Soft-bodied Fossils Research Articles

New occurrence of the Guanshan Lagerstätte (Cambrian Series 2, Stage 4) in the Kunming area, Yunnan, southwest China, with records of new taxa

As a classic Burgess-Shale-Type Konservat-Lagerstätte, the Guanshan biota is well known for its abundant soft-bodied fossils with high taxonomic diversity. Here, we report a new locality of the Guanshan biota in the Kunming area, Yunnan, South China with several new forms being first documented. In particular: a naraoiid euarthropod, the dorsal exoskeleton of which has characters of both naraoiids and Emucaris; and a discoidal animal, identified as a small eldonioid-like metazoan, which has a counterpart from the Emu Bay Shale Lagerstätte of South Australia, and which further strengths the biogeographic connection between the early Cambrian biotas of South China and Australia. Our new discoveries not only expand the palaeogeographic distribution of the Guanshan biota, but also reveal that its biodiversity is much higher than previously thought, and can be justifiably regarded as a community successor to the Chengjiang biota.Jun Zhao [zhaojunpaleo@126.com], Yunnan Key Laboratory for Palaeobiology, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China; Yunnan Key Laboratory of International Rivers and Transboundary Eco-Security, Yunnan University, Kunming 650091, China; MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China; Yujing Li [yujingli@ynu.edu.cn], Yunnan Key Laboratory for Palaeobiology, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China; MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China; Paul A. Selden [selden@ku.edu], Department of Geology, University of Kansas, 1457 Jayhawk Boulevard, Lawrence, KS 66045, USA; Natural History Museum, Cromwell Road, London SW7 5BD, UK; Peiyun Cong [cong@ynu.edu.cn], Yunnan Key Laboratory for Palaeobiology, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China; MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Institute of Palaeontology, Yunnan University, Kunming, Yunnan, 650091, China.

THE LIMITS OF BURGESS SHALE-TYPE PRESERVATION: ASSESSING THE EVIDENCE FOR PRESERVATION OF THE BLOOD PROTEIN HEMOCYANIN IN THE BURGESS SHALE

Cambrian deposits bearing exceptional preservation of soft-bodied fossils offer unique insights into the diversity and paleobiology of the Cambrian biota. In recent years, a growing number of studies have provided evidence that highly labile tissues, such as nervous tissues, may be more common in fossils from Burgess Shale-type deposits than previously appreciated. These discoveries have provided novel insights into the anatomy and phylogeny of Cambrian animals and have also challenged the basic understanding of the limits on the resolution of anatomical information that may be captured by Burgess Shale-type preservation. Evidence used to infer primary blood chemistry was recently reported from the iconic Burgess Shale fossil Marrella splendens, the first such report from a Cambrian Lagerstatte. Diminutive crystals of a copper sulfide mineral found in association with Marrella were interpreted as evidence that the animal's blood contained the Cu-bearing protein hemocyanin. Copper minerals in shales in general, and in the Burgess Shale in particular, however, may have a complex geologic history. Using microscopy and SEM-EDS elemental mapping of material from the Walcott Quarry as well as from the Burgess Shale fossil quarry at Marble Canyon, we assess the timing of emplacement of Cu-sulfides in fossil-bearing mudrocks of the Burgess Shale. These data demonstrate that copper minerals are most conspicuously concentrated within veins that are oriented sub-perpendicular to bedding. These late-stage fluid flow paths allowed Cu-mineralizing fluids to infiltrate bedding plane-parallel cracks, which commonly run through Burgess Shale fossils and extend into the matrix around them. Based upon the relationships of mineral phases associated with fossils, it appears that Cu-mineralization occurred after metamorphic volatilization of a large proportion of the carbonaceous material that originally comprised the fossils. We find that copper minerals in the Burgess Shale are of demonstrably post-Cambrian origin, a secondary overprint that cannot be used to infer original blood chemistry or tissue composition of its fossils.

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 (635-539Ma). 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.

Soft-bodied fossils from the upper Valongo Formation (Middle Ordovician: Dapingian-Darriwilian) of northern Portugal.

Soft-bodied preservation is common in the Cambrian but comparatively rare in the Ordovician. Here, a new deposit preserving soft-bodied fossils is reported from the Middle Ordovician (Dapingian-Darriwilian) upper Valongo Formation of northern Portugal. The deposit contains the first known occurrences of soft-bodied fossils from the Middle Ordovician (Dapingian-Darriwilian) of Portugal and is the first Ordovician example of soft-tissue preservation involving carbonaceous films from the Iberian Peninsula. It also represents the lone deposit of soft-bodied fossils from the Middle Ordovician of northern Gondwana. Thus temporally, it lies between the exceptional deposits of the Lower Ordovician of Fezouata (Morocco) and the Upper Ordovician of the Soom Shale (South Africa); it also serves as a biogeographic link between these and the various Ordovician soft-bodied deposits in Laurentia. The soft-bodied fossils come from the deep-water slates of the upper part of the Valongo Formation and include a discoidal fossil questionably referable to Patanacta, wiwaxiid sclerites, and a possible pseudoarctolepid arthropod.

First report on Guanshan Biota (Cambrian Stage 4) at the stratotype area of Wulongqing Formation in Malong County, Eastern Yunnan, China

Exceptionally preserved fossils, such as those from Cambrian Burgess Shale-type fossil-Lagerstätten are critical because of their unique contributions to knowledge of the phylogenetic radiation and palaeoecological expansion of metazoans during the Cambrian explosion. Critically, these deposits provide information that is usually unobtainable from shelly and skeletonized fossils alone. The Guanshan Biota (Cambrian Series 2, Stage 4) in the Yunnan Province of South China, has produced abundant and diverse, exquisitely preserved fossils that often retain soft tissues and organs. To date, most fossils from the Guanshan Biota have been collected from localities such as Gaoloufang and Gangtoucun, which have become inaccessible due to new urban expansions and constructions of residential buildings. Here we present the first report of soft bodied fossils from a new section at Kanfuqing, close to the Wulongqing village in Malong County, approximately 3 km east of the Wulongqing Formation stratotype section. Fossils retain soft morphology, and include brachiopods with delicate marginal setae, priapulids with well-preserved sclerites and vetulicolians with entire sections of body. In addition, this fauna includes rare occurrences of trilobites preserved with soft tissues replicated as pyrite pseudomorphs after weathering. This discovery represents an important palaeogeographical extension of soft-bodied fossils of the Guanshan fauna to the east of the Xiaojiang Fault (related to Tsinning tectonic movements ca. 700 Ma). The fauna from the new Kanfuqing section is similar to that reported from the Wulongqing Formation west of the Xiaojiang Fault, and thus has significant implications for early Cambrian palaeogeography, faunal successions and palaeoenvironments of eastern Yunnan.

Presence of Naraoia Walcott, 1912 (Nektaspida, Arthropoda) in the middle Cambrian of Europe (Murero, NE Spain)

The genus Naraoia Walcott, 1912, a Burgess Shale-type fossil known from the lower and middle Cambrian of British Columbia (Canada), Idaho and Utah (USA), as well as from Yunnan and Guizhou provinces (China), is now reported from the middle Cambrian of Murero (Zaragoza, Spain), which is the first record in the Acadobaltic province. The only fragmented specimen found is determined as Naraoia sp., its age being Pardailhania multispinosa Zone (Drumian Stage). This new datum reinforces the hypothesis of the existence of a cosmopolitan faunal substrate in early Cambrian times, which is to some extent refl ected in the mid Cambrian by faunal groups of low evolutionary potential as the family Naraoiidae and other soft-bodied fossil taxa.

Soft-bodied fossil of a lizard from the Parachute Creek Member, Green River Formation (Eocene), Utah

A rare specimen of soft tissue preservation of a lizard from the Parachute Creek Member of the Eocene Green River Formation, Uinta Basin, Utah, is described. The preservation is unusual in that it is a miner­alized body lacking the skeleton. This, and other small boneless vertebrate specimens also from the Para­chute Creek, indicate occasional demineralizing conditions in Lake Uinta, but not apparently in the other two lakes of the Green River Formation—Fossil Lake and Lake Gosuite.

Influence of redox conditions on animal distribution and soft-bodied fossil preservation of the Lower Cambrian Chengjiang Biota

The exceptionally preserved Chengjiang Biota (Yunnan, China) is significant for understanding the rapid development of complex animal-rich ecosystems during the evolutionary radiation of the Cambrian. However, the ecological signal provided by the fossils captured in this deposit may not reflect accurately the in-life community, with transport and decay of carcasses being the principal processes responsible for potential modification. The principal fossil-bearing interval (Maotianshan Shale Member, Yu'anshan Formation) is comprised of claystones of two distinct depositional origins: the “background” beds represent slow hemipelagic deposition into deep waters, while the “event” beds represent distal turbidites or storm-generated beds. Each bed type has a distinct fossil assemblage and preservation mode underscoring the importance of interpreting the palaeoenvironment of each bed type. Here, we interpret palaeo-redox conditions for both the background beds and the event beds by conducting a systematic geochemical study using iron speciation, δ34Spy, δ13Ccarb,δ13CTOC and molybdenum abundance. These data are from the most complete core recovered from the Chengjiang fossiliferous units and allow us to distinguish redox conditions where the animals lived, and where they were buried and exceptionally preserved. Our results demonstrate that background beds were dominated by dysoxic conditions; here, a diverse sponge community tolerant of low-oxygen conditions lived. In contrast, the shallower shelf and offshore-transitional environments, where event beds were sourced, were almost persistently oxic, and it was here where the diverse Chengjiang Biota flourished. Thus, the Chengjiang sediments record two distinct palaeocommunities within the background and the event beds. The contrasting redox conditions in close spatial proximity likely facilitated the soft-bodied fossil preservation. Our findings provide a valuable case study in the necessity to understand ecological composition and exceptional preservation informed by environmental context.

Interns and Volunteers Crucial in Curating and Digitizing Fossil Invertebrates in the Field Museum’s Fast Growing Mazon Creek Collection

The Mazon Creek region in Northeastern Illinois is home to a Middle Pennsylvanian (~307 million years old) soft-bodied fossil Lagerstätte of animals and plants that lived along a subtropical swampy coastline. This area was strip mined for coal from 1928 to 1974 and museum geologists and amateur collectors acquired large fossil collections during this time by collecting and splitting millions of nodules unearthed at the mines. These large collections are important because of the rarity of many of the species in the Mazon Creek biota. There are about 250 described fossil invertebrate species from the Mazon Creek region. Fifty-one of these species (mostly insects and arachnids) are represented by just a single specimen in the Field Museum’s collection. Since the 1980’s collecting has decreased and the mines have been restored to parks and wildlife areas. The Field Museum maintained a collection of 34,000 Mazon Creek invertebrate fossil for many decades. With the new donations from private collectors in the last three years this collection has grown by 20% and now represents 18% of the Fossil Invertebrate systematic collection. The Mazon Creek is also the most used fossil invertebrate collection accounting for about 38% of loans in the last five years. Dealing with these large and often unexpected donations adds to the already large workload of the collection staff, so interns and volunteers are utilized to process, catalog, digitize, and integrate these fossils into the museum’s collection. In the summer of 2016, interns Mackenzie Best and Yaal Dryer unpacked and sorted into drawers the Thomas V. Testa collection, and digitized the first 1,000 fossils. In 2017, two Women in Science interns, Kate Hodge and Dana Kahn, spent 6 weeks entering the data for 5,000 fossils into our database, numbering these fossils, and printing their labels. Having a well curated collection, as well as volunteer Jack Wittry, who has expert knowledge of Mazon Creek fossils, has also been crucial to the success of these projects. Mane Pritza, a Field Museum volunteer, began photographing these collections and has captured over 11,000 images. Janel Nelson, a former volunteer, has uploaded these images into our multimedia database and linked them to the corresponding records in the catalog module. James and Sylvia Konecny donated their 4,000-specimen Mazon Creek collection in December of 2017, ensuring that interns and volunteers will continue their curation work for at least the next two years.

Origin and the Early Evolution of the Phylum Mollusca

The paper gives an overview of the modern hypotheses on the origin of the phylum Mollusca and the formation of its main classes. The Cambrian stage of molluscan evolution is characterized based on the paleontological material. The doubtfulness of assignment of the Precambrian (Vendian) soft-bodied fossil Kimberella to mollusks is argued. Judging from the interpretation of the morphologically diverse Cambrian fossils, it is suggested that the classes Polyplacophora, Monoplacophora, Gastropoda and Bivalvia formed already near the Precambrian–Cambrian boundary, i.e., from the beginning of the paleontologically documented evolutionary history of the phylum, whereas the assumption of the later origin of these taxa is unconvincing. The remaining classes of mollusks arose later, i.e., Cephalopoda, in the Late Cambrian; Scaphopoda, in the Ordovician; and Aplacophora, in the Silurian.

New Megasphaera-like microfossils reveal their reproductive strategies

New occurrence of Ediacaran Weng’an Biota-type microfossil assemblage has been discovered from the base of the Dengying Formation in Zhenba County, Shaanxi Province, South China. The assemblage is dominated by spherical microfossils ranging from 200 to 1200μm in diameters. Based on the size and morphological features, two genera and three species have been identified, i.e. Megasphaera inornata, M. ornata and Spiralicellula bulbifera. Morphological analyses reveal three types of reproduction, i.e. binary fission, budding, and multiple fission, which are widely distributed among protist and metazoan lineages. Besides, a physiological phenomenon of “ornament growth” has been discovered among M. ornata. These again raise difficulties in determining biological affinities of Megasphaera-like microfossils. The new microfossil assemblage shares the most common elements of embryo-like microfossils with the Weng’an biota. However, those Large Acanthomorphic Acritarchs (LAAs) with great capacity for biostratigraphic correlation are not present. The age of the new assemblage is constrained within the Ediacaran, contemporary with or slightly younger than the Weng’an Biota by regional correlation. The new finding not only extends the geographical distribution of the Ediacaran phosphatized biotas in South China, but also provides valuable fossil materials for studying ontogenesis and evolution of early microscopic organisms and taphonomy of phosphatized soft-bodied fossil Lagerstätte.

Disc-shaped fossils resembling porpitids or eldonids from the early Cambrian (Series 2: Stage 4) of western USA.

The morphology and affinities of newly discovered disc-shaped, soft-bodied fossils from the early Cambrian (Series 2: Stage 4, Dyeran) Carrara Formation are discussed. These specimens show some similarity to the Ordovician Discophyllum Hall, 1847; traditionally this taxon had been treated as a fossil porpitid. However, recently it has instead been referred to as another clade, the eldonids, which includes the enigmatic Eldonia Walcott, 1911 that was originally described from the Cambrian Burgess Shale. The status of various Proterozoic and Phanerozoic taxa previously referred to porpitids and eldonids is also briefly considered. To help ascertain that the specimens were not dubio- or pseudofossils, elemental mapping using energy dispersive X-ray spectroscopy (EDS) was conducted. This, in conjunction with the morphology of the specimens, indicated that the fossils were not hematite, iron sulfide, pyrolusite, or other abiologic mineral precipitates. Instead, their status as biologic structures and thus actual fossils is supported. Enrichment in the element carbon, and also possibly to some extent the elements magnesium and iron, seems to be playing some role in the preservation process.

The same picture through different lenses: quantifying the effects of two preservation pathways on Green River Formation insects

AbstractInsects in the fossil record are generally preserved in lacustrine shales or in amber. For those in lacustrine shales, preservation is usually via keroginization or mineralization. Given the extended period of microbial decay required to generate ions for mineralization, there is a predicted inherent bias toward lower preservation quality for this pathway by most taphonomic indices compared with keroginization. This study tests this hypothesis by comparing multiple measures of preservation quality between sites with similar sedimentology in the Eocene Green River Formation of Colorado. Here, insects are either mineralized in iron oxides (likely after pyrite) at the Paleoburn site or keroginized at the Anvil Points site.Generally, the prediction that keroginization preserves soft-bodied fossils with higher preservational quality than mineralization is affirmed, but with some caveats. Beetles, known for their robust cuticles, are proportionately more abundant at the Paleoburn site, but eight of the nine orders recorded are shared between sites. As predicted, insects show lower preservation fidelity at the Paleoburn site, but they also show higher degrees of disarticulation. This second bias should be acquired primarily during the biostratinomy stage, and not early diagenesis. Nonetheless, higher-energy biostratinomic conditions may be compatible with taphonomic conditions that promote mineralization over keroginization.Comparing the inherent taphonomic bias of different preservation pathways is often difficult, since fossil deposits may be preserved millions of years or thousands of kilometers apart. By studying two different preservation pathways of insects within the same formation, we can affirm that keroginization does indeed preserve recalcitrant organic matter with higher quality than pyritization or iron-oxide mineralization. Additionally, some guidelines can be proposed concerning the body parts and taxa that can be compared, and for what purpose, when contrasting mineralized and keroginized soft-bodied deposits.

An Outline and New Aspects of the Regional Stratigraphy of Mongolia

This paper concerns deals with the Mongolian regional stratigraphic classification of sedimentary, sedimentary-volcanogenic sediments, and metamorphic complex spread over the territory of Mongolia according to the new International Chronostratigraphic Chart [1]. The regional stratigraphy of Precambrian-Cenozoic rocks of the Mongolian territory was studied by Dorjnamjaa et al. [2]. So, this paper provides for the stratigraphy and correlation of key sections through the Archean-Proterozoic-Mesozoic-Cenozoic of various areas of Mongolia. In this paper, the author introduces the most important results indicating the key reference sections for the first time and put into a broad regional context. The characterization and geologic mapping of lithological units provided an essential basis for the lithostratigraphy, biostratigraphy and chemostratigraphy. Especially key biostratigraphic aspects are the phenomenon of faunal succession. The basic lithostratigraphic unit is the formations and groups which have been successful mapped. At present in Mongolia for Precambrian period (systems and series) 30 formations and groups, for Paleozoic—about 250 formations and groups, for Mesozoic—102 formations and groups, for Cenozoic—27 formations, members, and beds are distinguished. The auxiliary stratigraphic units are members, beds which should be mapped depending on great extent of detailed study. In the regional stratigraphical classification, there are two primary kinds of stratotypes [3]: a) unit stratotypes, which serve as the standard of definition for a stratigraphic unit, and b) boundary stratotypes, which serve as the standard of definition for a stratigraphic boundary. These aspects, in particular gradational boundaryas far as possible were examined for example for Archean-Paleoproterozoic (by geochronologic data), Ediacaran-Early Cambrian (by soft-bodied fossils, trace fossils and small shelly fossils), Ordovician-Silurian (by beyrichiidostracodes), Devonian-Carboniferous (by conodonds), Paleocene-Eocene (by mammals), Oligocene-Miocene (by small rodents) systems and series.

A new model of the formation of Pennsylvanian iron carbonate concretions hosting exceptional soft-bodied fossils in Mazon Creek, Illinois.

Preservation of Pennsylvanian-aged (307Ma) soft-bodied fossils from Mazon Creek, Illinois, USA, is attributed to the formation of siderite concretions, which encapsulate the remains of terrestrial, freshwater, and marine flora and fauna. The narrow range of positive δ34 S values from pyrite in individual concretions suggests microenvironmentally limited ambient sulfate, which may have been rapidly exhausted by sulfate-reducing bacteria. Tissue of the decaying carcass was rapidly encased by early diagenetic pyrite and siderite produced within the sulfate reduction and methanogenic zones of the sediment, with continuation of the latter resulting in concretion cementation. Cross-sectional isotopic analyses (δ13 C and δ18 O) and mineralogical characterization of the concretions point to initiation of preservation in high porosity proto-concretions during the early phases of microbially induced decay. The proto-concretion was cemented prior to compaction of the sediments by siderite as a result of methanogenic production of 13 C-rich bicarbonate-which varies both between Essex and Braidwood concretions and between fossiliferous and unfossiliferous concretions. This work provides the first detailed geochemical study of the Mazon Creek siderite concretions and identifies the range of conditions allowing for exceptional soft-tissue fossil formation as seen at Mazon Creek.

The eyes of Tullimonstrum reveal a vertebrate affinity.

Tullimonstrum gregarium is an iconic soft-bodied fossil from the Carboniferous Mazon Creek Lagerstätte (Illinois, USA). Despite a large number of specimens and distinct anatomy, various analyses over the past five decades have failed to determine the phylogenetic affinities of the 'Tully monster', and although it has been allied to such disparate phyla as the Mollusca, Annelida or Chordata, it remains enigmatic. The nature and phylogenetic affinities of Tullimonstrum have defied confident systematic placement because none of its preserved anatomy provides unequivocal evidence of homology, without which comparative analysis fails. Here we show that the eyes of Tullimonstrum possess ultrastructural details indicating homology with vertebrate eyes. Anatomical analysis using scanning electron microscopy reveals that the eyes of Tullimonstrum preserve a retina defined by a thick sheet comprising distinct layers of spheroidal and cylindrical melanosomes. Time-of-flight secondary ion mass spectrometry and multivariate statistics provide further evidence that these microbodies are melanosomes. A range of animals have melanin in their eyes, but the possession of melanosomes of two distinct morphologies arranged in layers, forming retinal pigment epithelium, is a synapomorphy of vertebrates. Our analysis indicates that in addition to evidence of colour patterning, ecology and thermoregulation, fossil melanosomes can also carry a phylogenetic signal. Identification in Tullimonstrum of spheroidal and cylindrical melanosomes forming the remains of retinal pigment epithelium indicates that it is a vertebrate; considering its body parts in this new light suggests it was an anatomically unusual member of total group Vertebrata.

Palaeoscolecid worms from the Lower Ordovician Fezouata Lagerstätte, Morocco: Palaeoecological and palaeogeographical implications

The Lower Ordovician Fezouata Lagerstätte from Morocco (central Anti-Atlas, Zagora area) has yielded abundant and diverse soft-bodied fossils. Most described taxa were epibenthic or pelagic, so that little is known about the endobenthic components of the fauna. Here we report the discovery of a locally abundant element of the biota, a palaeoscolecid worm, which may have played an important ecological role in the community. Palaeoscolecids had a long, annulated body with an eversible tooth-bearing pharynx. They probably represent stem priapulids, but unlike them, they were protected by biomineralized (phosphatic) microscopic plates covering most of the body. These sclerites are commonly found isolated within SSF assemblages, thus considerably extending the fossil record of the group that ranges from the Cambrian Series 2 (Stage 3) to the upper Silurian. Like priapulids, palaeoscolecids might have been important bioturbators, contributing to the substrate colonization in the lower Palaeozoic. The discovery of abundant material (38 specimens) in the Fezouata Shale (Lower Ordovician, Morocco) allows us to reconsider their possible lifestyle, habitat and feeding habits. All the specimens belong to Palaeoscolex? tenensis, a species previously only known from a fragment of cuticle from the Floian of Bohemia (Czech Republic). Some exhibit remains of the gut and aboral spines, features that are highly informative with regard to the ecology of these extinct organisms, but rarely preserved. The phosphatic nature of the plates is demonstrated by compositional analyses and interpreted with regard to the diagenetic context of the Fezouata Shale as primary. We also hypothesize that (at least) some palaeoscolecids were makers of Tomaculum-type ichnofossils, which are elongate clusters of faecal pellets, and show that they ingested a notable amount of sediment during the normal course of feeding. The presence of Palaeoscolex? tenensis in the Lower Ordovician of Morocco further documents the great faunal similarities between Perunica and northern Gondwana at that time.

The 'Tully monster' is a vertebrate.

Problematic fossils, extinct taxa of enigmatic morphology that cannot be assigned to a known major group, were once a major issue in palaeontology. A long-favoured solution to the 'problem of the problematica', particularly the 'weird wonders' of the Cambrian Burgess Shale, was to consider them representatives of extinct phyla. A combination of new evidence and modern approaches to phylogenetic analysis has now resolved the affinities of most of these forms. Perhaps the most notable exception is Tullimonstrum gregarium, popularly known as the Tully monster, a large soft-bodied organism from the late Carboniferous Mazon Creek biota (approximately 309-307 million years ago) of Illinois, USA, which was designated the official state fossil of Illinois in 1989. Its phylogenetic position has remained uncertain and it has been compared with nemerteans, polychaetes, gastropods, conodonts, and the stem arthropod Opabinia. Here we review the morphology of Tullimonstrum based on an analysis of more than 1,200 specimens. We find that the anterior proboscis ends in a buccal apparatus containing teeth, the eyes project laterally on a long rigid bar, and the elongate segmented body bears a caudal fin with dorsal and ventral lobes. We describe new evidence for a notochord, cartilaginous arcualia, gill pouches, articulations within the proboscis, and multiple tooth rows adjacent to the mouth. This combination of characters, supported by phylogenetic analysis, identifies Tullimonstrum as a vertebrate, and places it on the stem lineage to lampreys (Petromyzontida). In addition to increasing the known morphological disparity of extinct lampreys, a chordate affinity for T. gregarium resolves the nature of a soft-bodied fossil which has been debated for more than 50 years.

Decay and mineralization of soft tissue in Early Devonian boring ctenostome bryozoans

The Early Devonian of Podolia, Ukraine, has yielded phosphatized colonies of the boring ctenostome bryozoan Podoliapora doroshivi with 3-D preservation of soft tissues. However, the feeding zooids are not anatomically complete, their preserved soft tissues comprising decay-resistant structures such as the protective cuticular polypide sacs with presumed parietal muscles inside the wall of the sacs, the setigerous collars, the membranous orificial walls and remains of the muscle tissues. Early diagenetic apatite mineralization occured in numerous feeding zooids of Podoliapora at different stages of decay and may be important for the interpretation of decay processes in these colonial soft-bodied fossil organisms. A setigerous collar, which is a characteristic of extant ctenostomes, occurs in P. doroshivi in several stages of decay showing progressive collapse and eventual complete loss. This study indicates that the morphological changes of collars induced by decay often resulted in connection with the membranous orificial wall, producing false anatomical structures, unrelated to structures observed in the earlier stages of decay or to the anatomical structures of extant ctenostomes. The most decay-resistant cuticular polypide sacs mineralized as cryptocrystalline apatite in early stage of decay became degraded in later stages of decay. These data provide evidence that the anatomical interpretation of soft-bodied fossils preserved only in the later stages of decay may have led to imprecise morphological interpretations.

The Cambrian explosion

The sudden appearance of fossils that marks the so-called 'Cambrian explosion' has intrigued and exercised biologists since Darwin's time. In On the Origin of Species, Darwin made it clear that he believed that ancestral forms 'lived long before' their first fossil representatives. While he considered such an invisible record necessary to explain the level of complexity already seen in the fossils of early trilobites, Darwin was at a loss to explain why there were no corresponding fossils of these earlier forms. In chapter 9 of the Origin, entitled 'On the imperfection of the geological record', he emphasized the 'poorness of our palaeontological collections' and stated categorically that 'no organism wholly soft can be preserved'. Fortunately much has been discovered in the last 150 years, not least multiple examples of Cambrian and Precambrian soft-bodied fossils. We now know that the sudden appearance of fossils in the Cambrian (541-485 million years ago) is real and not an artefact of an imperfect fossil record: rapid diversification of animals coincided with the evolution of biomineralized shells. And although fossils in earlier rocks are rare, they are not absent: their rarity reflects the low diversity of life at this time, as well as the low preservation potential of Precambrian organisms (see Primer by Butterfield, in this issue).