An ophiuroid trackway from the Lower Devonian Hunsrück Slate, Germany

A new trace fossil, Arcichnus saltatus , from the Lower Devonian Hunsrück Slate, Germany, is interpreted as the trackway of a protasterid ophiuroid, Taeniaster. The trackway consists of a series of horseshoe-shaped impressions, produced by a pair of arms, a feature characteristic of some ophiuroid trackways. This new trace fossil demonstrates that protasterids lived both infaunally and epifaunally.     

Annelids from the Lower Devonian Hunsrück Slate (Lower Emsian, Rhenish Massif, Germany)

Abstract:  Four new genera and species of annelid, Hunsrueckochaeta hohensteini , Ewaldips feyi , Crocancistrius lutzi and Scopyrites magnus , are described from the Hunsrück Slate (Lower Emsian) of Germany, as well as new material of Bundenbachochaeta eschenbachensis   Bartels and Blind, 1995 . The specimens preserve details of the appendages and other aspects of the morphology as a result of pyritization. A phylogenetic analysis using the morphological data matrix of Rouse and Fauchald places four of the five genera basal to the Aciculata; the fifth Ewaldips falls within the Scolecida. The diversity of annelids represented in the Hunsrück Slate is similar to that in the other major Palaeozoic Konservat-Lagerstätten that yield them, including the Cambrian Burgess Shale and the Late Carboniferous Mazon Creek biota.     

A NEW SEA SPIDER (ARTHROPODA: PYCNOGONIDA) WITH A FLAGELLIFORM TELSON FROM THE LOWER DEVONIAN HUNSRÜCK SLATE, GERMANY

Abstract:  A new Lower Devonian sea spider (Arthropoda: Pycnogonida) from the Hunsrück Slate, Germany, is described as Flagellopantopus blocki gen. et sp. nov. This is only the sixth fossil pycnogonid species to be described. Its most remarkable and unique aspect is the long, flagelliform telson. Although our fossil apparently lacks chelifores (an apomorphy), the retained telson and the segmented trunk end behind the last pair of legs resolve F. blocki to a fairly basal position in the pycnogonid stem lineage. It probably lies between Palaeoisopus problematicus Broili, which has a lanceolate telson and the most trunk segments of any sea spider, and all other Silurian–Recent Pycnogonida. Our new material shows that at least two fossil pycnogonids retained a telson, albeit with very different morphologies, and further supports the idea that a greater diversity of body plans existed among the Palaeozoic pycnogonid taxa.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

Ichnological evidence on the behaviour of mitrates: two trails associated with the Devonian mitrate Rhenocystis

The mitrates are controversial marine Palaeozoic fossils, assigned either to the chordates or to the echinoderms. This paper describes two trails associated with the mitrate Rhenocystis from the Lower Devonian Hunsrück Slate, Bundenbach, Germany. They indicate that, just before death, the animals were moving tail-first with the flat dorsal surface of the head upward. This behaviour is consistent with the chordate theory of mitrates, rather than with either of the two current echinoderm interpretations.     

Preservation of tube feet in an ophiuroid (Echinodermata) from the Lower Devonian Hunsrück Slate of Germany and a redescription of<Emphasis Type="Italic">Bundenbachia beneckei</Emphasis> and<Emphasis Type="Italic">Palaeophiomyxa grandis</Emphasis>

The preservation of non-mineralized tissues in the fossil record is extremely rare. The Lower Devonian Hunsrück Slate of Germany has long been known for the preservation of non-mineralized tissues in pyrite but whether or not these remnants represent true soft tissues has been questioned. This is especially true for struetures visible only on radiographs that are too delicate for excavation by traditional methods. Here we report the discovery of well-preserved pyritized tube feet in six fully prepared specimens of the protasterid brittle starBundenbachia beneckei from the Hunsrück Slate. This discovery represents the first report of fossilized ophiuroid tube feet in the fossil record. The successful excavation of the delicate tube feet was made possible by improved airbrasive techniques developed by German fossil collectors. The relatively large size of the fossil tube feet inBunden-bachia beneckei is consistent with earlier inferences on size based on the presence of large podial basins. Protasterid ophiuroids lack the specialized arm musculature and articulations that provide increased flexibility and strength to the arms of modern ophiuroids with typically reduced tube feet. How-ever, tube foot form and perhaps function inBundenbachia might have been similar to those of living asteroids in which large tube feet are used primarily for locomotion and food-manipulation thus compensating for a lack of specialized arm musculature and articulation. Hence, feeding and life mode of protasterid ophiuroids was not necessarily limited to sedentary, infaunal microphagy as traditionally suggested. Two Hunsrück protasterid ophiuroids,Bundenbachia benecki andPalaeophiomyxa grandis are redescribed and compared.      

ICHNOLOGICAL INSIGHTS INTO MITRATE PALAEOBIOLOGY

Abstract:  Mitrates are a controversial group of extinct deuterostomes; there is little agreement over their affinities, functional morphology or even the orientation of their upper and lower surfaces. Four slabs of slate from the Lower Devonian Hunsrück Slate (Bundenbach, Germany) are here described, showing trace fossils ( Vadichnites transversus igen. et isp. nov.) associated with the mitrate Rhenocystis latipedunculata . These new findings clearly demonstrate that the mitrate appendage was used in locomotion and that this movement took place appendage-first. Such a functional interpretation suggests that mitrates were oriented with the flat body surface upwards in life and argues against a phylogenetic position in the echinoderm crown-group.     

First record of eurypterids (Chelicerata,Eurypterida) from the Lower Devonian (Lower Emsian) Hunsrück Slate (SW Germany)

For the first time, fossils of unquestionable eurypterid origin are documented from the renowned Lower Devonian Hunsrück Slate Lagerstätte in southwestern Germany. The gnathobase of a coxal plate and a body segment can be attributed to pterygotid eurypterids, probably to Jaekelopterus rhenaniae. These body parts from the Hunsrück Slate once more confirm the extraordinary large size that could be attained by this Rhenish eurypterid. Based on the usual occurrence in marine–terrestrial transitional facies elsewhere in the Rhenish Devonian, rareness in the Hunsrück Slate, and poor preservation, the Hunsrück Slate pterygotid remains are most probably allochthonous, and eurypterids are considered nongenuine elements of the ‘Hunsrück Slate biota.     

Devonohexapodus bocksbergensis is a synonym of Wingertshellicus backesi (Euarthropoda) – no evidence for marine hexapods living in the Devonian Hunsrück Sea

The Devonian Hunsrück Slate fossil Devonohexapodus bocksbergensis Haas, Waloszek & Hartenberger, 2003 has been interpreted as a stem-lineage representative of the Hexapoda, implying their marine origin and independent terrestrialisation within the ‘Atelocerata’. Devonohexapodus bocksbergensis was based on a single specimen embedded in a lateral position. Reinvestigation of that holotype and of all known specimens of the Hunsrück Slate arthropod Wingertshellicus backesi Briggs & Bartels, 2001 demonstrates that all this material represents a single species. The latter is redescribed, its taxonomic diagnosis is emended, and the name Devonohexapodus bocksbergensis is treated as a junior synonym of Wingertshellicus backesi. The phylogenetic position of W. backesi neither is that of a stem-lineage representative of Hexapoda, nor does it fall within the crown group Mandibulata. The Hunsrück Slate provides no evidence of an independent terrestrialisation within the ‘Atelocerata’ or of a marine origin of the Hexapoda.     

A new specimen ofWeinbergina opitzi (Chelicerata: Xiphosura) from the Lower Devonian Hunsriick Slate, Germany

A new specimen of the synziphosurine arthropodWeinbergina opitzi is described from the Lower Devonian (Lower Emsian) Hunsrück Slate of Germany (Rhenish Slate Mountains). It is the smallest and only the fifth specimen of this taxon to be described and is preserved in ventral aspect with exceptional preservation of prosomal and opisthosomal appendages. This specimen confirms the presence of a seventh appendage, similar in morphology to the preceding prosomal appendages, associated with opisthosomal segment one. In addition, at least three opisthosomal plates fringed with teeth are confirmed. Correlation of prosomal appendage podomeres betweenWeinbergina and selected chelicerate taxa shows that appendage structure is most similar to eurypterid appendages III–IV and Araneae appendages III–VI. This is in contrast to modern horseshoe crabs which have fewer podomeres in appendages II–V due to an undifferentiated tibiotarsus.      

Re-evaluation of the Devonian family Helianthasteridae Gregory, 1899 (Asteroidea: Echinodermata)

Because of their taxonomic and morphologic diversity, the asteroids of the Lower Devonian Hunsrück Slate of Germany are important to both an understanding of the history of the class Asteroidea and to the interpretation of community evolution during the Paleozoic. Helianthaster Roemer, 1863, a large multiarmed Hunsrück asteroid, is redescribed. The Helianthasteridae Gregory is restricted to Helianthaster and Arkonaster Kesling, 1982 (Middle Devonian, Canada); Lepidasterella Schuchert is similar to the other two genera but known specimens are of poor quality and as a result the status of the genus is uncertain. Helianthaster is noteworthy in part because its size, multiarmed state, and presence of pedicellariae suggest certain crown-group genera, yet aspects of the arrangement of the ambulacral column are characteristic of the Paleozoic asteroid evolutionary grade. Neither Helianthaster nor other Hunsrück asteroids appear closely linked to the crown group; instead, functional patterns apparently re-emerged through time, although identification of specific behavior of ancient asteroids is difficult to impossible.      

Machaeridian locomotion

The discovery that machaeridians (class Machaeridia Withers, 1926) are annelids allows their mode of locomotion to be interpreted in the context of the body plan of this phylum. The Plumulitidae were errant epibenthic forms, moving with parapodia. The body of Turrilepadidae and Lepidocoleidae, however, was enclosed largely within the mineralized plates that make up the skeleton. Articulated specimens indicate that these machaeridians were able to burrow like other annelids using peristaltic locomotion. A lepidocoleid specimen indicates that multiple waves of shortened and contracted regions moved over the body. This is in contrast to the mode of locomotion in earthworms and most polychaetes, but similar to peristaltic progression in Polyphysia (Scalibregmidae). Either the rugose sculpture (turrilepadids) and/or the margins of the overlapping shell plates functioned as a burrowing sculpture, allowing forward movement but preventing backwards slipping. A trace from the Devonian Hunsrück Slate associated with a lepidocoleid indicates that considerable flexing of the skeleton was possible, but this is an escape trace and does not represent normal locomotion. Features of the skeleton of machaeridians are convergent on those of molluscs where the shells likewise function in protection and burrowing.     

Captopodus poschmanni gen. et sp. nov. a new stem-group arthropod from the Lower Devonian Hunsrück Slate (Germany)

A new arthropod from the Lower Devonian Hunsrück Slate is described on the basis of four specimens. The body of Captopodus poschmanni comprises a head, a trunk with an anal portion. The high number of trunk appendages (≥66 segments) is unusual. The function of one pair of cupola-like structures of the head shield is unclear. The presence of large grasping appendages in the head superficially resembles the ‘short great appendages’ of other euarthropods and grasping appendages of thylacocephalans. The phylogenetic position of the arthropod cannot be determined in detail, though several morphological aspects indicate a phylogenetic position as a stem lineage representative of the Euarthropoda, the morphology of the trunk appendages seem to indicate a more advanced phylogenetic position. This new taxon underlines the exceptional diversity of arthropods within the Hunsrück Slate in comparison to other Devonian fossil sites and highlights the significance of the Hunsrück Slate for the evolution of early arthropods.     

Presumed postlarval pentacrinoids from the Lower Devonian Hunsrück Slate,Germany

Several tiny crinoids with crowns as small as 1 mm, or less, in width are newly recognized from the Hunsrück Slate of southwestern Germany. The presence of erect arms above an amorphous calyx in some specimens can be inferred. Based on comparison with the size and gross morphology of developmental stages in living crinoids, these tiny Hunsrück crinoids are judged to be at an early postlarval stage that is analogous to the pentacrinoid stage just after development from the stalked, but armless, smaller cystidean larval stage found in both living comatulids and isocrinids. Some of these tiny crinoids have a stalk up to 4 mm long attached to a now pyritized former substrate. Their clustered occurrence suggests gregarious settlement of larvae. Taxonomic identification of these presumed pentacrinoids is not possible, even to the sub‐class level, although they are preserved with larger juveniles of the cladids Propoteriocrinus and Lasiocrinus. These larger juveniles exhibit 3‐D pyritized calcite plates, whereas the probable pentacrinoids appear to be preserved as flattened, micro‐crystalline pyritized dermal tissues that enclosed lightly calcified, porous ossicles. The pentacrinoids were likely buried within weeks or months of hatching, based on developmental stages in similar‐sized living crinoids. These tiny crinoids, presumably pentacrinoids, are a further example of the extraordinarily detailed preservation of delicate tissues in pyrite from the Hunsrück Slate. They are most likely the pentacrinoid stage from one or more of the crinoid taxa (30 genera) present in the Hunsrück Slate. Assuming these are not microcrinoids, they are the first report of pentacrinoids from the fossil record and document that a Palaeozoic sister group to modern crinoids had similar developmental stages.     

Redescription and palaeobiology of Palaeoscorpius devonicus Lehmann, 1944 from the Lower Devonian Hunsrück Slate of Germany

Abstract: Palaeoscorpius devonicus Lehmann, 1944 is known from only a single specimen, found in the Eschenbach Pit near Bundenbach in the Lower Devonian Hunsrück Slate of Germany. It is a key fossil, having been interpreted both as the most basal member of the Scorpiones and as one of the order’s most likely candidates for an aquatic mode of life. Prepared both ventrally and dorsally, some aspects of its morphology remain problematic. Here, with the aid of new techniques, including computed tomography, we present a re‐investigation of this scorpion’s anatomy and a new reconstruction, with a particular focus on the species’ original habitat. On the basis of the environmental interpretation of the Hunsrück Slate and the completeness of the specimen, previous authors concluded that P. devonicus was marine, but none offered convincing morphological evidence. Recent studies of the deposit’s environment suggest that the Hunsrück Sea was part of an intrashelf basin, relatively close to the coastline, and fossils of land plants show that terrestrial wash‐in occasionally occurred. Our revised interpretation of the fossil’s morphology demonstrates that the scorpion was most probably terrestrial. Internal mesosomal organs are interpreted as book lungs, but other terrestrial adaptations are lacking. The absence of both coxapophyses and gnathobases makes determining the scorpion’s feeding mechanism difficult. Interpreting the scorpion’s character states within a phylogenetic framework, especially the possible presence of book lungs, implies either that the plesiomorphic position of P. devonicus is no longer supported or that the development of book lungs had already taken place early in the scorpion lineage.     

An atypical fauna in the Lower Devonian Hunsrück Slate of Germany

The Hunsrück Slate is a world renowned conservation lagerstätte, stretching SW-NE in a narrow band between the villages of Bundenbach and Gemünden, Hunsrück region, Germany. A great variety of complete Lower Devonian fossils is preserved with their soft parts pyritised due to rapid burial by sediment. The fossils of the Hunsrück Slate outside the narrow band are scarcely known. They represent the normal Situation: quiet low energy Sedimentation. This paper describes this overall “Rhenish” neritic fauna under the heading “atypical”, in contrast to the famous fossils of the conservation lagerstätte. The fauna described here lacks starfishes, mitrates and chelicerates. The paper begins with an overview of the recent relevant literature (stratigraphic position of the Hunsrück Slate, palaeoecology). The slate fossils were collected in two quarries WNW and NE of Bundenbach (Lingenbach and Karschheck, respectively) and most closely resemble that of the Wisper Valley in the Taunus region. Their geologic age is Lower Devonian, early Emsian, Ulmen Substage. Special interest is given to Community structures, rugose corals, bivalves, gastropods, trilobites, conulariids, brachiopods and crinoids. In the systematic part, the crinoidOrthocrinus simplex is redescribed. Also, two new species are introduced: the rugose coralVolgerophyllum karschheckensis n. gen., n. sp., and the crinoidAcanthocrinus spinosus n. sp.     

A pyritized lepidocoleid machaeridian (Annelida) from the Lower Devonian Hunsrück Slate,Germany

A machaeridian, Lepidocoleus hohensteini sp. nov., is described from the Hunsrück Slate (Lower Emsian) of Germany. The available material includes a unique example preserving evidence of the soft tissues, only the second machaeridian specimen to do so and the first lepidocoleid. This specimen shows that the plates are attached to alternate segments in the trunk. The morphology is consistent with an annelid affinity of the Lepidocoleidae and confirms the unity of the Machaeridia. This discovery adds an important group to the known diversity of this famous late Palaeozoic marine Konservat-Lagerstätte.     

New arthropods from the Lower Devonian Hunsru¨ck Slate (Lower Emsian, Rhenish Massif, western Germany)

Four new genera and species of arthropod, Cambronatus brasseli Wingertshellicus backesi Eschenbachiellus wuttkensis and Magnoculus blindi , are described from the Hunsru¨ck Slate (Lower Emsian) of Germany. All four occur in the Wingertshell Member in the vicinity of Bundenbach. They preserve remarkable details of the ventral morphology, including the appendages, as a result of pyritization. In each case the body consists of just two tagmata, a cephalon and a large number of similar trunk somites. Both Cambronatus Wingertshellicus have fluke-like appendages making up a tail fan. The affinities of these arthropods lie with the Crustacea, apart from Magnoculus , which is an arachnomorph, but they do not fall within those clades with modern representatives. They show that morphologies other than those represented by trilobites and modern arthropod groups persisted long after the Cambrian, at least in muddy bottom marine settings.     

The acanthodian<Emphasis Type="Italic">Machaeracanthus</Emphasis> from the Lower Devonian Hunsrück Slate of the Hunsrück region (Germany)

Two recently collected slabs from the Lower Devonian Hunsrück Slate of Bundenbach, Hunsrück region, Germany, with spines of the acanthodianMachaeracanthus hunsrueckianum n. sp. are described. Both are associations of large and small spines and are the first to show groupings of symmetrical pairs; the spines are not homologous with those of other acanthodians. A pair of small spines ofMachaeracanthus peracutus Newberry, 1857 from the Karschheck quarry near Oberkirn, Hunsrück region, Germany, is articulated with the pectoral girdle and is the first such complex to be described. The only spines whichMachaeracanthus appears to have had were a pair of large and small pectoral spines on each side of the body. These spines could have helped to prevent the fish from sinking into the mud while resting on the sea floor.