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New insights on Tournaisian–Visean (Carboniferous, Mississippian) athyridide, orthotetide, rhynchonellide,
and strophomenide brachiopods from southern Belgium

Bernard Mottequin and Eric Simon

Plain Language Abstract

Brachiopods are benthic, microphagous suspension-feeder and marine invertebrates known from the Lower Cambrian to the present that were particularly abundant and diverse during the Palaeozoic. Thanks to their organophosphatic or organocarbonate shell, their preservation potential is high and thus, they can be used for a large panel of applications, such as biostratigraphy, geochemistry, or palaeogeography. This paper investigates 12 Lower Carboniferous (Tournaisian and Visean) brachiopod species belonging to seven genera of the orders Athyridida, Orthotetida, Rhynchonellida, and Strophomenida from southern Belgium (Namur–Dinant Basin), which is one of the best documented areas for Lower Carboniferous rocks, from both lithostratigraphic and biostratigraphic viewpoints. Two species are new (Leptagonia franca sp. nov., Serratocrista scaldisensis sp. nov.), and a new name is proposed (Nucleospira hannoniae nom. nov.) to solve the primary homonymy between Athyris globulina de Koninck and A. globulina Waagen. Taxonomic discussion involves selection of the holotype of Retzia intermedia de Koninck (definitely not an athyridide) and of lectotypes for several species of athyridides (Athyris globulina de Koninck, A. vittata de Koninck, and Retzia davidsoni de Koninck), and strophomenides (Orthis cylindrica M'Coy, Schellwienella radialiformis Demanet, and S. ornata Demanet). The palaeoecology of the investigated species and their biotic interactions with other organisms (e.g., drill holes, epizoans) are described. The analysis of the published data on the athyridides, orthotetides, and strophomenides occurring in southern Belgium during the Devonian–Lower Carboniferous reveals that our knowledge of these brachiopods from this part of the southern margin of the Laurussian continent is currently insufficient to assess the aftermath of the different biological crises that took place during this time span, notably the Hangenberg Crisis at the end of the Devonian.

Resumen en Español

Nuevas perspectivas sobre los braquiópodos Athyridida, Orthotetida, Rhynchonellida y Strophomenida del Tournaisiense-Viseense (Carbonífero, Misisípico) del sur de Bélgica

Doce especies carboníferas seleccionadas pertenecientes a siete géneros de los órdenes Athyridida, Orthotetida, Rhynchonellida y Strophomenida (Brachiopoda) del sur de Bélgica son revisadas y/o discutidas y algunas veces fotografiadas por primera vez desde su primera descripción que data del siglo XIX. El material investigado proviene de las áreas tipo históricas del Tournaisiense y Viseense (Tournai y Visé, respectivamente) y de las acumulaciones de carbonato "Waulsortiense" desarrolladas en la cuenca de Namur-Dinant. Dos especies son nuevas (Leptagonia franca sp. nov., Serratocrista scaldisensis sp. nov.), y se propone un nuevo nombre (Nucleospira hannoniae nom. nov.) para resolver la homonimia primaria entre Athyris globulina de Koninck y A. globulina Waagen. La discusión taxonómica implica la selección del holotipo de Retzia intermedia de Koninck (definitivamente no se trata de un Athyridida) y de lectotipos de varias especies de Athyridida (Athyris globulina de Koninck, A. vittata de Koninck, y Retzia davidsoni de Koninck) y de Strophomenida (Orthis cylindrica M'Coy, Schellwienella radialiformis Demanet, y S. ornata Demanet). Se describe la paleoecología de las especies investigadas y sus interacciones bióticas con otros organismos (por ejemplo, agujeros de perforación, epizoos). El análisis de los datos publicados sobre los Athyridida, Orthotetida y Strophomenida que se desarrollaron en el sur de Bélgica durante el Devónico- Misisípico revela que nuestro conocimiento de estos suspensívoros en esta parte del margen meridional de Laurusia es actualmente insuficiente para evaluar las consecuencias de las diferentes crisis biológicas que tuvieron lugar durante este lapso de tiempo, notablemente la Hangenberg Crisis al final del Fameniense.

Palabras clave: Brachiopoda; taxonomía; nuevas especies; paleobiología; Devónico-Misisípico; Bélgica

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Approches nouvelles des brachiopodes athyridides, orthotetides, rhynchonellides et strophomenides d'âge tournaisien-viséen (Carbonifère, Mississippien) du Sud de la Belgique

Douze espèces sélectionnées du Carbonifère du Sud de la Belgique, appartenant à sept genres des ordres Athyridida, Orthotetida, Rhynchonellida et Strophomenida (Brachiopoda), sont révisées et/ou discutées, et parfois photographiées pour la première fois depuis leur première description remontant au 19ème siècle. Le matériel étudié provient des régions types historiques des étages Tournaisien et Viséen, à savoir celles de Tournai et de Visé, ainsi que des monticules carbonatés waulsortiens développés dans le bassin de Namur-Dinant. Deux espèces sont nouvelles (Leptagonia franca sp. nov., Serratocrista scaldisensis sp. nov.) et un nouveau nom est proposé (Nucleospira hannoniae nom. nov.) pour résoudre l'homonymie primaire entre Athyris globulina de Koninck and A. globulina Waagen. Les discussions taxonomiques impliquent la sélection de l'holotype de Retzia intermedia de Koninck (rejeté définitivement des athyridides) et des lectotypes de plusieurs espèces d'athyridides (Athyris globulina de Koninck, A. vittata de Koninck, et Retzia davidsoni de Koninck) ainsi que de strophomenides (Orthis cylindrica M'Coy, Schellwienella radialiformis Demanet, et S. ornata Demanet). La paléoécologie des espèces investiguées et leurs interactions biologiques avec d'autres organismes (e.g., perforations, épizoaires) sont décrites. L'analyse des données publiées sur les athyridides, les orthotetides, et les strophomenides reconnus dans les successions dévoniennes et carbonifères du Sud de la Belgique révèle que notre connaissance de ces suspensivores de cette partie de la marge méridionale de Laurussia est encore insuffisante pour évaluer les conséquences des différentes crises biologiques majeures qui se sont déroulées durant cet intervalle, notamment la Crise de Hangenberg à la fin du Famennien.

Mots-clés : Brachiopoda; nouvelles espèces; paléobiologie; Dévonien-Mississippien; Belgique

Translator: Authors

Deutsche Zusammenfassung

Neue Einblicke in athyridide, orthotetide, rhynchonellide und strophomenide Brachiopoden aus dem Tournai–Visé (Karbon, Mississippium) von Südbelgien

Es werden 12 Arten aus sieben Gattungen der Ordnungen Athyridida, Orthotetida, Rhynchonellida und Strophomenida (Brachiopoda) aus Südbelgien überarbeitet und/oder besprochen, sowie zum ersten Mal seit ihrer Erstbeschreibung im 19ten Jahrhundert fotografiert. Das untersuchte Material stammt aus den historischen Typlokalitäten des Tournai und Visé und aus den Waulsortischen Karbonat-Buildups, die sich im Namur-Dinant Becken entwickelt haben. Zwei Arten sind neu (Leptagonia franca sp. nov., Serratocrista scaldisensis sp. nov.) und es wird ein neuer Name vorgeschlagen (Nucleospira hannoniae nom. nov.), um die primäre Homonomie zwischen Athyris globulina de Koninck und A. globulina Waagen aufzulösen. Taxonomische Diskussion betrifft die Auswahl des Holotyps von Retzia intermedia de Koninck (definitiv kein Athyridide) und die Lectotypen einiger Athyrididen (Athyris globulina de Koninck, A. vittata de Koninck und Retzia davidsoni de Koninck) und Strophomeniden (Orthis cylindrica M'Coy, Schellwienella radialiformis Demanet und S. ornata Demanet). Es wird die Paläoökologie der untersuchten Arten und ihre biotischen Interaktionen mit anderen Organismen (z.B. Bohrlöcher, Epizoen) beschrieben. Die Untersuchungen der publizierten Daten über die Athyrididen, Orthotetiden und Strophomeniden die während des Devon bis Mississippium in Südbelgien auftraten, zeigen, dass unsere Kenntnisse über diese Suspensionsfresser in diesem Teil des südlichen Randes von Laurussia derzeit nicht ausreichen, um die Folgen verschiedener biologischer Krisen in dieser Zeit zu beurteilen, hier besonders die Hangenberg Krise am Ende des Famenne.

Schlüsselwörter: Brachiopoda; Taxonomie; neue Art; Paläobiologie; Devon–Mississippium; Belgien

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

FIGURE 1. General context of Lower Carboniferous sedimentation in north-western Europe showing the distribution of emergent areas and Waulsortian buildups at the end of the Tournaisian (modified from Ziegler, 1990; Devuyst and Dehantschutter, 2007).

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FIGURE 2. Schematic geological map of southern Belgium with location of the fossiliferous localities (modified from de Béthune, 1954). Abbreviations: D, Dréhance; F, Furfooz; G, Germany; HSMOTS, Haine-Sambre-Meuse Overturned Thrust Sheets; L, Grand Duchy of Luxemburg; N, The Netherlands; S, Sosoye.

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FIGURE 3. Late Tournaisian sedimentation areas in the Namur-Dinant Basin (not palinspastically restored; modified from Hance et al., 2001; Poty et al., 2006; Poty, 2016) with with location of the fossiliferous localities. Abbreviations: ASA, Avesnois sedimentation area; CSA, Condroz sedimentation area; D, Dréhance; DSA, Dinant sedimentation area; F, Furfooz; G, Germany; HSA, Hainaut sedimentation area; L, Grand Duchy of Luxemburg; N, The Netherlands; NSA, Namur sedimentation area; S, Sosoye; VSA, Visé-Maastricht sedimentation area.

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FIGURE 4. Distribution of the athyridides, orthotetides and strophomenides described and/or discussed herein within the essentially carbonate Tournaisian and Visean succession of southern Belgium (see text). Stratigraphy, lithostratigraphy (incompletely represented here for the Hainaut and Visé-Maastricht sedimentation areas; formations and members are in roman and italic letters, respectively), and biostratigraphy are adapted from Poty et al. (2002, 2006, 2014). The lenticular Vignobles Member is not represented here as it is rarely developed (it is comprised between the Vaulx Member of the Tournai Formation and the Antoing Formation). Note that the “Grande Brèche” is an informal stratigraphic unit resulting from the dissolution of evaporitic levels in the Grands Malades Formation; other brecciated horizons occur in the Lives Formation.

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FIGURE 5. Leptagonia franca sp. nov. from the Tournai area, Tournai Formation (Tournaisian). 1-6, RBINS a5891 (holotype), articulated specimen, almost complete, in ventral (perpendicular to the surface of the alae), dorsal, lateral, posterior, anterior, and ventral (perpendicular to the disc) views. 7-12, RBINS a5892, articulated specimen, almost complete, in ventral (perpendicular to the surface of the alae), dorsal, lateral, posterior, anterior, and ventral (perpendicular to the disc) views. 13-18, RBINS a13099, articulated specimen (with an auloporid tabulate fixed to its right ventral flank), almost complete, in ventral (perpendicular to the surface of the alae), dorsal, lateral, posterior, anterior, and ventral (perpendicular to the disc) views. Scale bar equals 10 mm.

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FIGURE 6. Leptagonia franca sp. nov. from the Tournai area, Tournai Formation (Tournaisian). 1-6, RBINS a5829, complete articulated specimen (with Petrocrania? ryckholtiana (de Koninck, 1843) attached to ventral and dorsal valves) in ventral (orientation as in Figure 5.1), ventral (perpendicular to the visceral disc), dorsal, lateral, posterior, and anterior views. 7-12, RBINS a5895, complete articulated specimen (with Petrocrania? ryckholtiana and microconchid attached to ventral valve), in ventral (orientation as in Figure 5.1), lateral, ventral (perpendicular to the visceral disc), dorsal, posterior and anterior views. 13-17, RBINS a13100, articulated specimen, almost complete, in ventral (orientation as in Figure 5.1), dorsal, lateral, posterior, and anterior views. 18-22, RBINS a13101, articulated specimen, almost complete, in ventral (orientation as in Figure 5.1), dorsal, lateral, posterior and anterior views. 23-27, RBINS a13102, articulated specimen, almost complete, in ventral (orientation as in Figure 5.1), dorsal, lateral, posterior, and anterior views. Scale bar equals 10 mm.

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FIGURE 7. Leptagonia franca sp. nov. from the Tournai area, Tournai Formation (Tournaisian). 1-3, Detail of the small pseudodeltidium and apical foramen of three articulated specimens with the ventral valve on top: RBINS a5896 (1), RBINS a13103 (2), and RBINS a5895 (3). 4-5, RBINS a5902, incomplete ventral valve showing the posterior morphology. 6-7, RBINS a a13104, incomplete ventral valve interior showing the posterior morphology and posterior view notably displaying the prominent teeth. 8, RBINS a5903, incomplete ventral valve interior showing the posterior morphology. 9, RBINS a5899, almost complete dorsal valve interior in ventral view. 10-13, RBINS a5897, almost complete dorsal valve interior in posterior and ventral views, detail of the lateral vascular canals, and close-up of the cardinal process and the muscular field. Scale bars equal 2.5 mm (1-3), 5 mm (4-8, 12-13), 10 mm (9-11).

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FIGURE 8. Scatter diagrams of Leptagonia franca sp. nov. N: number of specimens measured. 1, Relation between shell width and length. 2, Relation between shell width and shell length/shell width ratio. 3, Relation beween shell width and thickness. 4, Relation between shell width and shell thickness/shell witdth ratio. 5, Relation between unrolled ventral valve length and ratio between unrolled ventral trail length (Vtl) and unrolled ventral disc length (Vdl). 6, Relation between unrolled dorsal valve length and ratio between unrolled dorsal trail length (Dtl) and unrolled dorsal disc length (Ddl).

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FIGURE 9. 1-4, Leptagonia sp. indet. from Sosoye, Waulsort Formation (Tournaisian, Ivorian), RBINS a11747, almost complete ventral valve in ventral, ventral (perpendicular to the disc), lateral and anterior views. 5-26, Leptagonia cf. caledonica Brand, 1972 from Visé, Visé Formation (Visean, Warnantian). 5-9, RBINS a5915, incomplete articulated specimen in ventral (perpendicular to the disc), dorsal, lateral, posterior, and anterior views. 10-14, RBINS a5914, incomplete articulated specimen in ventral (perpendicular to the disc), dorsal, lateral, posterior, and anterior views. 15-20, ULg.PA. 2016.12.25/1, articulated specimen, almost complete, in ventral, ventral (perpendicular to the disc), dorsal, lateral, posterior and anterior views. 21-26, ULg.PA.2016.12.25/2, articulated specimen, almost complete, in ventral, lateral, ventral (perpendicular to the disc), dorsal, posterior, and anterior views. Scale bar equals 10 mm.

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FIGURE 10. Leptagonia cf. caledonica Brand, 1972 from Visé, Visé Formation (Visean, Warnantian). 1-5, 19-20, ULg.PA.2016.12.25/3, internal mould in ventral, dorsal, lateral, posterior and anterior views, and detail (SEM) of the muscular platform (latex cast). 6-10, 21-22, ULg.PA.2016.12.25/4, internal mould in ventral, dorsal, lateral, posterior, and anterior views, and detail (SEM) of the muscular platform (latex cast). 11-15, 23-24, ULg.PA.2016.12.25/5, internal mould in ventral, dorsal, lateral, posterior, and anterior views, and detail (SEM) of the muscular platform (latex cast). 16-18, ULg.PA.2016.12.25/6, distorted internal mould in dorsal view and close-up (SEM) of the muscular platform (latex cast). Scale bars equal 5 mm.

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FIGURE 11. Leptagonia cf. caledonica Brand, 1972 from Visé, Visé Formation (Visean, Warnantian). 1-4, ULg.PA.2016.12.25/7, distorted, articulated internal mould in ventral view and detail of the latex mould of the muscular platform in three different views. Scale bar equals 5 mm (1), 2.5 mm (2-4).

 

 

FIGURE 12. Schellwienella radialiformis Demanet, 1934 from the Tournai area, Tournai Formation (Tournaisian). 1-5, RBINS a5924, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views. 6-10, RBINS a5928, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views. 11-15, RBINS a5929, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior and anterior views. 16-20, RBINS a5930 (lectotype), articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views. 21-25, RBINS a5926, almost complete articulated specimen (with a microconchid attached to the ventral valve, close to the anterior margin) in ventral, dorsal, lateral, posterior, and anterior views. Scale bars equal 10 mm.

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FIGURE 13. Schellwienella radialiformis Demanet, 1934 from the Tournai area, Tournai Formation (Tournaisian). 1-5, RBINS a5927, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior and anterior views. 6-10, RBINS a13105, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views. 11-12, RBINS a5924, detail (SEM) of the pseudodeltidium and of the koskinoid perforations penetrating in the umbonal part of the ventral valve. 13, RBINS a5928, detail of the microornament on dorsal valve (SEM). 14-17, RBINS a5925, incomplete ventral valve in external and internal views and detail (SEM) of the posterior internal and external morphology. 18-24, RBINS a13106, almost complete dorsal valve in dorsal, lateral, posterior, and internal views and detail (SEM) of the posterior morphology (cardinal process). Scale bars equal 10 mm (1-10, 14, 18-22), 1 mm (11), 0.25 mm (12), 0.5 mm (13), 2 mm (17, 23), 2.5 mm (24), 5 mm (15).

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FIGURE 14. Schellwienella radialiformis Demanet, 1934 from the Tournai area, Tournai Formation (Tournaisian). 1-4, RBINS a13107, almost complete dorsal valve in dorsal, lateral and posterior views and detail of the chilidium and cardinal process lobes. 5-10, RBINS a5921, articulated internal mould with shelly remains in ventral, dorsal, lateral, posterior and anterior views and close-up of the ventral interarea (5). Scale bars equal 10 mm (1-2, 4-6), 1 mm (4, 5).

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FIGURE 15. Schellwienella ornata Demanet, 1934 from Visé, Visé Formation (Visean, Warnantian). 1-8, RBINS a5922 (lectotype), articulated internal mould in ventral, enlarged ventral (showing the extent of the traces left by the dissolved dental plates), dorsal, lateral, posterior, and anterior views, and internal views of a latex cast (SEM) showing the internal morphology of the posterior part of the specimen (7-8). 9-14, RBINS a5923 (paralectotype), articulated specimen (partly sectioned, probably by Demanet) in ventral, dorsal, lateral, posterior, and anterior views, and close-up (SEM) of the parvicostellate ornamentation in the central part of the dorsal valve. Scale bars equal 10 mm (1, 3-6, 9-13), 5 mm (2), 2.5 mm (7-8), 1.25 mm (14).

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FIGURE 16. Schellwienella cylindrica (M’Coy, 1844), from the Visean (Brigantian) of Castle Espie (Comber, Co. Down, Ireland). 1-3, NMING:F6267 (lectotype), almost complete dorsal valve in dorsal, lateral, and posterior views. Scale bar equals 10 mm.

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FIGURE 17. Serratocrista scaldisensis sp. nov. from the Tournai area, Tournai Formation (Tournaisian) (all SEM). 1-6, RBINS a13108, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views, and close-up of the umbo showing the absence of dental plates. 7-11, RBINS a13109, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 12-16, RBINS 13110 (holotype), articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 17-21, RBINS a13111, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 22-25, RBINS a13112, almost complete ventral valve in ventral, lateral, posterior, and anterior views. Scale bars equal 1 mm (6), 5 mm (1-5, 7-25).

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FIGURE 18. Serratocrista scaldisensis sp. nov. from the Tournai area, Tournai Formation (Tournaisian). 1, 6, RBINS a13110 (holotype), close-up of the strongly convex pseudodeltidium, and detail of the microornament (central part of the dorsal valve). 2-3, 19-21, RBINS a13111, detail of the strongly convex pseudodeltidium and of the cardinal process, and close-up of the koskinoid perforations developed in the ventral umbonal area and on the interarea. 4-5, RBINS a13113, incomplete articulated specimen in dorsal view and close-up of the posteriorly grooved lobes of the cardinal process. 7-13, RBINS a13114, dorsal valve in external, internal and posterior (10) views, detail of the chilidium (9), the cardinal process (11), and oblique views of the internal posterior region (12-13). 14-18, RBINS a13115, almost complete dorsal valve in external, internal and posterior views and oblique views of the internal posterior region. Scale bars equal 1 mm (1-3, 5-6, 9, 11, and 19-20), 2.5 mm (4), 5 mm (7-8, 10, 14-16, 12-13, and 17-18), 0.1 mm (21).

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FIGURE 19. Serratocrista scaldisensis sp. nov. from the Tournai area, Tournai Formation (Tournaisian). 1-4, RBINS a13116, articulated specimen, incomplete, in dorsal view (1) with detail of the extropunctae arranged radially along the axis of the costellae (2, 3), and of the basal part of the pseudodeltidium and of the teeth (4). Scale bars equal 2 mm (1), 0.5 mm (2), 0.025 mm (3), 1 mm (4).

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FIGURE 20. Lamellosathyris lamellosa (Léveillé, 1835) from the Tournai area, Tournai Formation (Tournaisian). 1-5, RBINS a13119, juvenile articulated specimen in ventral, dorsal, lateral, posterior and anterior views. 6-10, RBINS a13120, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 11-15, RBINS a5468, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 16-20, RBINS a13121, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 21-25, RBINS a13122, articulated specimen (with two microconchids attached to the right dorsal flank) in ventral, dorsal, lateral, posterior and anterior views. 26-30, RBINS a13123, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. Scale bars equal 10 mm.

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FIGURE 21. Lamellosathyris lamellosa (Léveillé, 1835) from the Tournai area, Tournai Formation (Tournaisian). 1-2, RBINS a13124, partial posterior view of an articulated specimen showing the foramen and detail of the lamellose ornamentation in the anterior part of the dorsal valve. 3-8, RBINS a13125, complete ventral valve (juvenile) in external and internal views and detail of the teeth and the dental plates. 9-11, RBINS a13126, complete ventral valve in external and internal views and detail of teeth supported by almost vertical dental plates. 12, RBINS a13127, incomplete internal mould of ventral valve. 13-18, ULg.PA.2016.12.25/8, complete dorsal valve (with two microconchids attached to its internal surface (arrows), close to the anterior margin) in external and internal view and detail of the cardinalia. 19-23, RBINS a5470, incomplete dorsal valve in internal view and detail of the cardinalia. Scale bars equal 5 mm (1-2, 3-4, 10, 11), 2 mm (5-8, 20-23), 2.5 mm (15-18), 10 mm (9, 12-14).

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FIGURE 22. Lamellosathyris lamellosa (Léveillé, 1835) from the Tournai area, Tournai Formation (Tournaisian). 1-4, RBINS a13128, almost complete dorsal valve in external and internal views, with close-up of the cardinal plate in ventral and oblique lateral views. 5-6, RBINS a13129, incomplete ventral valve with partly preserved spiralia and detail of the jugal saddle and the primary lamella. 7-10, RBINS a13130, articulated specimen with top of the ventral valve removed and details of the crura, spiralia and fimbria. 11-13, ULg.PA.2016.12.25/9, articulated specimen with ventral valve partly removed in general view and details of the top of the jugal stem and the same in oblique view. Scale bars equal 10 mm (1, 7, 11), 5 mm (2, 5), 1 mm (3-4, 10), 2 mm (6, 9), 2.5 mm (8, 12-13).

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FIGURE 23.1-7, Lamellosathyris lamellosa (Lévéillé, 1835) from Visé, Visé Formation (Visean, Warnantian). 1-5, RBINS a5471, distorted specimen in ventral, dorsal, lateral, posterior and anterior views. 6-7, RBINS a13131, incomplete ventral valve in ventral view and anterior views. 8-17, Athyris vittata de Koninck, 1887 (here placed in synonymy with Lamellosathyris lamellosa (Léveillé, 1835)) from Furfooz, Waulsort Formation (Tournaisian, Ivorian). 8-12, RBINS a5472 (lectotype) distorted specimen in ventral, lateral, dorsal, posterior, and anterior views. 13-17, RBINS a13132 (paralectotype), distorted and incomplete specimen in ventral, dorsal, posterodorsal, posterior, and lateral views. Scale bars equal 10 mm.

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FIGURE 24. Scatter diagrams of Lamellosathyris lamellosa (Léveillé, 1835). N: number of specimens measured, r2: coefficient of linear regression. 1, Relation between shell width and length. 2, Relation between shell width and shell width/shell length ratio. 3, Relation beween shell width and thickness. 4, Relation between shell width and shell thickness/shell width ratio.

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FIGURE 25. Nucleospira hannoniae nom. nov., from the Tournai area, Tournai Formation (Tournaisian). 1-5, RBINS a13134, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior,  and anterior views. 6-10, RBINS a5466 (paralectotype), articulated specimen, almost complete, in ventral, dorsal, lateral, posterior, and anterior views. 11-15, RBINS a5465 (lectotype), articulated specimen in ventral, dorsal, lateral, posterior, and anterior views. 16-21, RBINS a13135, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views, and close-up of the interarea, Lemay quarry, Vaulx Member. 22-27, RBINS a13136, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views, and close-up of the solid spines (right flank of the dorsal valve). Scale bars equal 2 mm (1-20, 22-26), 1 mm (21), 0.2 mm (21, 27).

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FIGURE 26. Nucleospira hannoniae nom. nov., from the Tournai area, Tournai Formation (Tournaisian). 1-2, RBINS a13137, distorted and incomplete specimen in dorsal view showing two whorls of the spiralium and detail of one of the cyrtomatodont teeth. 3-4, RBINS a13138, distorted articulated specimen in dorsal view displaying a circular drill hole and detail of the latter. 5, RBINS a13139, distorted articulated specimen with a circular drill hole in ventral view. Scale bars equal 2 mm (1, 3, 5), 0.5 mm (2, 4).

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FIGURE 27. Scatter diagrams of Nucleospira hannoniae nom. nov. N: number of specimens measured. 1, Relation between shell width and length. 2, Relation between width and shell length/shell width ratio. 3, Relation beween shell width and thickness. 4, Relation between shell width and shell thickness/shell width ratio.

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FIGURE 28. Coveenia ulothrix (de Koninck, 1843) from the Tournai area, Tournai Formation (Tournaisian). 1-5, 16-18, ULg.PA.2016.12.25/10, articulated specimen in ventral, dorsal, lateral, posterior and anterior views, and detail (SEM) of the foramen and the symphytium, and of circular drill hole developed close to the dorsal anterior margin. 6-10, RBINS a13140, articulated specimen in ventral, dorsal, lateral, posterior, and anterior views, Dutoit quarry, Pont-à-Rieu. 11-15, RBINS a13141, articulated specimen, almost complete, in ventral, dorsal, lateral, posterior and anterior views, Dutoit quarry, Pont-à-Rieu. Scale bar equals 10 mm (1-15), 1 mm (16-18).

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FIGURE 29. 1-4, Coveenia davidsoni (de Koninck, 1887), RBINS a5508 (holotype), articulated specimen in ventral, dorsal, lateral and posterior views, from Dréhance, Waulsort Formation (Tournaisian, Ivorian). 5-9, ‘Retzia’ intermedia de Koninck, 1887 (here considered as a rhynchonellide species), RBINS a5510 (lectotype), articulated specimen in ventral, dorsal, lateral, posterior and anterior views, from Furfooz, Waulsort Formation (Tournaisian, Ivorian). Scale bars equal 2 mm.

figure29

 

 

TABLE 1. Measurements in mm of Leptagonia franca sp. nov. Abbreviations for tables 1-4: Ddl, unrolled length of dorsal disc; Dtl, unrolled length of dorsal trail; Hi, height of the ventral interarea; L, length of the shell; T, thickness of the shell; Vdl, unrolled length of ventral disc; Vtl, unrolled length of ventral trail; W, width of the shell; Wi, width of the ventral interarea.

  W L Vdl Vtl Ddl Dtl T L/W T/W Vtl/Vdl Dtl/Ddl
N 17 17 17 17 13 13 16 17 16 17 12
Mean 51.4 28.9 23.8 22.2 21.6 8.1 17.4 0.57 0.33 0.93 0.38
MIN 36.2 24 20 8 18 2 9.3 0.48 0.26 0.36 0.10
MAX 70.8 38.1 28 32 25 15 25.8 0.70 0.40 1.28 0.67

 

TABLE 2. Measurements in mm of Serratocrista scaldisensis sp. nov. Abbreviations: see Table 1.

RBINS W L T Hi Wi
a13108 8.1 6.5 2.9 1.2 7.2
a13109 9.8 7.1 2.4 2.2 8.8
a13110 10.4 7.7 2.7 1.8 9.6
a13111 13.1 9.8 3.3 2.2 12.1
a13112 15.6 10.8 / 2.0 /
a13117 9.4 8 3.2 2.3 7.6
a13118 7.7 6.1 2.5 1.0 5.7

 

TABLE 3. Measurements in mm of Lamellosathyris lamellosa (Léveillé, 1835). Abbreviations: see Table 1.

  W L T L/W T/W T/L
N 60 60 50 60 50 50
Mean 30.1 20.5 14.5 0.70 0.49 0.70
Standard deviation 9.3122 5.1365 5.6788 0.0761 0.0667 0.1282
Standard error ±1.2021 ±0.6631 ±0.8031 ±0.0098 ±0.0094 ±0.0181
MIN 9.2 8.1 4.8 0.53 0.33 0.53
MAX 51.4 30.5 30.5 0.90 0.66 1.02

 

 

TABLE 4. Measurements in mm of Nucleospira hannoniae nom. nov. Abbreviations: see Table 1.

  W L T L/W T/W
N 18 19 16 18 16
Mean 7.9 7.5 4.6 0.94 0.59
MIN 5.7 5.5 2.7 0.85 0.46
MAX 10.1 9.5 6.0 1.04 0.69
 

author1Bernard Mottequin. Royal Belgian Institute of Natural Sciences, Operational Directorate Earth and History of Life, rue Vautier 29, B 1000 Brussels, Belgium. This email address is being protected from spambots. You need JavaScript enabled to view it.

Bernard Mottequin is a palaeontologist with a special interest in Devonian and Mississippian brachiopods (e.g., taxonomy, palaeoecology), especially those from Western Europe and North Africa, and their applications. After his PhD in sciences at the University of Liège (ULg) and the Royal Belgian Institute of Natural Sciences (RBINS, Brussels), he was postdoctoral research fellow in palaeontology at the University of Dublin, and then research assistant and honorary lecturer at the ULg, mainly involved in the modernization programme of the Geological map of southern Belgium. He is currently researcher at the RBINS.

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author2Eric Simon. Royal Belgian Institute of Natural Sciences, Operational Directorate Earth and History of Life, rue Vautier 29, B 1000 Brussels, Belgium. This email address is being protected from spambots. You need JavaScript enabled to view it.

Eric Simon made his studies and Ph. D thesis (Biology) at the Free University of Brussels. His works concern the study of fossils and extant brachiopod species. They deal with ontogeny, evolution, ecology and palaeo-ecology, morphological analyses and taxonomic problems. Improving the perception of biological characteristics in the field of paleontology is for him an important concern.