Clionolithes alcicornis ( Vogel, Golubic & Brett, 1987 ) Wisshak, 2017

Wisshak, Max, 2017, Taming an ichnotaxonomical Pandora’s box: revision of dendritic and rosetted microborings (ichnofamily: Dendrinidae), European Journal of Taxonomy 390, pp. 1-99 : 32-34

Clionolithes alcicornis ( Vogel, Golubic & Brett, 1987) comb. nov.

Fig. 9

Ramodendrina alcicornis Vogel et al., 1987: 272 , fig. 6.

Clionolithes bullahirsuta Plewes, 1996: 178 , pl. 27, figs 3–7, pl. 28, figs 1–8 .

Ramodendrina – Vogel 1987: fig. 5.

Original diagnosis

Rosette-shaped repeatedly branched systems up to 2 mm (1.41 ± 0.34 [16]) in diameter. Branches diverge from the main trunk spreading radially, curved in a characteristic clawlike fashion at angles ranging from 40° to 115° (72 ± 17 [50]). The main trunk is 75–190 µm (142 ± 40 [7]) wide. The primary branches range from 53 to 136 µm (95 ± 19 [50]), the secondary branches from 31 to 66 µm (51 ± 11 [50]), and the tertiary branches from 12 to 45 µm (26 ± 9 [37]) in diameter. They have smooth and rounded surfaces, and are circular in cross section. The branches taper toward pointed tips. There is a significant decrease in diameter from primary to secondary and tertiary (distal) branches, and the diameter between individual branching points remains constant or decreases. The branching varies in density, without anastomoses. The main aperture to the surface was not detected.

Emended diagnosis

Branches diverge from the flat and elongated main trunk, spreading radially and in one plane, often in straight lines, ramifying in varying density at angles between 40° and 115° without forming anastomoses, tapering toward pointed tips. Often with a distinctive cuspate texture and fine filaments scattered over the entire surface, but most numerous on the edges and sides of the branches.

Original description

n/a, but see detailed original diagnosis.

Supplementary description

When erecting the junior synonym (and nomen nudum) C. bullahirsuta, Plewes (1996) made the following additional observations, particularly concerning the surface texture that is only poorly preserved in the Devonian holotype: “The overall oval shape and the characteristic texture of the trace are constant attributes (although the presence of the texture is dependent on the preservation and the quality of the cast). Whole rosette up to 3 mm in diameter. Even within the holotype the branches vary in their morphology, and frequency. However, there are recurring features: branches may have a sweeping shape and taper towards the pointed, but not filamentous, ends. Although branches do anastomose, sometimes they do not fuse and appear on the casts as still separate. The width of branches varies between 40 µm and 170 µm with a mean of 93 µm (n = 16, SD = 48.52). Some specimens show a curious loop on one side. The pseudochip sculpture is also highly variable and measured across the widest point can be up to 50 µm or as small as 7 µm the average is 18 µm (n = 19, SD = 9.99). The hairs also vary in length, although this is probably due more to the preservation of the substrate than variation between individuals. The width of the hairs is more constant, with a mean of 2.5 µm (n = 15, SD = 0.92).”

Type material, locality and horizon

The holotype ( Fig. 9 A–B) was epoxy cast in a Mediospirifer brachiopod shell from the Windom Member, Middle Devonian, sampled at Barnum Creek quarry just W of NY highway 89 and 0.2 km S of the Ovid/Romulus town line, Seneca County, New York, USA. It is deposited at the Institut für Geowissenschaften , Goethe-Universität , Frankfurt , Germany ( Bo 1/8).

Remarks

Originally established within the ichnogenus Ramodendrina, which is a junior synonym of Clionolithes . The nomen nudum (not formally published) Clionolithes bullahirsuta Plewes, 1996 , from the Lower Jurassic of Yorkshire, UK, is herein regarded as a junior synonym.

Differs from other ichnospecies of Clionolithes by its wide and straight central trunk, the flat overall morphology, with an orientation of the entire trace more or less along the same plane parallel to the substrate surface, the often straight path of the ramifying galleries, and (if preserved) the distinctive cuspate microsculpture ( Fig. 9 D–E).

Figures

Fig. 9. Clionolithes alcicornis (Vogel, Golubic & Brett, 1987) comb. nov. A–B. SEM planar and oblique views of the holotype epoxy cast (together with bryozoan borings) recorded in a Devonian brachiopod shell from New York State, USA. C. SEM of the holotype of junior synonym C. bullahirsuta Plewes, 1996 from the Lower Jurassic of Yorkshire, UK (reproduced from Plewes 1996: pl. 28, fig. 1). D. Close-up of C,illustrating the characteristic cuspatemicrotexture (reproduced from Plewes1996:pl. 28, fig. 3). E. SEM close-up of another specimen showing additional hairy filaments (reproduced from Plewes 1996: pl. 28, fig. 7).

Fig. 1 (page 7). Morphological characters of dendrinid microborings with respect to overall shape and symmetry of the trace, its vertical profile, openings to the substrate surface, orientation of entrance tunnel (where present), branching pattern and surface texture (in order of decreasing relevance as ichnotaxobase).

Fig. 2. Dendrina dendrina (Morris, 1851) comb. nov. A. Original illustration of belemnite with various traces, including D. dendrina (arrows; reproduced from Quenstedt 1849: pl. 30, fig. 36); provenance unresolved. B. Original amalgam, including D. dendrina (arrows and label “d”; reproduced from Quenstedt 1885: pl. 38, fig. 39). C. Re-illustration of original Quenstedt Dendrina (reproduced from Plewes 1996: pl. 22, fig. 5). D. Original illustrations of Talpina dendrina (= D. dendrina; reproduced from Morris 1851: pl. IV, figs 4–7; 5 = Talpina ramosa; 6a = Trypanites solitarius). E. Morris’ original belemnite from the Upper Cretaceous of Norfolk, UK, including the lectotype of D. dendrina (encircled). F–G. Close-up of lectotype under incipient and transparent light; the inlet canal is clearly visible. H–I. Three paralectotypes in the same belemnite. J–K. Planar and oblique views of the holotype of junior synonym D. fluensis Hofmann, 1996, illustrating the typical multi-tier occurrence, irregular branching, and meandering galleries; SEM of epoxy cast of a belemnite from the lower Maastrichtian at Kronsmoor, Germany. L. Holotype of junior synonym D. crassa Hofmann, 1996, within the morphological range of D. dendrina; SEM of epoxy cast of a belemnite from the lower Maastrichtian at Kronsmoor, Germany.

Fig. 3. Dendrina belemniticola Mägdefrau, 1937. A. Original illustration of belemnite with various traces including D. belemniticola (arrow; reproduced from Quenstedt 1849: pl. 30, fig. 36); provenance unresolved. B. Original amalgam, including D. belemniticola (arrow; reproduced from Quenstedt 1885: pl. 38, fig. 39). C. Re-illustration of original Quenstedt Dendrina (reproduced from Plewes 1996: pl. 22, fig. 3); inlet tunnel roughly at 11h in this and the following four sub-figures. D. Lectotype of D. belemniticola in an upper Senonian belemnite from Rosenthal near Peine, Germany (compare to Mägdefrau 1937: pl. IV, fig. 1). E. Holotype of junior synonym D. orbiculata Hofmann, 1996. SEM of epoxy cast of an upper Campanian belemnite from Lüneburg, Germany. F. Paratype of junior synonym D. constans Hofmann, 1996. SEM of epoxy cast of a lower Maastrichtian belemnite from Lüneburg, Germany. G. Holotype of junior synonym D. constans. SEM of epoxy cast of a lower Maastrichtian belemnite from Lüneburg, Germany. E–G illustrate the morphological range of D. belemniticola from almost completely fused radiating galleries, to partially fused galleries forming anastomoses, to rarely fused radiating galleries. H. Detail of partly obscured inlet tunnel connecting to centre of a paratype of junior synonym D. orbiculata. SEM of epoxy cast of an Upper Cretaceous belemnite from Lüneburg, Germany. I. Rarely observed overlap of D. belemniticola in a lower Maastrichtian belemnite from Rügen, Germany. J. Specimen of D. belemniticola (left) next to a D. dendrina (right) at the very tip of a lower Campanian belemnite from Höver, Germany.

Fig. 4. Dendrina lacerata Hofmann, 1996. A. SEM of holotype in an epoxy cast of an upper Campanian belemnite from Misburg, Germany. B. Paratype on the same cast as the holotype. C. Several paratypes in the periphery of a Calcideletrix anomala from the type locality. D. Oblique view of a specimen in a belemnite from the upper Campanian of Kronsmoor, Germany, illustrating a tubular inlet tunnel on the left and peripheral connections to the substrate surface on the right. E. Surface view of a solitary specimen with inlet canal in a belemnite from the upper Campanian of Misburg, Germany.

Fig. 5. Clionolithes radicans Clarke, 1908. A. Original illustration of the lectotype from the Devonian of Mansfield, Pennsylvania, USA (reproduced from Clarke 1908: pl. 11, fig. 1). B–C. Overview and close-up of a shell of the brachiopod Atrypa bearing the lectotype and a number of further specimens. D. Holotype of junior synonym Olkenbachia hirsuta Solle, 1938 from the Devonian near Koblenz, Germany, reproduced from Solle (1938: fig. 2). E–F. Overview and SEM close-up of one of the paratypes of junior synonym Olkenbachia hirsuta (compare with Solle 1938: fig. 4).

Fig. 6. Clionolithes palmatus Clarke, 1908. A. Original drawing of the lectotype, a natural cast in a shell of Loxopteria dispar Sandberger & Sandberger, 1857 from the Upper Devonian of New York State, USA (reproduced from Clarke 1908: pl. 12, fig. 1). B. The lectotype in its present state, either slightly damaged or originally idealised. C. Original drawing of the paralectotype, a natural cast in a Loxonema danai Clarke, 1904 from the type locality (reproduced from Clarke 1908: pl. 12, fig. 2). D. Significantly damaged paralectotype. E–F. A number of natural casts in a shell of Leptostrophia Hall & Clarke, 1892 from the Devonian of Victoria, Australia, including the holotype (close-up in F) and several paratypes of junior synonym Clionolithes sollei Talent, 1963 (compare with Talent 1963: pl. 9, figs 3–8). G. SEM of a specimen cast in epoxy resin by Plewes (1996) from the Devonian of Iowa, USA, illustrating the morphological range within one trace, comprising sheet-like fans and galleries terminating in fine ramifications (reproduced from Plewes 1996: pl. 1, fig. 1).

Fig. 7. Clionolithes pannosus (Solle, 1938) comb. nov. A. Holotype preserved as natural cast in a Devonian brachiopod shell from the Mosel Valley near Münstermaifeld, Germany. B–C. SEM overview and close-up of the holotype. D–F. SEM planar view, oblique view and close-up of pyritised and resinembedded holotype of junior synonym Platydendrina platycentrum Vogel et al., 1987 in a shell of Mediospirifer Bublichenko, 1959 from the Middle Devonian of New York State, USA.

Fig. 8. Clionolithes cervicornis (Vogel, Golubic & Brett, 1987). A. SEM of pyritised and epoxycast holotype in a Devonian brachiopod shell from New York State, USA. B–C. SEM overview and close-up of another specimen from the original material of Vogel et al., recorded in a coral substrate. D–E. Overview and close-up of several natural casts that are part of the suite of paratypes of Olkenbachia hirsuta Solle, 1938 (junior synonym of Clionolithes radicans Clarke, 1908), from the Devonian near Koblenz, Germany (compare with Solle 1938: fig. 5). F. Holotype of nomen nudum “Chondrites” symmetricus (Solle 1938), a morphologically similar (cf.) but unusually large specimen from the Devonian near Koblenz, Germany.