- Accueil
- volume 18 (2015)
- number 1
- Catalogue of the types and illustrated specimens recovered from the ‘black marble’ of Denée, a marine conservation-Lagerstätte from the Mississippian of southern Belgium
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Catalogue of the types and illustrated specimens recovered from the ‘black marble’ of Denée, a marine conservation-Lagerstätte from the Mississippian of southern Belgium
Abstract
The Viséan ‘black marble’ of Denée is one of the best preserved Mississippian fossil assemblages. This Lagerstätte is famous for the exceptional preservation of several groups of macro-organisms (fishes, echinoids, graptolites). However, only a part of the fauna has received attention from specialists and most of the phyla may benefit from a modern revision based on new investigative techniques such as 3D imagery and CT-scanning. Almost all the specimens illustrated in the literature have been traced and we present here a comprehensive catalogue of this material. Representatives of several minor groups are photographically illustrated for the first time as well as five emblematic specimens of the styracopterid genus Benedenius (Pisces). For purposes of nomenclatural formality, the lectotypes of Benedenius deneensis Traquair (in de Koninck, 1878), B. soreili Fraipont, 1890, Oligoporus soreili Fraipont, 1904, Taeniaster? fournieri Fraipont, 1904, and Scaphiocrinus longicaudatus Fraipont, 1904 are here selected.
Table des matières
1. Introduction
1The discovery of a well-preserved chondrostean fish by Hubert Thomson, a quarryman from Denée (Anhée, Namur province, Belgium) (Fig. 1), in 1866 (Van Beneden, 1867, 1871; Henrard, 1951) within the Moliniacian (Viséan) ‘black marble’ of Denée, a black coloured limestone, was the first of a long series of astonishing findings, which contributed to the fame of the locality among the palaeontological community. For example, Jackson (1929, p. 6) pointed out that ‘the Denée locality is not only remarkable for the number of genera and species of Echini found there, but also for the number of individual specimens of certain species that have been found’. Moreover, Kier (1962, p. 1) reported that the locality of Denée includes the ‘largest echinoids known, fossil or recent’. However, Soreil (1895) highlighted the scarcity of the fossils.
Figure 1. General context of Lower Carboniferous sedimentation in north-western Europe showing the distribution of emergent areas and Waulsortian mounds at the end of the Tournaisian (modified from Ziegler, 1990, and Devuyst & Dehantschutter, 2007).
2Most of the studies dedicated to the macrofauna from this particular level were published between 1871 and 1941 (e.g. Fraipont, 1904; Woodward, 1924; Van Straelen, 1926; Delépine, 1928; Fournier & Pruvost, 1928; Demanet, 1929; Jackson, 1929; Ubaghs, 1941). After Ubaghs’ publication, these faunas received scant attention from the palaeontological community despite their great scientific value (Kier, 1962; Maisey, 2008). This is partly due to the difficulties in collecting new specimens as most of the quarries have been disused since the 1940ths and/or are flooded (Henrard, 1951). Nevertheless, in the meanwhile, great progress has been made in the understanding of the palaeoenvironment and sedimentological framework of the ‘black marble’ of Denée (Mamet, 1964; Overlau, 1966; Hance, 1988; Hance et al., 1994, 2001; Lees, 1997; Poty et al., 2002; Mottequin, 2004, 2008; Devuyst et al., 2005).
3The aim of this paper is to provide for the first time a comprehensive list of the types and illustrated specimens as well as ichnofossils and unusual sedimentary structures (e.g. skip marks; see Mottequin & Poty, 2011) recovered from the ‘black marble’ of Denée and to document, if possible, their provenance and position within the sedimentary succession. Moreover, several groups of macroorganisms such as gastropods and conularids, that were known only from list of species (Fournier & Kaisin, 1929) are illustrated here for the first time in order to provide a better account of the diversity and the mode of preservation.
2. Geological and geographical setting
2.1. Lithostratigraphical and palaeoenvironmental context
4The ‘black marble’ of Denée is now included in the Molignée Formation of Early Viséan age (Moliniacian; Mississippian Foraminifer Zone MFZ 9 to MFZ 10 according to Devuyst & Hance in Poty et al., 2006) (Fig. 2). This formation (c. 60 m thick) consists of a succession of thin-bedded, commonly laminated black limestones (the typical ‘black marble’ facies) which alternate with thick-bedded, dark-grey limestones (‘thick beds’) (e.g. Mamet, 1964; Mottequin, 2004, 2008) (Figs 2-3). It developed in a confined intra-platform basin (central part of the Dinant sedimentation area [DSA]) located south of the Brabant Massif (Fig. 1), which was progressively filled by distal calcareous turbidites originating from the southward prograding shelf (e.g. Hance, 1988; Hance et al., 2001; Mottequin, 2008) (Figs 4-5). This basin was bordered to the south by a discontinuous barrier of Waulsortian mudmounds built against a major synsedimentary fault separating the DSA from the southern Avesnois sedimentation area (e.g. Hance et al., 2001; Pirotte, 2006 [see Mottequin, 2008, fig. 3]). The alternations of laminated and bioturbated lithofacies occurring within the Molignée Formation implies that the palaeoenvironment recorded several anoxic to dysoxic periods alternating with more oxygenated ones due to sea-level fluctuations of low magnitude (Mottequin, 2008). This periodic confinement of the central part of the DSA took place during a third-order sequence characterized by a low sea level, namely sequence 5 of Hance et al. (2001) (Fig. 5). Low oxygen concentrations are also suggested by the existence of organisms characteristic of dysaerobic facies such as the bivalves of the ‘paper pecten’ morphotype (e.g. Allison et al., 1995) and the remarkable preservation of the benthic and nektonic macrofauna (Mottequin, 2008).
Figure 2. Partial log of the Salet road section (stratotype of the Molignée and Salet formations) (modified from Poty et al., 2006) (see also Fig. 3). Bio. = Biostratigraphy; Chrono. = chronostratigraphy; Litho. = lithostratigraphy; MFZ = Mississippian Foraminifer Zones of Devuyst & Hance (in Poty et al., 2006). Bed numbers are those of Overlau (1966).
Figure 3. The Molignée Formation in its stratotype (Salet road section) showing the alternating thicker bedded (‘thick beds’) and thinner bedded (‘black marble’) units (see also Fig. 2). The beds are overturned and their numbering is that of Overlau (1966) (modified from Mottequin, 2008).
Figure 4. Late Tournaisian sedimentation areas in the Namur-Dinant Basin (not palinspastically restored; modified from Hance et al., 2001). ASA = Avesnois sedimentation area; CSA = Condroz sedimentation area; DSA = Dinant sedimentation area; HSA = Hainaut sedimentation area; NSA = Namur sedimentation area; VASA = Vesdre-Aachen sedimentation area; VSA = Visé sedimentation area.
Figure 5. The Condroz sedimentation area (CSA) and the Dinant sedimentation area (DSA) during the third-order sequence 5 of Hance et al. (2001) – their other sequences are represented by the black arrows (modified from Hance et al., 2001).
2.2. The quarries of the Denée area
5Most of the fossils were collected in the Denée area (Fig. 6) at the end of the 19th century and at the beginning of the 20th century by quarrymen when the ‘black marble’ was intensively and manually quarried. Most of the excavations were subterranean and some of them went down to more than 60 m in depth and were more than 100 m long (Marote, 1923; Fournier & Pruvost, 1928). The geological context of these quarries has been recently revised by Delcambre & Pingot (2004), and Delcambre & Pingot (in press). A comprehensive list of the quarries was published by Mamet (1964). Among them, the Le Cerisier, Meurisse and Piette quarries, located around the village of Denée, yielded most of the material (Fig. 6) according to the unpublished catalogue of Dom Grégoire Fournier housed at the Maredsous Abbey. The Plantis Thomas quarry can also be added to this list as it yielded a specimen of the rare styracopterid Benedenius.
Figure 6. Simplified geological map of the Denée area (modified from Delcambre & Pingot, 2004, and Delcambre & Pingot, in press) with location of the quarries that yielded most of the macrofossils (according to Fournier & Pruvost, 1928, and unpublished notes of Fournier housed at the Centre Grégoire Fournier of Maredsous Abbey).
6As is generally the case with old collections, the origin of the specimens is usually not known with precision, despite numerous pieces of information gathered by G. Fournier, except for some fossils which on their label mention the quarried level (e.g. ‘La Veine’, Les Drîs’). We can reasonably suspect that the bulk of the specimens were collected at Denée, from the exploited levels figured by Fournier (in Fournier & Pruvost, 1928) (Fig. 7).
Figure 7. Distribution of fish and some iconic invertebrates within the ‘black marble’ of Denée (Molignée Formation) as exposed in the quarries of the Denée area (modified from Fournier in Fournier & Pruvost, 1928).
3. Catalogue of the type and illustrated specimens
7The bulk of the material recovered from the ‘black marble’ of Denée was gathered by Dom Grégoire Fournier (1863-1931) (Kaisin, 1932), who amassed c. 1200 specimens from 1888 to about 1927 thanks to the expertise of several quarrymen, notably Alexis and Désiré Moreau from Denée (Fournier & Pruvost, 1928; Henrard, 1951). According to Jackson (1929, p. 72), the possession of very large echinoids such as Proterocidaris gigantea de Koninck, 1882 and the other species found in Denée, ‘should make the Musée de Maredsous (the former name of the Centre Grégoire Fournier) a Mecca for all lovers of Echini’. This invaluable collection is housed at the Maredsous Abbey (Centre Grégoire Fournier [CGF]). The collection deposited at the University of Liège (ULg) consists of specimens collected by Gustave Soreil (c. 1880) and Dom Grégoire Fournier (Jackson, 1929; Henrard, 1951). The collections of the Royal Belgian Institute of Natural Sciences (Brussels) (IRScNB) include specimens collected notably by Edouard Dupont as well as numerous plaster casts of the specimens deposited at the Centre Grégoire Fournier. Some specimens donated to R. T. Jackson by G. Fournier are curated at the Museum of Comparative Zoology of the Harvard University (MCZ).
8We here mainly focus on the types and the illustrated specimens. Particular sedimentary structures (skip marks), some of them, only known from drawings or poor photographs, are re-illustrated. We have also figured some rare, but unidentified specimens in order to show the diversity of the fauna.
9Most of the invertebrate phyla strongly need to be revised and/or studied for the first time, as the last comprehensive list of the faunas dates back to Fournier & Kaisin (1929)! Many specimens are still unidentified and rank among the incertae sedis awaiting study based on modern investigative techniques.
3.1. Fishes
10CGF 3 (Fig. 8A): Benedenius deneensis Traquair in Boulenger (1899, pl. 9) and Henrard (1951, unnumbered figure); Denée, Les Neuf Bonniers quarry.
11CGF 4 (Fig. 9A): Benedenius deneensis Traquair in Boulenger (1899, pl. 10, figs 1-4); Denée, Demaret or Meurisse quarry.
12CGF 5 (Fig. 9B): Benedenius deneensis Traquair in Boulenger (1902, without illustration); Salet, Plantis Thomas quarry.
13IRScNB P 01261 (counterpart CGF 1) (Fig. 8B): Palaeoniscus de Denée in Van Beneden (1871, pl. 4); lectotype of Benedenius deneensis Traquair, 1878 in (Traquair in de Koninck, 1878, pl. 2), Traquair (1879, pl. 3, fig. 17), Boulenger (1899, pl. 10, fig. 5), Groessens (in Dupuis et al., 1993, fig. 19.4), Groessens (1994, fig. 2), and Mottequin (2009, fig. 7A); Denée (the specimen was found in 1866, but its precise origin is unknown).
14ULg 6136 (Fig. 8C): lectotype here selected of Benedenius soreili Fraipont, 1890 in Fraipont (1890, pl. 5); Denée, Les Neuf Bonniers quarry.
15CGF 33 (not 233 as incorrectly reported by Pruvost in Fournier & Pruvost, 1928): holotype of Cratoselache pruvosti Woodward, 1924 (unnumbered plate) and Pruvost (in Fournier & Pruvost, 1928, pl. 6); Denée, Le Cerisier quarry, Les Nutons.
16CGF 34 (not 234 as incorrectly reported by Pruvost in Fournier & Pruvost, 1928): Cladodusmirabilis Agassiz in Pruvost (in Fournier & Pruvost, 1928, pl. 5, fig. 2); Denée, Les Béguines quarry, Grande Veine or Les Gahîs.
17CGF 35 (not 230 as incorrectly reported by Pruvost in Fournier & Pruvost, 1928) (Fig. 10A): ‘Helodus turgidus Agassiz’ in Pruvost (in Fournier & Pruvost, 1928, p. 22, not illustrated); Denée, Le Cerisier quarry, Les Nutons.
18CGF 36 (not 236 as incorrectly reported by Pruvost in Fournier & Pruvost, 1928) (Fig. 10B): ‘Ctenacanthus’ sp. in Pruvost (Fournier & Pruvost, 1928, p. 21, not illustrated); Denée, Grande Veine.
19CGF 40 (not 240 as incorrectly reported by Pruvost in Fournier & Pruvost, 1928, and Ivanov & Derycke, 2005): holotype of Sphenacanthus delepinei Pruvost (in Fournier & Pruvost, 1928, pl. 5, fig. 1-1a) and Ivanov & Derycke (2005, fig. 1J-P); Denée, Meurisse quarry, Les Drîs.
20Remarks. According to Fournier & Pruvost (1928), most of the specimens (23) assigned to Denaea fournieri were collected before 1890 at the Le Cerisier quarry (Les Nutons) but eleven additional ones were recovered from the Meurisse quarry (Les Drîs) after the publication of Fournier & Pruvost (1922).
21CGF 201 (Fig. 10C): holotype of Denaea fournieri Pruvost (in Fournier & Pruvost, 1922) in Fournier & Pruvost (1928, pl. 1, pl. 4, figs 2, 6, 7, 9, 10), Ivanov & Derycke (2005, fig. 1A-I), and Maisey (2008, fig. 2A); Denée.
22CGF 202: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 2, fig. 2), Maisey (2008, fig. 2F); Denée.
23CGF 203: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 4, fig. 5, 11, pl. 5, fig. 3); Denée.
24CGF 205: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 2, fig. 1); Denée.
25CGF 209: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 5, fig. 4); Denée.
26CGF 212: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 4, fig. 4); Denée.
27CGF 214: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 4, fig. 3); Denée.
28CGF 215: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 3, fig. 2); Denée.
29CGF 216: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 4, fig. 8); Denée.
30CGF 217: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 4, fig. 1); Denée.
31CGF 218: D. fournieri Pruvost in Fournier & Pruvost (1928, pl. 3, fig. 1); Denée.
32CGF 219: D. fournieri Pruvost in Maisey (2008, fig. 2B); Denée.
33CGF 208: D. fournieri Pruvost in Maisey (2008, fig. 2D); Denée.
34CGF 224: D. fournieri Pruvost in Maisey (2008, fig. 2E); Denée.
Figure 8. A-B. Benedenius deneensis Traquair (in de Koninck, 1878), Molignée Formation. A. CGF 3, Denée, Les Neuf Bonniers quarry. B. Lectotype, IRScNB P 01261, Denée, accurate origin unknown. C. Benedenius soreili Fraipont, 1890, lectotype, ULg 6136, Denée, Les Neufs Bonniers quarry. Scale bars: 10 mm.
Figure 9. A-B. Benedenius deneensis Traquair (in de Koninck, 1878), Molignée Formation. A. CGF 4, Denée, Meurisse quarry. B. CGF 5, Salet, Plantis Thomas quarry. Scale bars: 10 mm.
Figure 10.A. Helodus turgidus Agassiz fide Pruvost (in Fournier & Pruvost, 1928), Denée, Le Cerisier quarry, Les Nutons. B. ‘Ctenacanthus’ sp., Denée, Grande Veine. C. Denaea fournieri Pruvost (in Fournier & Pruvost, 1922), holotype, CGF 201, Denée, most probably from Le Cerisier quarry, Les Drîs.Scale bars: 10 mm, except A (1 mm).
3.2. Echinoderms
3.2.1. Crinoids
35CGF 44 (Fig. 11C): Scaphiocrinus longicaudatus Fraipont, 1904; Denée, Le Cerisier quarry, Les Nutons.
36CGF 49 (Fig. 11B): undetermined crinoid; Denée, Grande Veine.
37CGF 51 (Fig. 13B): undetermined crinoid; Denée.
38CGF 191 (Fig. 11A, D): Denée, Meurisse quarry, La Veine.
39ULg 11,219: lectotype (here selected) of Scaphiocrinus longicaudatus Fraipont, 1904 in Fraipont (1904, pl. 1, fig. 1); Graphiocrinus longicaudatus in Mottequin (2009, fig. 6F); Denée.
40ULg 11,220: Scaphiocrinus longicaudatus Fraipont, 1904 in Fraipont (1904, pl. 1, fig. 2); Denée, Minet quarry, Les Nutons.
Figure 11. Crinoids from the Molignée Formation. A, D. Complete undetermined crinoids with close-up of the holdfast, Denée, Meurisse quarry, La Veine, CGF 191. B. Undetermined crinoid, Denée, Grande Veine, CGF 49. C. Scaphiocrinus longicaudatus Fraipont, 1904, CGF 44, Denée, Le Cerisier quarry, Les Nutons. Scale bars: 10 mm.
3.2.2. Echinoids
41CGF 69: Archaeocidaris wervekei Tornsquist in Fraipont (1904, pl. 1, fig. 7); Denée.
42CGF 70: Archaeocidaris wervekei Tornsquist in Fraipont (1904, pl. 1, fig. 6); Denée.
43CGF 71: paratype of Perischodomus fraiponti Jackson, 1929 in Jackson (1929, pl. 5, fig. 4); Proterocidaris gigantea de Koninck, 1882 in Kier (1962, pp. 3, 8); Denée, Meurisse quarry, Les Drîs.
44CGF 72: Rhoechinus elegans (M’Coy) in Fraipont (1904, pl. 2, fig. 9); Palaeechinus elegans M’Coy in Jackson (1929, pl. 2, fig. 17); Proterocidaris gigantea de Koninck, 1882 in Kier (1962, pl. 4, fig. B); Denée.
45CGF 73: Palaeechinus elegans M’Coy in Jackson (1929, pl. 2, fig. 18); Denée.
46CGF 74: Palaechinus lacazei Julien in Fraipont (1904, pl. 2, fig. 4); Denée, Grande Veine.
47CGF 75: Palaechinus lacazei Julien (?) in Fraipont (1904, pl. 2, fig. 3); Denée, Grande Veine.
48CGF 76: Palaechinus lacazei Julien (?) in Fraipont (1904, pl. 1, fig. 8); Denée, Grande Veine.
49CGF 78: holotype of Perischodomus fraiponti Jackson, 1929 in Jackson (1929, pl. 5, fig. 3) and Henrard (1951, unnumbered figure); Proterocidaris gigantea de Koninck, 1882 in Kier (1962, pp. 3, 8); Denée, Piette quarry, Grande Veine.
50CGF 81: Palaechinus lacazei Julien (?) in Fraipont (1904, pl. 2, fig. 6); Perischodomus fraiponti Jackson, 1929 in Jackson (1929, p. 50); Denée.
51CGF 94: Maccoya sphaerica (M’Coy) in Jackson (1929, pl. 3, fig. 1); Denée.
52CGF 100: Archaeocidaris wervekei Tornsquist in Jackson (1929, pl. 1, fig. 7); Denée.
53CGF 101: holotype of Lepidechinus belgicus Jackson, 1929 in Jackson (1929, pl. 5, fig. 1); Denée.
54CGF 123: Lovenechinus lacazei (Julien) in Jackson (1929, pl. 3, fig. 4); Palaeoechinus elegans in Groessens (1994, pl. 7, unnumbered figure); Denée, Meurisse quarry.
55CGF 125: Oligoporus soreili Fraipont, 1904 in Fraipont (1904, pl. 4, fig. 2); Proterocidaris giganteus de Koninck, 1882 in Jackson (1912, pl. 65, fig. 3, pl. 67, figs 6-7); P. gigantea in Jackson (1929, text-fig. 8); Denée.
56CGF 127: Proterocidaris gigantea de Koninck, 1882 in Jackson (1929, pl. 7), Mortensen (1935, p. 71, fig. 43), Groessens (in Dupuis et al., 1993, fig. 19.3 [?]), and Groessens (1994, pl. 6); Denée.
57CGF 138: Lovenechinus anglicus Jackson, 1929 in Jackson (1929, pl. 4, fig. 1); Denée.
58CGF 140: holotype of Fournierechinus deneensis Jackson, 1929 in Jackson (1929, pl. 9); Denée, probably from Les Drîs.
59CGF 141: paratype of Fournierechinus deneensis Jackson, 1929 in Jackson (1929, pl. 10, fig. 2) and Mortensen (1935, p. 72, fig. 44); Denée.
60CGF 142: paratype of Fournierechinus deneensis Jackson, 1929 in Jackson (1929, pl. 10, fig. 1) and Mottequin (2009, fig. 6D); Proterocidaris deneensis in Kier (1962, text-figs 5-6, pls 5-6); Denée.
61CGF 147: holotype of Deneechinus tenuispinus Jackson, 1929 in Jackson (1929, pl. 1, fig. 11) and Mortensen (1935, p. 63, fig. 34); Denée, Piette quarry, Grande Veine.
62CGF 151: Proterocidaris gigantea de Koninck, 1882 in Jackson (1929, pl. 8, fig. 1); Denée.
63IRScNB a7386: paratype of Lepidechinus belgicus Jackson, 1929 in Jackson (1929, pl. 5, fig. 2); Denée, Piette quarry.
64IRScNB a7388: Proterocidaris gigantea de Koninck, 1882 in Jackson (1929, pl. 8, fig. 2); Denée, Piette quarry.
65IRScNB a7383: Lovenechinus lacazei (Julien) in Jackson (1929, pl. 3, figs 2-3) and Maillieux (1933, pl. 9, fig. 132); Denée.
66MCZ 3330: Proterocidaris gigantea de Koninck, 1882 in Kier (1962, text-figs 1-2, pls 1-2) and Kier (1966, fig. 224.1a, b); Denée.
67ULg 11,221 (not traced!): Archaeocidaris urii (Fleming) in Fraipont (1904, pl. 1, fig. 5).
68ULg 11,222: Palaechinus lacazei Julien in Fraipont (1904, pl. 2, fig. 2); Denée.
69ULg 11,225: Palaechinus lacazei Julien in Fraipont (1904, pl. 2, fig. 5); Denée.
70ULg 11,226: Palaechinus sp. in Fraipont (1904, pl. 2, fig. 8); Lovenechinus lacazei (Julien) in Kier (1962, text-figs 7-9, pl. 4, fig. A); Denée.
71ULg 11,227: Palaechinus konincki Julien in Fraipont (1904, pl. 3, fig. 1); Palaeechinus? regnyiensis Jackson, 1929 (? proP. regnyi Jackson, 1912) in Jackson (1929, p. 38); Denée.
72ULg 11,228 (Fig. 12A): lectotype (here selected) of Oligoporus soreili Fraipont, 1904 in Fraipont (1904, pl. 3, fig. 2); Denée.
73ULg 11,230: Oligoporus soreili Fraipont, 1904 in Fraipont (1904, pl. 5, fig. 1); Proterocidaris giganteus de Koninck, 1882 in Jackson (1912, pl. 67, fig. 4); P. gigantea in Jackson (1929, text-fig. 9); Denée.
74ULg 11,231: Oligoporus soreili Fraipont, 1904 in Fraipont (1904, pl. 5, fig. 2); Proterocidaris giganteus de Koninck, 1882 in Jackson (1912, pl. 67, fig. 5); P. gigantea in Jackson (1929, text-fig. 10); Denée.
75ULg 11,239: Oligoporus soreili Fraipont, 1904 in Fraipont (1904, pl. 4, fig. 1); Denée.
76ULg 11,541: Palaechinus sp. in Fraipont (1904, pl. 2, fig. 7); Denée.
77ULg 35,400: Proterocidaris gigantea de Koninck, 1882 in Kier (1962, text-fig. 3, pl. 3) and Mottequin (2009, fig. 7C); Denée.
78ULg 2014-02-10/1: Palaechinus lacazei Julien in Fraipont (1904, pl. 2, fig. 1); Denée.
Figure 12. Echinoderms from the Molignée Formation. A. Proterocidaris gigantea de Koninck, 1882, ULg 11,228, lectotype of Oligoporus soreili Fraipont, 1904. B-C. Taeniaster? fournieri Fraipont, 1904, Denée, Les Neuf Bonniers quarry. B. CGF 68. C. CGF 67, lectotype, the arrow indicates the presence of a trace left by the animal before its death. D. undertermined ophiuroid, ULg 30,795. Scale bars: 10 mm.
3.2.3. Ophiuroids
79CGF 67 (Fig. 12C): lectotype (here selected) of Taeniaster? fournieri Fraipont, 1904 in Fraipont (1904, pl. 1, fig. 4), Groessens (1994, pl. 7, unnumbered figure), and Mottequin (2009, fig. 6E); Denée, Les Neuf Bonniers quarry.
80CGF 68 (Fig. 12B): Taeniaster? fournieri Fraipont, 1904 in Fraipont (1904, pl. 1, fig. 3); Denée, Les Neuf Bonniers quarry.
81ULg 30,795 (Fig. 12D): undetermined ophiuroid; Denée.
3.3. Brachiopods
82CGF 51 (Fig. 13B): unidentified productide; Denée.
83CGF 307: Productus mesolobus (Phillips) in Delépine (1928, pl. 3, fig. 25); Productus semireticulatus in Groessens (1994, pl. 7, unnumbered figure); Salet, La Veine (?)
84CGF 320: Productus mesolobus (Phillips) in Delépine (1928, pl. 3, fig. 24); Denée.
85CGF 322: Productus mesolobus (Phillips) in Delépine (1928, pl. 3, fig. 23); Denée.
86CGF 329: Productus semiréticulé (sic) in Delépine (1928, pl. 2, fig. 17); Denée.
87CGF 331: Productus (Pustula) interruptus Thomas in Delépine (1928, pl. 3, fig. 32); Denée.
88CGF 336: Productus (Pustula) interruptus Thomas in Delépine (1928, pl. 3, fig. 30); Denée, Minet quarry, Les Berwettis.
89CGF 341: Productus pyxidiformis de Koninck in Delépine (1928, pl. 6, fig. 70); Denée.
90CGF 350: Productus semireticulatus (Martin) forme crassispinus in Delépine (1928, pl. 3, fig. 21); Productus semireticulatus in Maillieux (1933, pl. 8, fig. 133); Salet.
91CGF 373: Productus corrugatus M’Coy in Delépine (1928, pl. 6, fig. 68); Denée.
92CGF 386: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 42); Denée.
93CGF 406: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 38); Denée.
94CGF 408: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 41); Denée.
95CGF 409: Productus semireticulatus (Martin) forme crassispinus Delépine, 1928 in Delépine (1928, pl. 2, fig. 13) and Groessens (in Dupuis et al., fig. 19.2); Denée.
96CGF 410: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 39); Denée.
97CGF 411: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 40); Denée.
98CGF 412: Productus corrugatus M’Coy in Delépine (1928, pl. 4, fig. 46); Denée.
99CGF 417: Productus corrugatus M’Coy in Delépine (1928, pl. 4, fig. 45); Denée.
100CGF 431: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 44); Denée.
101CGF 446: Productus cora d’Orbigny in Delépine (1928, pl. 4, fig. 43); Denée.
102CGF 470: Spirifer bisulcatus Sowerby in Delépine (1928, pl. 6, fig. 75); Denée.
103CGF 472: Productus concinnus Sowerby in Delépine (1928, pl. 2, fig. 12); Denée, Grande Veine.
104CGF 522: Productus aff. semireticulatus (Martin) forme ramispinus Delépine, 1928 in Delépine (1928, pl. 1, fig. 1); Denée.
105CGF 523 (Fig. 13A): Productus aff. semireticulatus (Martin) (? forme ramispinus) in Delépine (1928, pl. 2, fig. 15); Denée.
106CGF 525: Productus aff. semireticulatus (Martin) forme ramispinus in Delépine (1928, pl. 2, fig. 16); Denée.
107CGF 528: Productus aff. semireticulatus (Martin) forme ramispinus Delépine, 1928 in Delépine (1928, pl. 1, fig. 2) and Groessens (in Dupuis et al., 1993, fig. 19.1), unidentified productid brachiopods in Mottequin (2009, fig. 7K); Denée.
Figure 13. Brachiopods, crinoids and graptolite from the Molignée Formation in Denée. A. Undertermined productide with long ventral spines exceptionally preserved, CGF 523. B. Partly articulated undetermined crinoids and productide with ventral spines preserved, CGF 51. C. Ptiograptus fournieri Ubaghs, 1941, holotype, CGF 702. Scale bars: 10 mm.
3.4. Molluscs
3.4.1. Cephalopods
108CGF (not traced): Nomismoceras vittigerum (Phillips) in Delépine (1940, pl. 5, fig. 29); Denée.
109CGF 252 (Fig. 14C): undetermined goniatite; Denée, Piette quarry, Grande Veine.
110CGF 262 (Fig. 14D): undetermined orthoconic cephalopod; Denée.
111CGF 274 (Fig. 14F): undetermined orthoconic cephalopod; Denée, Grande Veine.
112CGF 278 (Fig. 14B): undetermined goniatite; Denée.
113CGF 284 (Fig. 14E): undetermined goniatite; Denée, Grande Veine.
114CGF 289 (Fig. 14A): undetermined goniatite; Denée.
115CGF 603: Nomismoceras frechi Schmidt in Delépine (1940, pl. 5, fig. 32); Denée.
Figure 14. A-H. Molluscs from the Molignée Formation in Denée. A-C, E. Undetermined goniatite. A. CGF 289. B. CGF 278. C. CGF 252, Piette quarry, Grande Veine. E. CGF 284, Grande Veine. D, F. Undetermined orthoconic cephalopods. D. CGF 262. F. CGF 274, Grande Veine. G. ‘Pterinopecten’ radiatus (Phillips, 1836), left valve, CGF 652. H. Undetermined gastropod, CGF 614. Scale bars: 10 mm.
3.4.2. Pelecypods
116CGF 646: Pterinopecten dumontianus (de Koninck) in Demanet (1929, pl. 2, fig. 8); ‘Pterinopecten’ dumontianus (de Koninck) in Mottequin (2009, fig. 7K [right]); Denée.
117CGF 649: Pterinopecten dumontianus (de Koninck) in Demanet (1929, pl. 2, fig. 10); Denée.
118CGF 652 (Fig. 14G): Pterinopecten radiatus (Phillips) in Demanet (1929, pl. 2, fig. 13); Denée.
119CGF 654: Pterinopecten radiatus (Phillips) in Demanet (1929, pl. 2, fig. 12); Denée.
120CGF 660: Pterinopecten dumontianus (de Koninck) in Demanet (1929, pl. 2, fig. 9); ‘Pterinopecten’ dumontianus (de Koninck) in Mottequin (2009, fig. 7K [left]); Denée.
121CGF 680: Aviculopecten stellaris (Phillips) in Demanet (1929, pl. 2, fig. 6); Denée.
122CGF 682: Pterinopecten radiatus (Phillips) in Demanet (1929, pl. 2, fig. 14); Denée.
123CGF 686: Aviculopecten stellaris (Phillips) in Demanet (1929, pl. 2, fig. 7); Denée.
124IRScNB a5773: Pterinopecten dumontianus (de Koninck) in Demanet (1929, pl. 2, fig. 11); Denée, Meurisse quarry.
125ULg 11,244: Pseudamusium anisotum (Phillips) in Demanet (1929, pl. 2, fig. 16); Denée.
126ULg 2014-02-03/1: slab with a ‘paper pecten’ in Mottequin (2004, pl. 4, fig. C); Maredret-Station (see Mottequin, 2004, fig. 6).
3.4.3. Gastropods
127CGF 614 (Fig. 14H): undetermined gastropod; Denée.
3.5. Corals
128CGF 1021: Caninophyllum sp. (det. E. Poty) in Mottequin (2000, pl. 12, fig. 1); Denée.
129ULg 2014-02-10/2: Caninophyllum sp. (det. E. Poty) in Mottequin (2000, pl. 12, fig. 1, 2004, pl. 3, fig. C, 2009, fig. 7I); Denée.
130ULg 2014-02-10/3: ‘Caninia du type patula’ in Salée (1911, fig. 1); Denée.
131Not traced: ‘Caninia du type patula’ in Salée (1911, fig. 2); Denée.
3.6. Conularids
132CGF 633 (Fig. 15A): Denée, Les Cerisiers quarry.
Figure 15. A. Undetermined conularid, CGF 633, Denée, Les Cerisiers quarry. B-D. Incertae sedis. B. this enigmatic fossil was tentatively identified as a holothurian by Kaisin (1926), CGF 759, Denée. C. Incertae sedis, CGF 801, Denée. D. Incertae sedis, CGF 763, Denée. E. Repichnia, CGF 778, Denée. F. Skip marks probably made by a gastropod shell, CGF 847, Denée. Scale bars: 10 mm.
3.7. Sponges
133ULg 2014-02-03/2: slab with ‘paper pecten’ in Mottequin (2004, pl. 4, fig. C) with a sponge showing long spicules, close to representatives of the genus Belemnospongia(Mottequin, 2008, p. 201); Maredret-Station (see Mottequin, 2004, fig. 6).
134CGF 1100: sponge (?) in Mottequin (2000, pl. 13, fig. 2); Denée, Meurisse quarry, top of La Veine.
135CGF 819 (Fig. 16A): cf. Phyllothallus sp. in Mottequin (2000, p. 13, fig. 1); this may be a non-rigid sponge similar to that illustrated by Dietl & Schweigert (2004, pl. 2, fig. 1); Denée, Meurisse quarry.
Figure 16.A. Probably a non-rigid sponge, CGF 8198, Denée, Meurisse quarry. B. Undertermined plant axis, Denée, Meurisse quarry, Les Drîs. C. Undetermined spermatophyte fertile branching system, CGF 770, Denée, La Veine. D. Most probably a non-calcareous alga, CGF 771, Denée, La Veine. Scale bars: 10 mm.
3.8. Graptolites
136CGF 701: ‘Dendrograptus’ sp. 1 in Ubaghs (1941, pl. 2, fig. 9); Denée.
137CGF 702 (Fig. 13C): holotype of Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 5, fig. 18) and Henrard (1951, unnumbered figure); Denée.
138CGF 704: ‘Dendrograptus’ sp. 2 in Ubaghs (1941, pl. 1, fig. 4); Denée, Les Drîs.
139CGF 706: paratype of Dictyonema ultimum in Ubaghs (1941, pl. 3, fig. 11); Denée, La Grande Veine.
140CGF 708: holotype of Dictyonema ultimum Ubaghs, 1941 in Ubaghs (1941, pl. 3, fig. 10); Denée.
141CGF 710: paratype of Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 2, fig. 8); Denée.
142CGF 714: Dictyonema fraiponti Ubaghs, 1941 in Ubaghs (1941, pl. 1, fig. 1), Groessens (in Dupuis et al., 1993, fig. 19.5) and Mottequin (2009, fig. 6H); Denée, Piette quarry, La Veine.
143CGF 724: holotype of Dictyonema fraiponti Ubaghs, 1941 in Ubaghs (1941, pl. 4, fig. 16); Denée.
144CGF 725: holotype of Desmograptus crassus Ubaghs, 1941 in Ubaghs (1941, pl. 3, fig. 14); Denée, Meurisse quarry, Les Drîs.
145CGF 1192: Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 5, fig. 19); Denée.
146IRScNB a7671: Dictyonema fraiponti Ubaghs, 1941 in Ubaghs (1941, pl. 4, fig. 15); Denée, Piette quarry, La Veine.
147IRScNB a7672A: Dictyonema fraiponti Ubaghs, 1941 in Ubaghs (1941, pl. 4, fig. 17); Denée, Meurisse quarry.
148IRScNB a7672B: Desmograptus sp. in Ubaghs (1941, pl. 5, fig. 20); Denée.
149ULg 11,280: paratype of Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 1, fig. 2); Denée.
150ULg 11,281: paratype of Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 2, fig. 7), Groessens (in Dupuis et al., fig. 19.6), Groessens (1994, fig. 1), and Mottequin (2009, fig. 7G); Denée.
151ULg 11,282: Ptiograptus fournieri Ubaghs, 1941 in Ubaghs (1941, pl. 1 fig. 3); Denée.
3.9. Incertae sedis
152CGF 759 (Fig. 15B): holothurian (?) (see Kaisin, 1926) in Mottequin (2000, pl. 11, fig. 1); Denée.
153CGF 1120: crustacean (?) in Mottequin (2000, pl. 11, fig. 2); Denée.
154CGF 763 (Fig. 15D): incertae sedis (holothurian?) in Mottequin (2000, pl. 12, fig. 3, 2004, pl. 4, fig. B); Denée.
155CGF 801 (Fig. 15C): incertae sedis in Mottequin (2000, p. 13, fig. 3); Denée.
156CGF 798: worm (?) in Mottequin (2000, pl. 14, fig. 1); Denée.
157CGF 873: worm (?) in Mottequin (2000, pl. 14, fig. 2); Denée.
3.10. Plants
158CGF 769 (Fig. 16B): This fossil consists of a dichotomous axis showing clear longitudinal striation. No other characteristic feature could be observed. It has been determined as Archaeosigillaria by Renier (unpublished data). The latter genus has recently been revised by Berry and Edwards (1997). It is characterized by lycopodiaceous stems covered with spirally arranged hexagonal leaf bases. None of these characters are present in the present plant fragment. We think this axis is impossible to determine with precision. Denée, Meurisse quarry, Les Drîs.
159CGF 770 (Fig. 16C):This fossil has been identified as Tristichia? sp. (det. M. Fairon-Demaret) in Mottequin (2000, pl. 12, fig. 4, 2004, pl. 3, fig. D). The genus name Tristichia corresponds to anatomically preserved three-ribbed protosteles presenting papilionoid leaf traces (Long, 1961). The lack of anatomical details in the present material thus invalidates this identification. The occurrence of a bidimensional dichotomous branching system distally bearing cupulate structures is however reminiscent of earliest seed plant organisation but a more detailed analysis is necessary in order to clarify its affinities. We will thus refer to this material as ‘undetermined spermatophyte fertile branching system’. Denée, La Veine.
160CGF 771 (Fig. 16D): This fossil consists of a long slender axe that all seem to arise from the same zone. The branching pattern is not very clear. Considering the morphology of the specimen, it very likely corresponds to a fossil non calcareous alga. Further investigations are necessary to clarify its affinities. Denée, La Veine.
3.11. Ichnofossils
161CGF 754: Medusina boulengeri Van Straelen, 1926 in Van Stralen (1926, figs 1-2) and Groessens (1994, pl. 7, unnumbered figure); ‘Medusina’ boulangeri (sic) Van Straelen in Harrington & Moore (1956, fig. 129.6) and Groessens (in Dupuis et al., 1993, fig. 19.7a-19.7b). This specimen is probably a burrow (for discussion, see Mottequin, 2004, p. 5); Denée, Piette quarry.
162CGF 773: ‘empreinte néréitiforme’ in Carpentier (1913, pl. 11, fig. 1); Denée.
163CGF 778 (Fig. 15E): repichnia, Denée.
164CGF 787: ‘empreinte néréitiforme’ in Carpentier (1913, pl. 11, fig. 2); Denée.
165CGF 887: Zoophycos sp. in Mottequin (2000, pl. 17, fig. 1); Denée.
166CGF 1117: repichnia in Mottequin (2000, pl. 15, fig. 1); Denée, Meurisse quarry, Les Drîs.
167ULg 30,016: ‘empreinte néréitiforme’ in Fraipont (1911, pl. 3); Denée.
168ULg 30,088: pascichnia in Mottequin (2000, pl. 16, fig. 1); Denée.
3.12. Skip and groove marks
169CGF 825: groove marks in Mottequin (2000, pl. 22, fig. 1); Denée, Piette quarry.
170CGF 847 (Fig. 15F): skip marks probably made by a gastropod shell in Mottequin (2000, pl. 20, fig. 1); Denée.
171CGF 906: skip mark moulds in Mottequin (2000, pl. 21, fig. 1); Denée, Meurisse quarry.
172CGF 1132: skip mark moulds produced by a broken cephalopod shell in Mottequin (2000, pl. 19, fig. 1, 2004, pl. 4, fig. A); Denée, Meurisse quarry, La Veine.
173ULg 30,082: skip mark moulds in Mottequin (2000, pl. 18, fig. 1); Denée.
4. Perspectives
174This catalogue rather than representing the last word constitutes the first step of a complete reassessment of the Denée “black marble” diverse fauna. This assemblage constitutes an invaluable window on Mississippian environments. The occurrence in the same beds of exquisitely preserved vertebrates and invertebrates as well as the presence of ichnofossils and indications of soft bodied organisms are in our opinion crucial to the reconstruction of past ecological conditions.
5. Acknowledgments
175We are greatly indebted to Thierry Hubin (Royal Belgian Institute of Natural Sciences) for taking the photographs, to Paul Van Genabeek for providing access to the collections curated at the Centre Grégoire Fournier of the Maredsous Abbey, and to Luc Hance (Carmeuse s.a.) and George Sevastopulo (Trinity College) for reviewing the manuscript.
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236Manuscript received 10.04.2014, accepted in revised form 02.09.2014, available on line 20.10.2014.