corgtaxon), which have lower carbon isotope values than TOC. The data suggest that oxygenic photosynthesizers (e.g., macroalgae, cyanobacteria, and eukaryotic phytoplankton) were not the only contributors to sedimentary TOC. Methylotrophs, photoautotrophs, and chemoautotrophs living in anoxic waters or at the oxycline may have fixed recycled CO2 derived from organic carbon remineralization, thus contributing a significant amount of 13C-depleted organic carbon to sedimentary kerogen in the Miaohe Member.

Electron microscopy reveals evidence for simple multicellularity in the Proterozoic fossil Chuaria

Abstract Multicellularity arose multiple times in the evolutionary history of eukaryotes, and simple multicellularity may have a deep history tracing back to the Paleoproterozoic. However, complex multicellular organisms with cellular and tissue differentiation did not appear in the fossil record until the Mesoproterozoic, and it is not until the Ediacaran Period (635–541 Ma) when diverse assemblages of complex multicellular eukaryotes evolved. In the intervening Tonian Period (ca. 1000–720 Ma), the fossil record of multicellular organisms is poorly documented. To address this knowledge gap, we investigated Chuaria and associated carbonaceous compression fossils from the Tonian Liulaobei Formation in North China. These fossils have been variously interpreted as unicellular or multicellular organisms. Our analysis using backscattered-electron scanning electron microscopy (BSE-SEM) revealed direct evidence for simple multicellularity in some of these fossils and suggests that Chuaria may have had a multicellular vegetative stage in its life cycle. This study demonstrates that BSE-SEM has the potential to unveil the hidden diversity of multicellular organisms in the Tonian Period, thus enriching our knowledge about the multiple origins of multicellularity in this critical geological period before Cryogenian glaciations.

The survival of benthic macroscopic phototrophs on a Neoproterozoic snowball Earth

The greatest ice ages in Earth’s history occurred during the 654–635 Ma Marinoan glaciation, when glaciers reached tropical oceans and our planet approached a snowball Earth condition. Paleontological and genomic data suggest that several eukaryotic groups must have survived the Marinoan glaciation. But their fossil record is scarce and limited to microbes, whose ecological and physiological ranges are poorly constrained, thus hampering a full understanding of how and where eukaryotic life—particularly macroscopic phototrophs—survived this snowball Earth. Here we report carbonaceous compression fossils from the Marinoan-age Nantuo Formation in South China. These fossils are preserved in thin black shales sandwiched between glacial diamictites deposited in inner shelf environments of the mid-latitudinal Yangtze block. Some of these fossils are interpreted as benthic macroalgae. Thus, the Marinoan glaciation must have been punctuated by episodes of open waters where habitable benthic substrates were available in the photic zone and along the coast of mid-latitudinal continents. Such open waters may have been the refugia where macroscopic phototrophs survived the Marinoan glaciation and subsequently diversified in the early Ediacaran Period.

The Ediacaran macroalga Zhongbaodaophyton Chen et al. from South China

Wang, Y., Wang, Y., Du, W. & Xang, X., 13.2.2015. The Ediacaran macroalga Zhongbaodaophyton Chen et al. from South China. Alcheringa 39, xxx–xxx. ISSN 0311-5518Zhongbaodaophyton Chen et al. (including two species Z. crassa Chen et al. and Z. robustus Wang et al.) is an Ediacaran macroscopic genus in the Wenghui biota from black shales of the upper Doushantuo Formation (ca 551–593 Ma) in northeastern Guizhou, South China. Zhongbaodaophyton has an unequally dichotomous thallus and a robust holdfast, resembling phaeophytes in morphology; however, its cytological features are not determinable from the carbonaceous compression fossils. Its thallus bears at least nine photosynthetic branches, and its holdfast consists of a globose tuberous rhizome with many unbranched or dichotomous filamentous rhizoids. In addition, these Ediacaran macroalgal species bear pith in the stipe and tuberous rhizome, which is a key feature of phaeophytes. Based on the measurements of 129 Z. robustus specimens, the degree of branching is closely related to the thallus length, the stipe width, and the width ratio of two primary branches; and the stipe width is correlated with the thallus length, the two primary branch widths, and the first branching angle. The thallus grew primarily upward to enhance its competitiveness for sunlight and/or nutrient uptake in the Ediacaran marine ecosystem. A turning point toward reduced growth rates may have occurred in the life history of Z. robustus individuals (at the fourth or fifth axial bifurcation).Yue Wang [gzyuewang@126.com], School of Resources and Environments, Guizhou University, Guiyang 550003, PR China; Ye Wang [gboywangye@126.com] and Xunlian Wang [wxl@cugb.edu.cn], School of Earth Sciences and Resources, China University of Geosciences, Beijing 100083, PR China; Wei Du [duwei@ea.c.u-tokyo.ac.jp], Department of Earth Science and Astronomy, The University of Tokyo, Tokyo 153-8902, Japan.

SEDIMENT EFFECTS ON THE PRESERVATION OF BURGESS SHALE-TYPE COMPRESSION FOSSILS

Experimental burial of polychaete (Nereis) and crustacean (Crangon) carcasses in kaolinite, calcite, quartz, and montmorillonite demonstrates a marked effect of sediment mineralogy on the stabilization of nonbiomineralized integuments, the first step in producing carbonaceous compression fossils and Burgess Shale–type (BST) preservation. The greatest positive effect was with Nereis buried in kaolinite, and the greatest negative effect was with Nereis buried in montmorillonite, a morphological trend paralleled by levels of preserved protein. Similar but more attenuated effects were observed with Crangon. The complex interplay of original histology and sediment mineralogy controls system pH, oxygen content, and major ion concentrations, all of which are likely to feed back on the preservation potential of particular substrates in particular environments. The particular susceptibility of Nereis to both diagenetically enhanced preservation and diagenetically enhanced decomposition most likely derives from the relative lability of its collagenous cuticle vs. the inherently more recalcitrant cuticle of Crangon. We propose a mechanism of secondary, sediment-induced taphonomic tanning to account for instances of enhanced preservation. In light of the marked effects of sediment mineralogy on fossilization, the Cambrian to Early Ordovician taphonomic window for BST preservation is potentially related to a coincident interval of glauconite-prone seas.

Does cement-induced sulfate limitation account for Burgess Shale-type preservation?

In a recent PNAS paper, Gaines et al. (1) proposed a mechanism of early diagenetic sealing and oxidant limitation to account for the preservation of Burgess Shale-type (BST) fossils (i.e., the “exceptional” preservation of carbonaceous compression fossils in Cambrian-aged marine mudstones) (1). It is an interesting hypothesis, but it raises a number of issues.

Megascopic carbonaceous compression fossils from the Neoproterozoic Bhima Basin, Karnataka, South India

Abstract An abundant well-preserved assemblage of annulated carbonaceous compressions and impressions has been recorded from the intra-cratonic argillaceous sedimentary sequence of the Bhima basin in south India. Impressions and carbonaceous compressions recorded in the Hulkal Formation belong to annulated forms similar to the previously reported Sinosabellidites huainanensis , Protoarenicola baiguashanensis and Pararenicola huaiyuanensis from China. This paper discusses the diversity, systematics, affinity, biostratigraphical potential and global significance of these remains. The previously proposed worm-like body fossil affinity for these organisms, based on similarity with the Chinese assemblage, has been reinterpreted as with pre-Ediacaran epibenthic organisms. Fresh investigations of the Indian assemblage of such specimens reveal their close proximity to the algal affinity. The occurrence of global marker events, such as phosphatization, the presence of complicated annulated carbonaceous remains in the Hulkal Formation and the absence of stromatolites in the carbonate sequences of the Bhima basin, collectively indicate a Neoproterozoic, possibly pre-Sturtian age for the Bhima Group.

Restudy of the worm-like carbonaceous compression fossils Protoarenicola, Pararenicola, and Sinosabellidites from early Neoproterozoic successions in North China

The carbonaceous compression fossils Protoarenicola baiguashanensis Wang, 1982, Pararenicola huaiyuanensis Wang, 1982, and Sinosabellidites huainanensis Zheng, 1980, from the early Neoproterozoic Liulaobei and Jiuliqiao formations in northern Anhui, North China, were previously interpreted as worm-like metazoans, largely on the basis of transverse annulations and purported proboscis structures. If correct, these would be some of the earliest known bilaterian animals and would provide a key paleontological calibration to molecular clock analyses. In this study, we examine a large population of these carbonaceous fossils, clarify their taxonomy, and provide new insights into their morphological, paleoecological, and phylogenetic interpretations. Although all three species are characterized by annulated tubes, P. baiguashanensis bears a bulbous terminal structure at one end of its tube. P. huaiyuanensis is characterized by a constricted opening at one end and a closed termination at the other. The two ends of S. huainanensis tubes are both closed and round. The bulbous terminal structure in P. baiguashanensis was previously interpreted as an animal proboscis, but new observations suggest that it was more likely a holdfast structure analogous to discoidal holdfast structures of the Mesoproterozoic Tawuia-like fossil Radhakrishnania Kumar, 2001, and the frondose Ediacara fossil Charniodiscus Ford, 1958. Furthermore, it is possible that at least P. baiguashanensis and P. huaiyuanensis may represent reproductive or taphonomic fragments of the same organism. This reinterpretation weakens the previous interpretation that P. baiguashanensis and P. huaiyuanensis were worm-like bilaterian animals. Instead, they can be alternatively interpreted as erect epibenthic organisms, possibly coenocytic algae reaching a tiering height of 30 mm. The predominance of discoidal holdfasts, as opposed to rhizoidal holdfasts, in pre-Ediacaran epibenthic organisms was probably related to more stable substrates in the presence of microbial mats and in the absence of bioturbating animals.

MACROSCOPIC CARBONACEOUS COMPRESSIONS IN A TERMINAL PROTEROZOIC SHALE: A SYSTEMATIC REASSESSMENT OF THE MIAOHE BIOTA, SOUTH CHINA

Carbonaceous compression fossils in shales of the uppermost Doushantuo Formation (ca. 555-590 Ma) at Miaohe in the Yangtze Gorges area provide a rare Burgess-Shale-type taphonomic window on terminal Proterozoic biology. More than 100 macrofossil species have been described from Miaohe shales, but in an examination of published and new materials, we recognize only about twenty distinct taxa, including Aggregatosphaera miaoheensis new gen. and sp. Most of these fossils can be interpreted unambiguously as colonial prokaryotes or multicellular algae. Phylogenetically derived coenocytic green algae appear to be present, as do regularly bifurcating thalli comparable to red and brown algae. At least five species have been interpreted as metazoans by previous workers. Of these, Protoconites minor and Calyptrina striata most closely resemble animal remains; either or both could be the organic sheaths of cnidarian scyphopolyps, although an algal origin cannot be ruled out for P. minor. Despite exceptional preservation, the Miaohe assemblage contains no macroscopic fossils that can be interpreted with confidence as bilaterian animals. In combination with other late Neoproterozoic and Early Cambrian body fossils and trace fossils, the Doushantuo assemblage supports the view that body-plan diversification within bilaterian phyla was largely a Cambrian event.

Macroscopic carbonaceous compressions in a terminal Proterozoic shale: A systematic reassessment of the Miaohe biota, south China

Carbonaceous compression fossils in shales of the uppermost Doushantuo Formation (ca. 555-590 Ma) at Miaohe in the Yangtze Gorges area provide a rare Burgess-Shale-type taphonomic window on terminal Proterozoic biology. More than 100 macrofossil species have been described from Miaohe shales, but in an examination of published and new materials, we recognize only about twenty distinct taxa, includingAggregatosphaera miaoheensisnew gen. and sp. Most of these fossils can be interpreted unambiguously as colonial prokaryotes or multicellular algae. Phylogenetically derived coenocytic green algae appear to be present, as do regularly bifurcating thalli comparable to red and brown algae. At least five species have been interpreted as metazoans by previous workers. Of these,Protoconites minorandCalyptrina striatamost closely resemble animal remains; either or both could be the organic sheaths of cnidarian scyphopolyps, although an algal origin cannot be ruled out forP. minor.Despite exceptional preservation, the Miaohe assemblage contains no macroscopic fossils that can be interpreted with confidence as bilaterian animals. In combination with other late Neoproterozoic and Early Cambrian body fossils and trace fossils, the Doushantuo assemblage supports the view that body-plan diversification within bilaterian phyla was largely a Cambrian event.

LECLERCQIA COMPLEXA (LYCOPSIDA, MIDDLE DEVONIAN): ITS ANATOMY, AND THE INTERPRETATION OF PYRITE PETRIFACTIONS

The development and/or modification of special acid etching and macerating techniques permits pyritized fossils of Leclercqia complexa to be separated into their carbonaceous wall remains and pyrite lumen—pit casts. These carbonized and pyritic portions can then be studied separately by both light and SEM microscopy and information from both modes compared with that obtainable from carbonaceous compression fossils of L. complexa. This combination of techniques, preservation modes and methods of analysis, allows a synthesis of information obtainable from each, as well as providing a check on errors of interpretation due to preservation mode and method of observation. The developmental sequence of the secondary wall thickenings of the protoxylem elements is shown to be annular to connected annular and spiral, with both exarch and mesarch maturation. The metaxylem shows a transition through reticulate to multiseriate round, oval, and elongate bordered‐pitted tracheids. Different wall facets of separated pyrite tracheid casts can be examined by the SEM and variations in wall structure and pitting characterized. Cells of the outer cortex are 6 to 10 times longer than wide, have thickened vertical walls and end walls that vary from nearly horizontal to angles of overlap of 50 degrees.

Leclercqia complexa (Lycopsida, Middle Devonian): Its Anatomy, and the Interpretation of Pyrite Petrifactions

The development and/or modification of special acid etching and macerating techniques permits pyritized fossils of Leclercqia complexa to be separated into their carbonaceous wall remains and pyrite lumen—pit casts. These carbonized and pyritic portions can then be studied separately by both light and SEM microscopy and information from both modes compared with that obtainable from carbonaceous compression fossils of L. complexa. This combination of techniques, preservation modes and methods of analysis, allows a synthesis of information obtainable from each, as well as providing a check on errors of interpretation due to preservation mode and method of observation. The developmental sequence of the secondary wall thickenings of the protoxylem elements is shown to be annular to connected annular and spiral, with both exarch and mesarch maturation. The metaxylem shows a transition through reticulate to multiseriate round, oval, and elongate bordered-pitted tracheids. Different wall facets of separated pyrite tracheid casts can be examined by the SEM and variations in wall structure and pitting characterized. Cells of the outer cortex are 6 to 10 times longer than wide, have thickened vertical walls and end walls that vary from nearly horizontal to angles of overlap of 50 degrees.