Royal H. Mapes

Elemental mobility in phosphatic shales during concretion growth and implications for provenance analysis

Shale-normalized rare-earth element (REE) patterns record variable redox conditions in coprolitic phosphorite concretions and their host shales from the Pennsylvanian-age cyclothems in the U.S. midwestern states of Oklahoma, Kansas, Nebraska, Iowa, and Ohio as well as the Mississippian-age Fayetteville Shale of Arkansas. Shale-normalized enrichments in middle REE (MREE) developed as the phosphatic concretions lithified under reducing conditions in pore waters of organic-rich muds. Depletions in MREE (Sm–Ho) in host shales correspond to the MREE enrichments in some nearby concretions. Rare and weak negative cerium depletions in both reworked and unreworked phosphorite concretions suggest exposure to oxygenated waters. Positive europium anomalies in thick phosphorite coatings on coprolitic phosphorite concretion cores suggest extreme reduction during lithification of the phosphorite coating. Flat REE patterns in both host shales and phosphorite concretions probably reflect detrital influence. Phosphate diagenesis can affect the trace element chemistry of black shale enough to alter provenance signals. Overall enrichment in REE in phosphorite relative to host shale suggests movement of REE as does the MREE-enriched phosphorite near the MREE-depleted host shale. Diagenetically mobile REE can affect La–Th–Sc relationships used in provenance analysis. The extent of the diagenetic modification of La–Th–Sc distribution is related to the amount of P2O5 present. Typical La–Th–Sc distribution in host shale results when P2O5 values are below 0.5%. High La abundance substantially skews La–Th–Sc distribution when P2O5 is above 5.0%, but when P2O5 is between 0.5% and 5.0% the extent of the effect is unclear. Oxides of aluminum and other major elements may be useful in recording detrital influence and proximity to the shoreline in phosphorite concretions. Most of the concretions nearest to shoreline sources of detrital input have both flat REE patterns and elevated values of Al2O3/(Al2O3+Fe2O3).

The youngest carpoid: occurrence, affinities, and life mode of a Pennsylvanian (Morrowan) mitrate from Oklahoma

Abundant, well-preserved specimens of a new peltocystidan mitrate carpoid, Jaekelocarpus oklahomensis n. gen. and sp., have been discovered in the Pennsylvanian (Morrowan Series) Gene Autry Shale Member, Golf Course Formation, in southern Oklahoma. The new carpoid postdates the youngest previously known carpoids (Middle Devonian) by about 65 million years. It is characterized by a small (to 4 mm in length) globose theca composed of six plates: two large adaulacophorals, two large marginals, and two relatively small plates that frame the main thecal orifice. One of the plates at the orifice bears a single stout spine. The theca and plates that comprise the theca display a high degree of bilateral symmetry. In contrast, the styloid has an asymmetrical array of spines and blades. We hypothesize that most mitrates, including this one, were adapted to an infaunal life mode. The streamlined, generally symmetrical body shape is an adaptation to moving on or through the substrate. Spines and blades on the lower surface of the styloid and proximal aulacophore are inferred to be adaptations for gripping the substrate as the animal pulled itself backwards (aulacophore first). The main thecal orifice opposite the aulacophore was the site of both mouth and anus. Evidence for a sluggish, epifaunal life mode in the cornute carpoids includes the typically flattened, asymmetrical body, the presence of downward projecting spines and knobs on the marginal thecal plates of many species, and the specialized thecal pores and slits that faced away from the substrate.

Late Paleozoic conifers of North America: structure, diversity and occurrences

Abstract Late Paleozoic conifer communities originated in moisture-stressed environments along the paleoequator on the Laurussian continent. By Late Permian time, northward movement of the continent displaced these communities toward subequatorial regions. North American deposits provide the most abundant and best preserved pre-Permian conifer fossils. They represent a wide spectrum of preservational modes including compression/impression, mold/cast, and cellular permineralization by carbonates, phosphates, iron sulfides, iron oxides and iron hydroxides. Some modes are associated with distinct biotic assemblages, and to some extent each can be correlated with certain depositional environments and taphonomic histories. Exceptional floras containing extensive conifer remains occur in the Rocky Mountains, the Midcontinent, and the Appalachian regions. These include: estuarine carbonates at Hamilton and Pomona, Kansas; transitional terrestrial to marine clastics at Kinney Brick Quarry, New Mexico, Garnett, Kansas and the 7–11 Mine in northeastern Ohio; and offshore marine shales in Kansas, Oklahoma and Texas that were deposited in dysoxic environments. Most specimens have been assigned to the form genus Walchia , but exceptionally well preserved specimens conform to species of Emporia (Emporiaceae) and to one or more undescribed species that may represent additional genera and/or families. These species add significantly to the growing diversity of Paleozoic conifers, and provide valuable data for testing hypotheses of conifer and coniferophyte relationships.

Larval and juvenile gastropods from a Carboniferous black shale: palaeoecology and implications for the evolution of the Gastropoda

A horizon in the late Visean Ruddle Shale from Arkansas contains the oldest well-preserved gastropod protoconchs known from the Americas. The gastropod fauna consists of a diverse larval shell assemblage and a low diversity assemblage of juvenile gastropods that probably had a benthic life habit. Gastropod larval shells are always isolated, i.e. the gastropods did not complete their life cycle (no metamorphosis) and were unable to become benthic. This was caused by unfavorable environmental conditions on the soft muddy bottom that was probably due to anaerobic to exaerobic conditions. The absence or scarcity of bioturbation caused by invertebrate detritus or sediment feeders in both shale and concretions (formed before compaction) favored preservation of the delicate larval shells. The lack or scarcity of infauna and bioturbation as well as the low diversity of the presumed benthos supports an interpretation of a quasi-anaerobic to exaerobic benthic environment. The superbly preserved larval shells demonstrate that there are more caenogastropod clades present in the late Palaeozoic than suggested previously. Some larval shell types have an openly coiled first whorl followed by a planktotrophic larval shell; openly coiled initial whorls are unknown from modern caenogastropods. The vetigastropods have a smooth protoconch of two whorls clearly demarked from the following whorls - a pattern unknown in modern vetigastropods which have a protoconch of less than one whorl and build no larval shell during their planktonic stage. This could indicate a link between Palaeozoic vetigastropods and the caenogastropods.

IDENTITY AND PHYLOGENY OF THE LATE PALEOZOIC SUBULITOIDEA (GASTROPODA)

Abstract The Subulitoidea have long been an enigmatic group of Paleozoic gastropods and share many characters of post-Paleozoic clades. Newly described protoconchs from several late Paleozoic subulitoid species have been employed in a phylogenetic analysis of the group. Late Paleozoic representatives, the Soleniscidae, are caenogastropods with an unornamented orthostrophic larval shell. The Meekospiridae have a smooth blunt protoconch of about two whorls. In contrast to previous interpretations, this protoconch is not heterostrophic or heterostrophy is not obvious. Therefore, a placement of the Meekospiridae with the genus Girtyspira in the Opisthobranchia is to be treated with caution. The new Ianthinopsis-like genus Imogloba has a gobular subulitid-like teleoconch but its protoconch consists of a smooth first whorl which is loosely coiled and a larval shell with a characteristic non collabral ornament. Therefore, a close relationship between Soleniscidae and Imogloba is unlikely and the new family Imoglobidae is proposed. We found no clearly heterostrophic groups within the late Paleozoic subulitoid gastropods. The phylogenetic analysis of all subulitoid genera with known protoconchs provides little support for the monophyly of the Subulitoidea, particularly for a close relation between the Meekospiridae and the Soleniscidae. However, the Soleniscidae and Meekospiridae are probably monophyletic groups. Most genera are restricted to the Paleozoic, although several Mesozoic genera may hold descendants of Paleozoic Subulitoidea. Finally, a hypothesized link between Subulitoidea and Neogastropoda based on the presence of an anterior notch in both groups is unlikely.

Cephalopods present and past : new insights and fresh perspectives

Part I: Phylogeny and Systematics 1. Phylogenetic Practices among Scholars of Fossil Cephalopods, with Special Reference to Cladistics Pascal Neige, Isabelle Rouget, and Sebastien Moyne 2. Patterns of Embryonic Development in Early to Middle Devonian Ammonoids Susan M. Klofak, Neil H. Landman, and Royal H. Mapes 3. Mode of Life of the Frasnian (Late Devonian) Ammonoid Manticoceras from Coumiac (Montagne Noire, France) Dieter Korn and Christian Klug 4. GONIAT - The Current State of the Paleontological Database System on Paleozoic Ammonoids Jurgen Kullmann 5. Ornamental polymorphism in Placenticeras kaffrarium (Ammonoidea Upper Cretaceous of India): Evolutionary implications Tapas K. Gangopadhyay and Subhendu Bardhan 6. A Late Carboniferous Coleoid Cephalopod from the Mazon Creek Lagerstatte (USA), with a Radula, Arm Hooks, Mantle Tissues, and Ink Larisa A. Doguzhaeva, Royal H. Mapes, and Harry Mutvei 7. On the Species Status of Spirula spirula (Linne, 1758) (Cephalopoda): A New Approach Based on Divergence of Amino Acid Sequences between the Canaries and New Caledonia Kerstin Warnke Part II: Morphology of Soft and Hard Tissues 8. Understanding Ammonoid Sutures: New Insight into the Dynamic Evolution of Paleozoic Suture Morphology Emily G. Allen 9. Cameral Membranes in Carboniferous and Permian Goniatites: Description and Relationship to Pseudosutures Kristin Polizzotto, Neil H. Landman, and Royal H. Mapes 10. Soft-Tissue Attachment of Middle Triassic Ceratitida from Germany Christian Klug, Michael Montenari, Hartmut Schulz, and Max Urlichs 11. The Preservation of Body Tissues, Shell, and Mandibles in the Ceratitid AmmonoidAustrotrachyceras (Late Triassic), Austria Larisa A. Doguzhaeva, Royal H. Mapes, Herbert Summesberger, and Harry Mutvei 12. Connecting Ring Ultrastructure in the Jurassic Ammonoid Quenstedtoceras with Discussion on Mode of Life of Ammonoids Harry Mutvei and Elena Dunca 13. Jaws and Radula of Baculites from the Upper Cretaceous (Campanian) of North America Neil H. Landman, Neal L. Larson, and William A. Cobban 14 Ultrastructure Analyses on the Conotheca of the Genus Belemnotheutis (Belemnitida: Coleoidea) Dirk Fuchs, Helmut Keupp, Vasilij Mitta, and Theo Engeser Part III: Biogeography, Biostratigraphy, Ecology, and Taphonomy 15. New Data on the Clymeniid Faunas of the Urals and Kazakhstan Svetlana Nikolaeva 16. Deformities in the Late Callovian (Late Middle Jurassic) Ammonite Fauna from Saratov, Russia Neal L. Larson 17. Biogeography of Kutch Ammonites During the Latest Jurassic (Tithonian) and a Global Paleobiogeographic Overview Subhendu Bardhan, Sabyasachi Shome, and Pinaki Roy 18: Ammonite Touch Marks in Upper Cretaceous (Cenomanian-Santonian) Deposits of the Western Interior Seaway Neil H. Landman and William A. Cobban 19: Some Data on the Distribution and Biology of the Boreal Clubhook Squid Moroteuthis robusta (Verrill, 1876) (Onychoteuthidae, Teuthida) in the Northwest Pacific Alexei M. Orlov 20: Habitat Ecology of Enteroctopus dofleini from Middens and Live Prey Surveys in Prince William Sound, Alaska D. Scheel, A. Lauster, and T.L.S. Vincent

Medullosa steinii sp. nov., a seed fern vine from the Upper Mississippian

Three specimens of a medullosan seed fern stem with attached leaf bases, from the Chesterian Series of Arkansas, USA, are the earliest unequivocal occurrence of the genus Medullosa and form the basis for a new species Medullosa steinii. The stems have numerous features characteristic of vines, including stem diameters of only 2–3 cm in diameter, numerous wide, plate-like rays, relatively wide tracheids, the presence of cortical spines and variable but numerous stelar segments. In M. steinii vascular segment numbers vary from two to eight. Primary xylem bundles consist of one to several protoxylem poles with metaxylem and abundant parenchyma; each bundle is surrounded by secondary vascular tissue. Leaf traces typically diverge from a vascular segment as pairs, with the resulting bundles separating from each other at an angle of 90–100°. Paired leaf traces are separated from each other by a wedge of secondary vascular tissue, thus producing a distinctive mode of leaf trace production for the new species. Leaf traces divide repeatedly to produce the Myeloxylon rachis base configuration at the margin of the stem cortex. Rachis bases are decurrent and are separated from one another by a discontinuous row of sclerotic bundles and from the stem by a prominent periderm. The discovery of this plant demonstrates that scrambling and climbing seed ferns were part of some plant communities by Mississippian time.

Predation on the Pennsylvanian ammonoid Gonioloboceras and its implications for allochthonous vs. autochthonous accumulations of goniatites and other ammonoids

Damage displayed by 18 of 954 relatively mature (> 35 mm diameter) specimens of Gonioloboceras goniolobum (Meek) from North American Midcontinent Upper Carboniferous sediments is interpreted to be bite marks caused by condrichthyans and other fish, particularly the symmoriid shark Symmorium reniforme. It is likely that these and other predators regularly preyed on Gonioloboceras goniolobum because all of the analyzed specimens of Gonioloboceras exhibit some evidence of unrepaired damage, including broached camerae and missing body chambers. Two new characteristics utilized with caution to detect predatory events on ammonoids are raised oval or circular pedestals on internal molds and the breakage and termination of septa in conjunction with a broken external shell. When sufficient damage to the Gonioloboceras conchs occurred during predatory attacks, the broached camerae flooded, the conch became negatively buoyant, and it sank in the vicinity of the attack. When preserved, these conchs form a variable component of an autochthonous fossil accumulation that can include numerous other cephalopod taxa, which also were probably subjected to predatory attacks. Other, but not all, ammonoid accumulations in the fossil record form similar autochthonous deposits, or even autochthonous portions of mixed deposits.

Anatomically preserved vojnovskyalean seed plants in Upper Pennsylvanian (Stephanian) marine shales of North America

ABSTRAcr-Upper Pennsylvanian dysoxic marine shales of midcontinent North America yield permineralized remains of apparently extrabasinal vegetation. A large percentage of the plants are surprisingly unlike the well known swamp, fluvial and lacustrian floras of the late Paleozoic paleotropics, revealing numerous aspects of the morphology, anatomy and reproductive biology of plants that may have been ancestral to the dominant taxa of the Mesozoic. Included among the assemblages are ovulate coniferophyte remains that demonstrate the occurrence of vojnovskyalean seed plants in equatorial Euramerica. Specimens consist of simple ovulate cones that are more-or-less clustered along eustelic stems in the axils of helically arranged, strap-shaped leaf bases, and are described as Sergeia neuburgii new genus and species. Individual cones are up to approximately 2 cm long and 1.5 cm in maximum diameter, with helically arranged scales and sporophylls that diverge from a eustelic axis. Scales occur in the basal region of the cone, and are laminar and pointed. Sporophylls are borne distally. One specimen shows about 45 sporophylls, each of which terminates as one erect ovule. Most of the other cones have abraded apices, and terminate in relatively terete foliar appendages that are interpreted to be sporophyll bases. Ovules are flattened and winged, approaching 1800 rotational symmetry. Integument histology, vascular tissue distribution and pollen chamber structure are similar to those of cordaiteans and callistophytalean seed ferns. Sergeia adds to the number of late Paleozoic conifer-like plants that do not conform to the Pinopsida as traditionally circumscribed, and poses additional questions to assumptions of monophyly for coniferophytes and for conifers sensu lato.

Pityostrobus milleri sp. nov., a pinaceous cone from the lower Cretaceous (Aptian) of southwestern Russia.

A new species of permineralized Pityostrobus cone, Pityostrobus milleri Falder et al., sp. nov. has been discovered in a calcareous nodule from shallow marine sediments of Early Cretaceous (latest Aptian; Clansenian) age in the Caucasus Mountains in southwestern Russia. Nodules of this type also contain wood fragments, gastropods, bivalves, and an extremely rich assemblage of well preserved cephalopods. Russian spore and pollen analyses indicate that deposition in this region may have occurred during a temperate climate regime. The cone is cylindrical, about 2.5 cm in diameter and individual scales contain up to 17 resin canals and 18 vascular strands. Although the fossil displays many features of modern Pinus cones, including inflated scale apexes, resin canals abaxial to vascular tissue in scale base, and scale strands curved on the adaxial side, it cannot be assigned to Pinus because the bract and scale trace are not united at their origin. Tunneled borings in the cortex of the cone axis are tightly packed with frass, and resemble tunnels of modern cone boring beetles such as Conophthorus. This cone further documents the worldwide diversity of pinaceous conifers during the Cretaceous, and demonstrates a well developed syndrome of insect herbivory in the Pinaceae by the late Mesozoic.