Fossilized eggs from the Pennsylvanian of Illinois

Nine siderite concretions from the Middle Pennsylvanian, Francis Creek Shale (Carbondale Formation, Desmoinesian Series, West‐phalian C‐D), in the Mazon Creek area, Will‐Kankakee Counties, Illinois, U.S.A. preserve clusters of impressions of small eggs. Differential staining of the matrix suggests that the eggs were originally spawn within gelatinous strings containing 1 or 2 rows of eggs. Unfortunately, these egg impressions lack the diagnostic features needed to identify the zoological taxon (taxa) from which they originated.     

A new cephalopod with soft parts from the Upper Carboniferous Francis Creek Shale of Illinois, USA

A ten armed fossil from the Pennsylvanian Francis Creek Shale and belonging to the famous Mazon Creek biota is described as a cephalopod. Apart from a superficial resemblance to the coleoids, there is little basis for an assessment of the animal's affinities. A mineralized Bask shaped structure preserved in calcite and seen in both part and counterpart in fragmentary form may represent the fossilized remains of an internal organ such as the stomach or ink sac A small, dark circular spot positioned anterior to the appendages could be the animal's eye although mere is no internal structure visible. The extremely rare occurrence of Palaeozoic cephalopods with soft parts has so far been Ended to the Francis Creek Shale and the Devonian Hunsrückschiefer of Germany. The Mazon Creek biota includes many fossils which show a remarkable standard of soft part preservation which in some cases includes the fossilization of eye spots and stomach traces. Previously deserted Mazon Creek cephalopods have included the tentacular impression of a squid complete with arm hooks, and the radulae and shells of both coleoids and nautiloids. The specimen described here is the first from the Palaeozoic to show any hint of organ preservation.     

A synopsis of medullosan pollen organs from the middle Pennsylvanian Mazon creek flora of northeastern Illinois, USA

Six kinds of medullosan pollen organs containing Monoletes-type pollen (prepollen) are described from the middle Pennsylvanian Mazon Creek flora of northeastern Illinois, USA. Codonotheca caduca Sellards and Schopfitheca boulayoides Delevoryas are redescribed and emended diagnoses and new reconstructions are provided. Three forms of Dolerotheca-type pollen organs are described for the first time, along with ultrastructural details of in situ pollen. A sixth kind of pollen organ, tentatively associated with the ovule Stephanospermum konopeonus (Langford) Drinnan, Schramke and Crane, is too poorly preserved for generic assignment but has pollen different from that of other Mazon Creek medullosan pollen organs. This new information on Mazon Creek plants provides important evidence on the diversity of medullosan “pteridosperms” in the flora. Characterization of in situ pollen provides a basis for continuing attempts to associate pollen organs with the variety of co-occuring dispersed ovules.     

STUDIES OF PALEOZOIC MARATTIALEANS: AN EARLY PENNSYLVANIAN SPECIES OF THE FERTILE FERN SCOLECOPTERIS

Remains of the fossil Marattiales are very rare in Lower Pennsylvanian sediments. The present report describes a new species of the fertile fern foliage Scolecopteris from the Lewis Creek, Kentucky locality (Lower or lower Middle Pennsylvanian). Scolecopteris conicaulis n. sp. has radial synangia composed of a ring of 4–7 elongate, exannulate sporangia. Most features of the synangia of S. conicaulis were previously hypothesized to be primitive in Scolecopteris based on geologically younger species. Supposed primitive characters include the large synangium pedicel with fiber core, an outer-facing sporangial wall lacking differentiation or zonation, and large spores. The anatomy of the sporangium walls, pinnule morphology, and general spore type support an association with the Minor group of Scolecopteris. The new species is similar in several important features to Scolecopteris (Cyathotrachus) altus, the only other anatomically preserved fertile marattialean known from this early time, and indicates a considerably earlier origin for fertile foliage of this type.     

STIGMARIAN PETRIFACTIONS FROM THE PENNSYLVANIAN OF COLORADO

The designation Stigmaria ficoides is restricted to casts and impressions. Petrifactions of Stigmaria arnoldii sp. n. occur in Pennsylvanian age strata in Trout Creek Pass, Colorado. The rootlet gaps of this species appear narrow and elongate in both cross and longitudinal sections. Rootlet traces are contiguous with the secondary xylem to its outer margin and are surrounded by a small parenchyma sheath in the middle cortex. The outer cortex consists of an inner layer of primary tissue and a relatively thick zone of periderm. S. arnoldii most closely resembles material described as S. ficoides from England. Various stigmarian characters are evaluated with regard to their systematic or ontogenetic significance.     

ZOSTEROGRAMMIDA, A NEW ORDER OF MILLIPEDES FROM THE MIDDLE SILURIAN OF SCOTLAND AND THE UPPER CARBONIFEROUS OF EURAMERICA

Abstract:  New Palaeozoic millipedes, Zosterogrammus stichostethus gen. et sp. nov. and Casiogrammus ichthyeros gen. and sp. nov., are described from the Francis Creek Shale Member of the Carbondale Formation (Upper Carboniferous: Pennsylvanian), Mazon Creek, Illinois, and the Hagshaw Hills Inlier (Middle Silurian: Wenlock) of the Midland Valley of Scotland, respectively. These millipedes, together with Purkynia lata Fritsch from the Upper Carboniferous (Westphalian D) of Nýřany, Czech Republic, are placed in the new family Zosterogrammidae within the new order Zosterogrammida. All of these millipedes have very broad terga with a microsculpture consisting of fine transverse terrace lines along the anterior of the terga and oblique terrace lines across the remainder. Zosterogrammida have a trunk-ring architecture consisting of an arched diplotergite, a pair of free ventral diplopleurites and a pair of free ventral sternites, indicating a relatively basal phylogenetic position within Chilognatha. Although the exact phylogenetic position is indeterminate, Zosterogrammus stichostethus has divided sternites, indicating a possible affinity with the Pentazonia. The Hagshaw Hills millipede contributes significantly to the high-rank diversity of Middle Silurian millipedes as all previously described taxa of this age belong to the Archipolypoda.     

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

Preservation of Pennsylvanian‐aged (307 Ma) 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 δ³⁴S 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 (δ¹³C and δ¹⁸O) 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 ¹³C‐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.     

Chemical signatures of soft tissues distinguish between vertebrates and invertebrates from the Carboniferous Mazon Creek Lagerstätte of Illinois

The chemical composition of fossil soft tissues is a potentially powerful and yet underutilized tool for elucidating the affinity of problematic fossil organisms. In some cases, it has proven difficult to assign a problematic fossil even to the invertebrates or vertebrates (more generally chordates) based on often incompletely preserved morphology alone, and chemical composition may help to resolve such questions. Here, we use in situ Raman microspectroscopy to investigate the chemistry of a diverse array of invertebrate and vertebrate fossils from the Pennsylvanian Mazon Creek Lagerstätte of Illinois, and we generate a ChemoSpace through principal component analysis (PCA) of the in situ Raman spectra. Invertebrate soft tissues characterized by chitin (polysaccharide) fossilization products and vertebrate soft tissues characterized by protein fossilization products plot in completely separate, non‐overlapping regions of the ChemoSpace, demonstrating the utility of certain soft tissue molecular signatures as biomarkers for the original soft tissue composition of fossil organisms. The controversial problematicum Tullimonstrum, known as the Tully Monster, groups with the vertebrates, providing strong evidence of a vertebrate rather than invertebrate affinity.     

Saccoglossus testa from the Mazon Creek fauna (Pennsylvanian of Illinois) and the evolution of acorn worms (Enteropneusta: Hemichordata)

The limited fossil record of enteropneust hemichordates (acorn worms) and the few external features that distinguish the four families have provided a challenge to our understanding of the evolution of the group and their various feeding adaptations. The middle Pennsylvanian Saccoglossus testa sp. nov. from the Mazon Creek, Westfalian D Carbonate Formation, Francis Creek Shale of northern Illinois provides evidence for the exploitation of surface sediments. Saccoglossus testa has a long proboscis characteristic of the extant genus Saccoglossus, a specialist in surface deposit feeding. The collar is as long as it is wide. The anterior trunk lacks a distinctively wide branchial region. These three features distinguish it from its sympatric enteropneust species Mazoglossus ramsdelli Bardack that has a proboscis characteristic of an infaunal deposit feeder. It is the seventh known species of fossil enteropneust, including a resting trace of a Lower Triassic fossil that has collar lips that characterize the extant deep‐sea family Torquaratoridae, and which represents a second parallel evolution of surface deposit feeding. An analysis of the seven fossils shows that the earliest Enteropneusta had a relatively simple harrimaniid‐like body plan, and that the spengelid, the torquaratorid and lastly the most complex ptychoderid body plan appeared in that chronological order.     

ETAPTERIS LECLERCQII SP. N., A ZYGOPTERID FERN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The zygopterid fern Etapteris leclercqii sp. n. is described from Lower Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. Isolated petioles extend up to 15.0 cm in length and 5.0 mm in diam; no lateral axes have been observed. The clepsydroid-shaped petiole trace is characterized by lateral arms that sharply taper and by the production of peripheral loops prior to the separation of pinna traces from the stele. Large irregularly shaped multicellular hairs are randomly scattered over the surface of the petioles. When compared with other currently recognized zygopterid ferns, E. leclercqii appears most similar to Metaclepsydropsis duplex. Based on the configuration of the leaf trace and the occurrence of peripheral loops, it is suggested that E. leclercqii may represent the most primitive species of the genus known to date.     

THE ULTRASTRUCTURE OF PALEOZOIC FERN SPORES: I. BOTRYOPTERIS

The spores of four species of the Paleozoic filicalean fern Botryopteris are examined at the ultrastructural level. Spores of B. cratis, B. forensis, B. globosa, and an unnamed species from the Lower Pennsylvanian, are compared on the basis of sporoderm stratification and the presence or absence of a sculptine layer. The species examined differ widely as to the type of reproductive unit in which they are borne and include forms that range throughout the Pennsylvanian. In all species the exine is homogeneous, lacking cavities and lamellae. A thin nexine is present in the Middle and Upper Pennsylvanian taxa, but is absent in the Lower Pennsylvanian spores. Only one spore type (B. cratis) possesses a clearly defined sculptine layer. Features of the sporoderm are compared with those of extant, homosporous pteridophyte spores.     

Discovery of radiolaria and conodonts in the Carboniferous–Permian of San Salvador Patlanoaya (Puebla,Mexico); biostratigraphic implications

New identifications of radiolaria and conodonts allow clarifying the biostratigraphy of the Pennsylvanian and Permian deposits in San Salvador Patlanoaya (Puebla State, Mexico). The radiolaria are sometimes relatively common in the series, but weakly diversified and endemic. Among the conodonts, Streptognathodus bellus is characteristic of the Late ‘Virgilian’ sensu Baars, probably coeval with the Early Wolfcampian sensu Thompson or Wilde, the Bursumian of Ross & Ross or the Orenburgian–Asselian stage or substage of Russia (sensu Davydov). San Salvador Patlanoaya is a key-section for the Pennsylvanian–Permian boundary in Mexico.     

STUDIES OF PALEOZOIC SEED FERNS: ANATOMY AND MORPHOLOGY OF MICROSPERMOPTERIS APHYLLUM

The discovery of numerous specimens of the monostelic pteridosperm genus Microspermopteris in Pennsylvanian coal ball petrifactions from the Lewis Creek and What Cheer localities provides additional information about the anatomical and morphological variability within the genus. Specimens are now known up to 1.1 cm in diam that bear epidermal appendages in the form of variously-shaped trichomes. The external surface of the stem is further ornamented by longitudinal flaps of cortical tissue. Petioles exhibiting a single C-shaped vascular strand with abaxial protoxylem are produced in a 2/5 phyllotaxy. Large petiole bases that clasp the stem produce primary pinnae alternately. The presence of axillary branching appears similar to that reported in Callistopliyton and Lyginopteris. Triarch to polyarch adventitious roots, some with secondary tissues, are produced at both nodal and internodal regions. Of the currently recognized monostelic seed fern genera, Microspermopteris is most similar to Heterangium. Information is presented that supports current ideas regarding the evolution of the gymnospermic eustele from protostelic Devonian ancestors.     

FERAXOTHECA GEN. N., A LYGINOPTERID POLLEN ORGAN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The pollen organ Feraxotheca gen. n. is described from Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. The fructifications consist of bilaterally symmetrical synangia composed of a basal pad supporting elongate sporangia that are laterally appressed for the entire length of the sporangial cavities. Sporangial tips extend over the center of the synangium and delimit a small open area, while the bases arise from a parenchymatous cushion that is bounded by short tracheid-like cells. Each synangium is borne on the surface of an expanded pinna tip and is surrounded by a small amount of laminar tissue that envelopes the base of the synangium. Ultimate pinnae are rectangular in transverse section, possess an elliptical vascular bundle surrounded by canals containing a yellow froth-like substance, and have a cortex of elongate cells that radiate from the center of the axis. Sporangia contain small (40–64 μm), radial, trilete spores ornamented by regularly spaced coni or blunt tipped grana. Feraxotheca is compared with the compression genus Crossotheca and some new ideas are advanced concerning the morphology of this compression genus. The obvious differences between Feraxotheca and other lyginopterid pollen organs strongly suggests that the Lyginopteridaceae, as it is currently interpreted, is an unnatural family.     

Paleozoic seed fern pollen organs

Paleozoic pollen organs exhibit numerous morphological forms that have been arranged in categories based on their probable organization. Progymnosperm ancestors are characterized by three dimensional branching systems bearing pairs of terminal sporangia. Early Mississippian examples of seed fern fertile branches appear little modified from the progymnosperms. These pteridosperm microsporangia are nonsynangiate and thin walled with longitudinal dehiscence. By Upper Mississippian time all forms show sporangial clustering into large or small groups, with several taxa exhibiting radially symmetrical synangia. In the Lower Pennsylvanian all pollen organs are synangiate and appear to consist of a uniseriate ring of sporangia that either surround a central hollow, or are bilaterally flattened. Sporangial dehiscence in all forms is longitudinal and toward the center of the synangium. In bilateral synangia with no central hollow, the sporangia either separate laterally or effective dehiscence areas are restricted to the free apical portions of the sporangia. Callistophytacean synangia resemble the lyginopterid type, but are abaxial on laminar foliage. This family is thought to have evolved from the lyginopterids during the Early Pennsylvanian. Middle Pennsylvanian medullosan pollen organs are all radial and may be solitary, aggregated into groups, or fused into a large compound synangium. Several pollen organ types are reinterpreted, and the possible evolutionary relationships among the various Paleozoic pollen organ forms are discussed based on synangial organization, patterns of frond branching, and pollen or prepollen morphology.     

New Pennsylvanian (Moscovian) echinoderms from Xinjiang Uyghur Autonomous Region,western China

The few Pennsylvanian echinoderm faunas reported from the Paleotethys are from China, Japan, and Australia. The Japanese and Chinese faunas contain camerates that are rare in Pennsylvanian faunas worldwide.Genera of one monobathrid camerate, two disparids, five cladids, one flexible, one blastoid, and one archaeocidarid are reported from the late Moscovian uppermost part of the Qijiagou Formation or lowermost part of the Aoertu Formation in the Taoshigo Valley near Taoshuyuan Village, Xinjiang Uyghur Autonomous Region, China. They are here referred to as the Aoertu fauna, which is most closely allied with Pennsylvanian crinoids reported from a slightly older fauna from the Qijiagou Formation in the Taoshigo Valley and with faunas known from Japan and North America. Most taxa are identified to the genus level and left in open nomenclature because of poor or partial preservation. In addition, camerate ossicles and a cup of Synbathocrinus are reported from the early Moscovian part of the Sanquanzi Formation at Yamansu, southeast of the Taoshigo Valley. New taxa described are: Synbathocrinus labrus n. sp., Stellarocrinus qijiagouensis n. sp. and Metaperimestocrinus aoertuensis n. sp.     

THE PENNSYLVANIAN-PERMIAN VEGETATIONAL TRANSITION: A TERRESTRIAL ANALOGUE TO THE ONSHORE-OFFSHORE HYPOTHESIS

An analysis of 68 floras from the Pennsylvanian and Early Permian of Euramerica reveals distinct patterns of environmental distribution. Wetland assemblages are the most commonly encountered floras from the Early and Middle Pennsylvanian. Floras from drier habitats characterize the Permian. Both wetland and dry-site floras occur in the Late Pennsylvanian, but floristic overlap is minimal, which implies strong environmental controls on the distributions of the component species. Drier habitats appear to be the sites of first appearance of orders that become prominent during the Late Permian and Mesozoic. Higher taxa originated in physically heterogeneous, drier habitats, which were geographically marginal throughout most of the Pennsylvanian. They then moved into the lowlands during periods of climatic drying in the Permian, replacing older wetland vegetation. This pattern is analogous to the marine onshore-offshore pattern of origination and migration. The derivation of Mesozoic wetland clades from the Permian dry-lowland vegetation completes the parallel. The similarities of the marine and terrestrial patterns suggest that the combination of evolutionary opportunity, created by physical heterogeneity of the environment, and migrational opportunity, created by changing extrinsic conditions, may be underlying factors that transcend the specifics of organism and environment.     

Late Paleozoic–early Mesozoic continental biostratigraphy — Links to the Standard Global Chronostratigraphic Scale

Nonmarine biostratigraphic/biochronologic schemes have been created for all or parts of the late Carboniferous–Middle Triassic using palynomorphs, megafossil plants, conchostracans, blattoid insects, tetrapod footprints and tetrapod body fossils, and these provide varied temporal resolution. Cross correlation of the nonmarine biochronologies to the Standard Global Chronostratigraphic Scale has been achieved in some parts of the late Carboniferous–Middle Triassic in locations where nonmarine and marine strata are intercalated, the nonmarine strata produce biochronologically significant fossils and the marine strata yield fusulinids, conodonts and/or ammonoids. Other cross correlations have been aided by magnetostratigraphy, chemostratigraphy and a growing database of radioisotopic ages. A synthetic nonmarine biochronology for the late Carboniferous–Middle Triassic based on all available nonmarine index fossils, integrated with the Standard Global Chronostratigraphic Scale, is presented here. The focus is on the nonmarine biostratigraphy/biochronology of blattoid insects, conchostracans, branchiosaurid amphibians, tetrapod footprints and tetrapod body fossils within the biochronological framework of land-vertebrate faunachrons. Correlation to the Standard Global Chronostratigraphic Scale presented here is divided into seven time intervals: Pennsylvanian, Carboniferous–Permian boundary, Cisuralian, Guadalupian, Lopingian, Permian–Triassic boundary and Early to Middle Triassic. The insects, conchostracans and branchiosaurs provide robust nonmarine correlations in the Pennsylvanian–Cisuralian, and the footprints and tetrapod body fossils provide robust correlations of varied precision within the entire Pennsylvanian–Middle Triassic. Radioisotopic ages are currently the strongest basis for cross correlation of the nonmarine biostratigraphy/biochronology to the Standard Global Chronostratigraphic Scale, particularly for the Pennsylvanian–Cisuralian. Chemostratigraphy and magnetostratigraphy thus far provide only limited links of nomarine and marine chronologies. Improvements in the nonmarine-marine correlations of late Paleozoic–Triassic Pangea require better alpha taxonomy and stratigraphic precision for the nonmarine fossil record integrated with more reliable radioisotopic ages and more extensive chemostratigraphic and magnetostratigraphic datasets.     

Arachnids from the Carboniferous of Russia and Ukraine,and the Permian of Kazakhstan

New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carboniferous of Russia and Ukraine and one complete, albeit poorly preserved, specimen from the Permian of Kazakhstan, are described. The spider genus Arthrolycosa is reported from the Late Carboniferous (Late Pennsylvanian: Kasimovian–Gzhelian) of Chunya in the Tunguska Basin of Siberia; it is the first find of a spider outside the Carboniferous tropics. Another fossil assigned to the same genus comes from the Late Carboniferous (Early Pennsylvanian: Bashkirian) of Kamensk–Shakhtinsky in the Donets Basin of Russia; it is probably the oldest fossil spider known. A thelyphonid (whip scorpion) carapace is described from the Late Carboniferous (Late Pennsylvanian: Kasimovian) of the adjacent Lugansk Province of the Donets Basin of Ukraine.