Ammonicrinus sulcatus Kongiel, 1958

Bohatý, Jan, 2011, Revision of the flexible crinoid genus Ammonicrinus and a new hypothesis on its life mode, Acta Palaeontologica Polonica 56 (3), pp. 615-639 : 625-627

Ammonicrinus sulcatus Kongiel, 1958

Figs. 1A , 5A, B , 10 , 13 , 14A , 15C .

pars cf. 1926 Ammonicrinus wanneri ; Springer 1926: pl. 6: 6 = A. cf. sulcatus .

1958 Ammonicrinus sulcatus Kongiel, 1958: 34–36 , fig. 6a, b.

1977 Ammonicrinus sulcatus Kongiel, 1958 ; Piotrowski 1977: 208, 211–213, figs. 2, 4, 5B, 6, pl. 17: 1a–c, 2a–c, 3, 4, 5a, b, pl. 18: 4, 5a, b, 6–8, 10.

1977 Ammonicrinus kongieli Piotrowski, 1977: 213–215 , pl. 18: 1a–c, 2, 3, 9, pl. 19: 1, 2a, b, 3, 4a, b, 5, 6a, b, 7a, b, 8, 9a, b, 10 (not “pl. 18, figs. 1–9” as indicated by Piotrowski [1977: 213]).

1978 Ammonicrinus sulcatus Kongiel, 1958 ; Ubaghs 1978: T78, fig. 57: 9.

2003 Ammonicrinus sulcatus Kongiel, 1958 ; Webster 2003: GSAwebpage.

2003 Ammonicrinus sulcatus Kongiel, 1958 ; Le Menn and Jaouen 2003: 208, fig. 1A.

Holotype: MZ−VIII−EP−1/1. The type locality is Grzegorzowice−Skały ( Holy Cross Mountains , Poland); and the stratigraphy is Member XIV of the Givetian Skały beds (Middle Devonian) (see Piotrowski 1977: 213).

Material.— Type material and new material nos. GIK−2104– 2120 ( Fig. 10A–Q ), GIK−2147–2155 ( Fig. 13A–I ).

Description.— Ammonicrinus sulcatus is distinguished by the fine tubercles on the cup ossicles ( Fig. 10Q 1 ). The mesistele shows nearly linear and wide external flanks and relatively short LCEE; extensions of the mesistele composed of regularly or irregularly arranged columnals with longer and shorter extensions; adult mesistele “pseudo−tuberculated” by echinoid−like spine−tubercles and articulated spines, or distinguished by additional, irregularly arranged, sometimes slightly meandering nodular tubercles bearing the spine−tubercles ( Fig. 10A–G ); columnals of the juvenile mesistele with strongly tuberculated extensions and external flanks ( Fig. 10L–O ); dististele either medium long and composed of numerous columnals (“encased runner−type”) short and composed of only few columnals, or nearly reduced (“settler−type”); the connection between disti− and mesistele is variously formed with floating transitions between those individuals with none (rare) or one to several columnals (characteristic) with laterally positioned enclosure extensions on the proximal−most, barrel−like dististele ( Fig. 10J, K ) and the following mesistele; the planispirally coiled, proximal column is relatively low, wide and barrel−shaped, due to the relatively short LCEE of the mesistele.

Discussion.— Ammonicrinus sulcatus is similar to A. leunisseni sp. nov. and, especially, to A. jankei sp. nov. A. sulcatus developed characteristic and nearly linear external flanks of the mesistele, with short LCEE in contrast to the longer extensions of A. leunisseni sp. nov. Several cup ossicles of A. sulcatus have rudimentary radiating ridges that are not known in A. leunisseni sp. nov. but developed much stronger in A. jankei sp. nov. Because of the longer extensions of the mesistele of A. jankei sp. nov., the shape of the coiled stem is oblate spheroidal, rather than wide and barrel−shaped in A. sulcatus .

Stratigraphic and geographic range.—Givetian (Member XIV of the Givetian Skały beds) of Grzegorzowice−Skały, Holy Cross Mountains ( Poland) and upper Eifelian (Bohnert Member of the Freilingen Formation) of Kerpen and Nollenbach within the Hillesheim Syncline (Eifel, Germany).

showing typical LCEE. E. View of external columnal flanks of a nearly uncoiled mesistele (“runner−type”) (GIK−2137). F. View of external columnal flanks and LCEE of a slightly compressed, coiled mesistele (GIK−2138). G. View of external columnal flanks of a nearly uncoiled mesistele (“runner−type”) (GIK−2139). H. View of external columnal flanks of the mesistele of a partly preserved specimen (GIK−2140). I. View of external columnal flanks of a nearly uncoiled mesistele (“runner−type”) (GIK−2141). J. View of external columnal flanks of the coiled mesistele of a weathered specimen (GIK−2142) on matrix. K–O. Ammonicrinus doliiformis Wolburg 1938a (for 1937). K, L. From the Eifel (locality 9, Appendix 1), Germany, upper Eifelian (Middle Devonian). M. From the Bergisches Land (locality 10, Appendix 1), Germany, Eifelian/Givetian threshold (Middle Devonian). N. From the Eifel (locality 4, Appendix 1), Germany, upper Eifelian (Middle Devonian). O. From the Eifel (locality 5, Appendix 1), Germany, upper Eifelian (Middle Devonian). K. Lateral view of a coiled specimen (GIK−2143) with lost dististele and cracked LCEE of the mesistele, exposing the coiled proxistele and several cup ossicles (arrow). L. Lateral view of a nearly completely coiled specimen (GIK−2144) with lost dististele and cracked LCEE of the mesistele, exposing distal−most part of the coiled proxistele and several cup ossicles (arrow). M. View of external columnal flanks of a preserved, coiled mesistele (GIK−2145) on matrix; the imprint of the uncoiled distal mesistele (“runner−type”), of the dististele and of the holdfast, which is attached to a fenestrate bryozoan (imprint, see arrow), is traced by a dashed line. N. Facet view of a coiled, adult specimen (GIK−2146) with exposed distal part of the proxistele and disarticulated remains of the arms (arrows). O. Perfect, three dimensionally preserved, adult specimen ( MB.E.−287, original of Krause 1927). Coiled mesistele in lateral view (O 1), dististele, attachment and spines missing, centres of tuberculated radials partly visible (arrow); the specimen is infested by a (?)craniid brachiopod (arrow on the left); opposite lateral view (O 2), centres of radials partly visible (arrow); oblique lateral view (O 3); view of the external flanks of the mesistele (O 4) (centre and upper part of the figure) and of the facet area of distal mesistele (below), showing wide barrel−shaped outline. Scale bars 10 mm.

doi:10.4202/app.2010.0020

Ammonicrinus doliiformis Wolburg, 1938a (for 1937) Figs. 2E , 3 , 4A , 9K–O , 15D .

1938a (for 1937) Ammonicrinus doliiformis Wolburg, 1938a: 230–241 , figs. 1–5, pl. 17: 1–5, 6a, b, 7, pl. 18: 1(?), 2a, b, 3, 4, 5–7(?), 8.

1927 Ammonicrinus wanneri Springer, 1926 ; Krause 1927: pl. 8: 1–6.

1952 Ammonicrinus doliiformis Wolburg, 1938a ; Ubaghs 1952: 216– 218, pl. 3: 1–5.

1978 Ammonicrinus doliiformis Wolburg, 1938a ; Ubaghs 1978: T64, fig. 44: 3.

1978 Ammonicrinus wanneri Springer, 1926 ; Ubaghs 1978: T78, fig. 57: 6, 7.

2003 Ammonicrinus doliiformis Wolburg, 1938a ; Webster 2003: GSAwebpage.

2005 “ Ammonicrinus wachtbergensis ” sp. nov.; Hauser 2005: 4, 23–25, figs. 1, 15a, b, second unnumbered fig. below on p. 34, pl. 1: 3a–c, front and backside covers of private publication.

Holotype: Due to world war damages, only one cast of the dististele of

Wolburg’s (1938a) type material is still preserved in the Geowissenschaftliches Zentrum der Universität Göttingen, Germany (without re−

pository−number) (Reimund Haude, personal communication 2009). The type locality is Plettenberg−Ohle at the Lenne River ( SE of Werdohl,

Märkischer Kreis, Sauerland; UTM unknown); and the stratigraphy is

“Selscheider Formation” sensu Wolburg (1938a: 230); more probable,

the type material came from the Odershäuser Formation of the Eifelian/

Givetian threshold (Middle Devonian) (Martin Basse, personal communication 2009).

Material.— Type material ( Figs. 2E , 9O ), originals of Wolburg (1938a) ( Fig. 3A–C ) and Ubaghs (1952) (no. SMF− XXIII−165a; Fig. 4A ) and new material nos. GIK−2143–2146 ( Fig. 9K–N ).

Description.— Ammonicrinus doliiformis is distinguished by fine tubercles on the surface of the cup ossicles ( Fig. 9K, L, O 1 , O 2 ). The arms are relatively long and formed by medium wide, short and straight or laterally somewhat curved brachials. The dististele ( Fig. 3A 1 , A 2 ) is long and composed of numerous columnals (“encased runner−type”), the distal−most dististele is connected with a substrate−controlled holdfast, typically in form of a relatively small attachment disc ( Fig. 3A 1 , A 2 ); the LCEE of the wide mesistele are composed of characteristic, regularly or irregularly (rare) arranged columnals with longer and shorter extensions. These are interconnected with several columnals with broadened extensions that could intermesh in a closed coiled position and are combined with smaller, “regular” columnals ( Figs. 9O 1 , O 2 , 15D ); connection between disti− and mesistele distinguished by a triangular columnal without extensions ( Fig. 3A 1 , A 3 ); columnals of the mesistele with long, less curved external flanks with relatively thin cross sections; shape of coiled stem wide barrel−shaped ( Fig. 9O 3 , O 4 ); the cup is partly visible in lateral radial view ( Fig. 9O 1 , O 2 ); mesi− and dististele covered by echinoid−like spine−tubercles, which bear articulated spines.

Discussion.— Ammonicrinus doliiformis is similar to A. leunisseni sp. nov. Wolburg’s (1938a) species has a wider diameter of the coiled stem and a characteristic connection between the disti− and mesistele, which is distinguished by a triangular columnal without extensions in opposition to the variously formed connection between the disti− and mesistele of A. leunisseni sp. nov.

After studying the holotype of “ Ammonicrinus wachtbergensis Hauser, 2005 ” (= original of Krause 1927, figured as A. wanneri ), it is clearly evident that the specimen is a typical adult that is a three−dimensionally preserved A. doliiformis . The specimen came from the Eilenberg Member of the uppermost part of the Freilingen Formation (upper Eifelian) of locality 5. This stratigraphic level is most famous for A. doliiformis and could be correlated with several localities within the Eifel (e.g., with the deposits of the Freilingen Formation of village Gondelsheim within the Prüm Syncline or with locality 4). Also the stratum typicum at the A. doliiformis type locality (locality 11, also see locality 10) correlates approximately with the Eifel findings. Therefore, “ A. wachtbergensis Hauser, 2005 ” is declared a subjective junior synonym of A. doliiformis .

Stratigraphic and geographic range.—Eifelian (Selscheid Formation) of Ohle, Sauerland ( Germany), Eifelian (Oderhäuser Formation) of Lindlar−Hartegasse ( Germany), Eifelian (Eilenberg Member of the Freilingen Formation) of Sötenich (Sötenich Syncline, Eifel), Gondelsheim (Prüm Syncline, Eifel), Niederehe (Hillesheim Syncline, Eifel) and “Rommersheim Formation” of the “Auburg” near Gerolstein (Gerolstein Syncline, Eifel) ( Germany).

Figures

Fig. 1. A. Reconstruction of the Ammonicrinus life time position (modified after Piotrowski 1977:208, fig. 2). B. Ammonicrinus plate diagram (not to scale).

Fig. 5. Schematic illustrations of Ammonicrinus sulcatus Kongiel, 1958 after Piotrowski (1977). A. Lateral cross section through the feeding crinoid (Piotrowski 1977: 209, fig. 3). B. Former reconstruction of life time position (Piotrowski 1977: 208, fig. 2). Not to scale.

Fig. 10. The lecanocrinid species Ammonicrinus sulcatus from locality 1 (A–G, I–Q) and 2 (H) (see Appendix 1). A–D. Facet views of GIK−2104–2107, showing nodular tubercles and spine−tubercles on exterior flanks of the columnals of the mesistele. E. Facet view (E1) and view of the exterior flank of a specimen (E2) (GIK−2108), showing tubercles and spine−tubercles on exterior flank of the columnal of the mesistele. F. Facet view of a specimen (GIK−2109), showing tubercles and spine−tubercles on exterior flank of the columnal of the mesistele. G. Facet view of a strongly sculptured columnal (GIK−2110) of the distal−most mesistele, showing connection to the dististele. H. Facet view of a columnal of the distal−most mesistele (GIK−2111), showing long LCEE and connection to the dististele.I. Facet view of a columnal of the distal−most mesistele (GIK−2112), showing relatively long LCEE and connection to the dististele. J. Interior view of a distal−most, barrel−like columnal of the mesistele (GIK−2113) with LCEE. K. Interior view of a distal−most, barrel−like columnal of the mesistele (GIK−2114), with partly preserved LCEE. L. Facet view of a juvenile distal columnal of the mesistele (GIK−2115) with nodular tubercles on exterior flank and on LCEE. M–N. Juvenile columnals of the proximal mesistele in facet views, showing well developed nodes on exterior flanks. M. GIK−2116. N. GIK−2117. O. Facet view of a juvenile distal columnal of the mesistele (GIK−2118) with nodular tubercles on exterior flank and on LCEE. P. Lateral view (P1) and view of the exterior flank (P2) of the partly preserved mesistele (GIK−2119); the specimen shows nodular tubercles, spine−tubercles and a few partly preserved spines (arrow). Q. Facet view (Q1) and lateral view (Q2) of a cracked, coiled mesistele (GIK−2120), showing several tuberculated and concave ossicles of the cup (arrows). Scale bars 10 mm.

Fig. 13. Postmortem epizoan encrusting on disarticulated columnals of the lecanocrinid crinoid species Ammonicrinus sulcatus Kongiel, 1958. A–G. From the Eifel (locality 1, Appendix 1), Germany, upper Eifelian (Middle Devonian). H, I. From the Eifel (locality 2, Appendix 1), Germany, upper Eifelian (Middle Dvonian). A. View of external flanks of a pluricolumnal of the mesistele (GIK−2147), encrusted by a trepostomate bryozoan (Leptrotrypella(?) sp.) (arrows). B. Internal view of a pluricolumnal of the distal−most mesistele (GIK−2148), encrusted by a cystoporate bryozoan (Eridopora(?) sp.) (arrows). C. Facet view of an isolated, distal−most columnal of the mesistele (GIK−2149), encrusted by a trepostomate bryozoan [Leptrotrypella(?) sp.] (arrows). D. External flanks of a pluricolumnal of the mesistele (GIK−2150), encrusted by a trepostomate bryozoan (Eostenopora(?) sp.) (see arrows below); the bryozoan is infested by a crinoid attachment disc (arrows in D2); general view (D1), detail view (D2). E. View of external flanks of a pluricolumnal of the mesistele (GIK−2151), encrusted by a crinoid holdfast (arrow). F. Facet view of an isolated columnal of the mesistele (GIK−2152), encrusted by a cystoporate bryozoan (Cyclotrypa(?) sp.) (arrows). G. Facet view of a pluricolumnal of the mesistele (GIK−2153), encrusted by a cystoporate bryozoan (Cyclotrypa(?) sp.) (arrows). H. Facet view of an isolated columnal of the mesistele (GIK−2154) (H1), encrusted by microconchid valves (see arrows in detail view (H2). I. Facet view of an isolated columnal of the mesistele (GIK−2155), encrusted by a holdfast of a fenestrate bryozoan (arrow). Scale bars 10 mm.

Fig. 14. A. Schematic reconstruction of Ammonicrinus (strongly modified after Piotrowski 1977: 209, fig. 3) of a feeding Ammonicrinus within low−intensity current water. Alternating water pressure was possibly generated in the interior of the enrolled proximal stem by slow, bellow−like partial opening and closing (red arrows) of the base of the central mass; due to the synarthrial (bifacial) articulation of the ammonicrinid mesistele that developed two huge ligamentary facets (orange), separated by the fulcrum, bellow−like partial opening could possibly enabled by stiffening of the outer ligaments (see orange bars in A1); closing could be controlled by stiffening of the inner ligaments (see orange bars in A2). A1, suction during opening may result from low−pressure (P−) and create an ingesting water flow (blue arrow); A, ejection during closure (red arrow) resulted from overpressure (P+); to minimise faecal recycling, 2 the water ejection may have occurred laterally (blue arrows), feasibly at both lateral centres, which have “openings”. B. Lobe−like enlarged appendages (framed in red) could possibly support the lateral water faecal−ejection (modified from Ubaghs 1952: pl. 1: 4). C. Reconstruction of a feeding “encased runner−type” of A. leunisseni sp. nov., attached to a tabulate coral (model); the spined specimen dwelled enrolled on the muddy seafloor; alternating water pressure was obviously generated in the interior of the enrolled proximal stem globe by non−muscular, probably MCT−controlled, slow, bellow−like partial opening and closing of the oblate sphere at its bottom (dashed arrow); suction during opening created an ingesting water flow (see arrow on the left), which was funnelled in a “canal”, formed by the unspined interior of the columnals of the mesistele, whose U−shaped LCEE additionally formed a protection against immersive sediment; ejection during closure resulted from overpressure; to minimise faecal recycling, the water ejection occurred supposably laterally, feasibly at both lateral centres, which accordingly show “openings” (see arrows on the right). Not to scale.

Fig. 15. Schematic sketches of different LCEE of the mesistele in uncoiled (above) and coiled positions (below), indicating evolution of perfecting the crown−encasing in coiled position by modifying the extensions form Emsian to Givetian. A. Lateral view of Ammonicrinus kerdreoletensis Le Menn and Jaouen, 2003, showing similar shaped columnals with very short LCEE; thus, the crown is laterally nearly unprotected in coiled position.B. Lateral view of Ammonicrinus wanneri Springer, 1926, with lengthened LCEE of the similar shaped columnals, which lattice−like guarded the crown in coiled position. C. Lateral view of Ammonicrinus sulcatus Kongiel, 1958, showing smaller columnals of the mesistele, which are interconnected with longer ones and afford lateral density of the coiled stem. D. Lateral view of Ammonicrinus doliiformis Wolburg, 1938a (for 1937), showing regularly or irregularly arranged columnals with longer and shorter LCEE, which were interconnected with several columnals showing broadened convex and concave extensions that could interlock in coiled position.

Fig. 2. The first figures of Ammonicrinus from Springer (1926) and Krause (1927). A. Ammonicrinus wanneri Springer, 1926 (from Springer 1926: pl. 6: 4a, b). A1, view of the extetrnal flanks of the coiled mesistele; A2, coiled mesistele in lateral view. B. Ammonicrinus leunisseni sp. nov. (= “A. wanneri” in Springer 1926: pl. 6: 5, 5b). B1, view of the extetrnal flanks of the coiled mesistele; B2, coiled mesistele in lateral view. C. Photograph of the holotype of Ammonicrinus wanneri Springer, 1926, USNM−S2115; lateral view of coiled mesistele; connection between mesi− and dististele, dististele and attachment missing (see fracture surface at distal mesistele). D. Photograph of USNM−S2115, the Springer (1926) original of “Ammonicrinus wanneri” (= A. leunisseni sp. nov. herein); lateral view of coiled mesistele; connection between mesi− and dististele, dististele and attachment missing (see fracture surface at distal mesistele). E. Ammonicrinus doliiformis Wolburg, 1938a (for 1937) (= “A. wanneri” in Krause 1927: pl. 8: 4, 2). E1, view of the extetrnal flanks of the coiled mesistele; E2, coiled mesistele in lateral view. A–D from the Middle Devonian, Eifel Limestone; Prüm, Eifel, Germany (Springer 1926: 25); E from the Middle Devonian of Sötenich, Eifel (Krause 1927: 456). A, B, E not to scale; C, D scale bars 10 mm.

Fig. 3. The lecanocrinid species Ammonicrinus doliiformis Wolburg, 1938a (for 1937) from the Selscheid Formation of Ohle, Sauerland (Wolburg 1938a: 230). A. Casts of nearly complete specimen. Specimen attached to a brachiopod valve (right arrow) (A1), showing the characteristic triangular connection between mesi− and dististele (left arrow) and slightly compressed mesistele (from Wolburg 1938a: pl. 17: 1); detail view of the attachment disc (arrow) (A2), encrusting the brachiopod (from Wolburg 1938a: pl. 18: 8); detail view of the triangular connection between mesi− and dististele (arrow) (A3) (from Wolburg 1938a: pl. 17: 6a); detail view of the coiled, slightly compressed mesistele (A4) (from Wolburg 1938a: pl. 17: 4). B. Former assumed reconstruction of life mode, figured with a crown that protrudes toward the lateral−exterior (arrow) (from Wolburg 1938a: 240, fig. 5). C. Former assumed reconstruction of the crown (1938a: 233, fig. 4). Not to scale.

Fig. 4. First illustration of the actual plate diagram and definition of genus Ammonicrinus as a lecanocrinid Flexibilia by Ubaghs (1952). A. Ammonicrinus doliiformis Wolburg, 1938a (for 1937), SMF−XXIII−165a from the “Rommersheim Formation” of the Auburg, Gerolstein, Eifel, Germany (Ubaghs 1952: 220). View of coiled mesistele (A1); view of exposed proxistele (A2) (taken from Ubaghs 1952: pl. 3: 1, 3). B. Anomalous crown of “Ammonicrinus wanneri” from the “Rommersheim Formation” of the Steineberg, N of Kerpen, Eifel, Germany (Ubaghs 1952: 220) (= holotype of A. jankei sp. nov., no. SMF−XXIII−167a) coiled by the mesistele.View of the coiled mesistele (B1) (Ubaghs 1952: pl. 1: 3); partly excavated crown (B2), showing radiating ridges on radials and one slightly lobe−like enlarged appendage that possibly could support the lateral water respectively faecal−ejection (arrow) (Ubaghs 1952: pl. 1: 4); excavated crown in lateral view (B3, B4), the second “radianal plate” respectively “supplementary plate” (see arrows) is based on an anomaly (Ubaghs 1952: pl. 2: 3, 2); plate diagram (B5), showing the two anomalous plates (arrows) (slightly modified after Ubaghs 1952: 205, fig. 1); schematic drawing of the coiled specimen (B6); reconstruction of the assumed living feeding position (B7) (Ubaghs 1952: 110, fig. 2; p. 223, fig. 5). Not to scale.

Fig. 9. Lecanocrinid Ammonicrinus species. A–J. Ammonicrinus wanneri Springer, 1926. A–I. From the Eifel (locality 3, Appendix 1), Germany, Lower Givetian (Middle Devonian). J. From the Eifel (locality 7, Appendix 1), Germany, Lower Givetian (Middle Devonian). A. Lateral view of a partly preserved specimen (GIK−2133) with coiled mesistele. B. Lateral view, respectively view of external columnal flanks of the coiled mesistele of a partly preserved specimen (GIK−2134) with one preserved, postulated cup ossicle (arrow). C. View of external columnal flanks of the mesistele of a partly preserved specimen (GIK−2135). D. Lateral view, respectively view of external columnal flanks of the coiled mesistele of a partly preserved specimen (GIK−2136), +