Thomasaria cf. altumbona Stainbrook, 1945

Mottequin, Bernard, 2008, New observations on Upper Devonian brachiopods from the Namur-Dinant Basin (Belgium), Geodiversitas 30 (3), pp. 455-537 : 512-534

Thomasaria cf. altumbona Stainbrook, 1945 (Figs 55-58)

Spirifer simplex – Dumon 1929: 164.

Reticularia simplex – Maillieux 1933 (e.p.): 81.

Reticularia (Eoreticularia) simplex – Maillieux 1940: 27.

Eoreticularia simplex – Maillieux 1941a (e.p.): 6.

Thomasaria altumbona Stainbrook, 1945: 58 , 59, pl. 4, figs 22-30, text-figs 2, 14.

Plectospirifer simplex – Vandercammen 1957c (e.p.): 12- 20, pl. 2, figs 7-15, 17, non pl. 2, figs 3-6, 16.

Thomasaria sp. – Mottequin 2004b: 52.

MATERIAL EXAMINED. — BM-2003-6 (one articulated specimen, one ventral valve); BM-2003-8 (six articulated specimens); BM-2003-9 (25 articulated specimens, seven FIG. 55. — Thomasaria cf. altumbona Stainbrook, 1945 , loc. BM- ventral valves); JG-1996-1 (30 articulated specimens, 2003-9,Neuville Formation: A -E, complete shell (IRScNB a12304); two ventral valves). F -J, complete shell (IRScNB a12305); A, F, ventral views; B, G, dorsal views; C, H, lateral views; D, I, posterior views; E, J, anterior views. Scale bar: 10 mm. DESCRIPTION

Shell medium-sized, wider than long(width/length ra- (1968: pl. 5, figs 5-15)]. Warrenella (W.) aquaealbae tio:1.13-1.57),ventribiconvex,rounded subpentagonal reaches a size clearly superior to the one of W. (W.) (those with an apsacline ventral area) to semi-elliptical adopecta Crickmay, 1953 and displays a wider sulcus (those with a procline ventralarea); hinge line shorter

Upper Devonian brachiopods from Belgium than greatest width (hinge line/width ratio: 0.68- bottom or corresponding to a shallow and poorly 0.91); widest at about mid-length; cardinal extremities defined groove; tongue 1.56-2.45 times wider than rounded; anterior commissure uniplicate. high, rounded to subtrapezoidal, perpendicular

Ventral valve hemipyramidal, in lateral profile, to commissural plane or bent dorsally; shoulder regularly and moderately curved or nearly flat in lines straight or concave; apical angle between 82- the case of the specimens with a procline interarea; 96°; beak straight or moderately curved; interarea flanks sloping moderately or strongly towards the 1.45-3.37 times wider than high, clearly defined, lateral commissure; sulcus originating close to beak, flat to concave, procline or pro-apsacline to cataeither well-defined, relatively wide (sulcus width/ apsacline; delthyrium partially closed by a convex width ratio: 0.35-0.57), deep with a flattened pseudodeltidium.

Mottequin B.

Dorsal valve wider than long (width/dorsal valve originating at about midvalve or close to the antelength:1.23-1.57), semi-elliptical in outline; postero- rior margin (absent in juvenile forms), rounded at lateral extremities inclined to become flat; fold top; interarea flat, orthocline, linear.

Upper Devonian brachiopods from Belgium

Flanks generally devoid of costae but up to of apical plates originating from the internal face three in some specimens; most internal costae of dental plates and dividing the post-delthyrial better defined and originating in the anterior cavity; lateral and central apical cavities poorly half of the shell (external ones only perceptible filled in. near anterior margin); growth lamellae close to Dorsal interior with ctenophoridium composed the commissure; no spine base observed (poor of more or less 10 thickened plates; dental sockets preservation). shallow with internal crests well-developed; myo-

Ventral interior with relatively long dental phragm short and slightly prominent; spiral cones plates supporting teeth of small dimensions; pair laterally oriented with at least five whorls.

Mottequin B.

DISCUSSION

These specimens are close to T. altumbona Stainbrook, 1945 on the basis of their similar size and outline as well as the development of the tongue and of the ventral area. Nevertheless, the specimens from the Namur-Dinant Basin display a generally wider sulcus (sulcus width/width ratio: 0.35- 0.57 vs 0.28-0.49 in T. altumbona ) and a convex pseudodeltidium. The latter has not been observed in the type material of Stainbrook’s species housed at the SUI.

In Belgium, these smooth spiriferids have always been misidentified with Spirifera simplex Phillips, 1841 , the type species of Pyramidalia Nalivkin, 1947 . The validity of the latter genus is not clear because it has been considered as a doubtful synonym of Cyrtinaella Frederiks, 1916 by Pitrat (1965: H678), synonym of Thomasaria by Baliński (1979: 71) or more recently, a synonym of Squamulariina Frederiks, 1916 by Carter et al. (1994: 361) and Johnson (2006: 1882). However, it was maintained as a distinct genus by Drot (1964: 78, 79) and Biernat (1966: 136). Cyrtinaella and Squamulariina have been assigned to the Cyrtinidae which implies that Spirifera simplex has a punctate shell. However, the shell structure has never been described in the English species which is very close to Spirifer pyramidalis Schnur, 1853 and Spirifer nudus Schnur, 1853 from the Eifel. A revision of Phillips’ species is urgently needed but it is beyond the scope of this work.

Vandercammen (1956: 29, pl. 1, figs 28-44, pl. 2, figs 1-6) described two Frasnian species that he assigned to Thomasaria ( T. gibbosa and T. parallela ), an opinion followed by Oleneva (2006: 421) but they have nothing in common with this genus and need to be revised.

DISTRIBUTION

Thomasaria cf. altumbona occurs in the upper part of the Neuville Formation on the southern flank of the Dinant Synclinorium. Representatives of the genus Thomasaria are known in the uppermost part

FIG. 59.— Cyrtina douvillei Rigaux,1908 ,loc.Couvin 6158p.,Grands of the Les Valisettes Formation as well as in the basal Breux Formation (Boussu-en-Fagne Member): A -E, complete shell part of the Matagne Formation, but their poor state (tral IRScNB views a; 12310 B, G,) dorsal; F -J, complete views; C shell, H, lateral (IRScNB views a12308; D, I),; A posterior, F, ven- of preservation does not permit to identify them views; E, J, anterior views. Scale bar: 10 mm. at the specific level.

0.45

Upper Devonian brachiopods from Belgium

0.65 0.85 1.15

Order SPIRIFERINIDA Ivanova, 1972 259, 260; 1933 (e.p.): 81; 1940: 27; 1941a (e.p.): 6. — Suborder CYRTINIDINA Maillieux in Asselberghs & Maillieux 1925: 166. — Dumon 1929: 164. — Brice 1988: 358, 359, pl. 43, Carter & Johnson (in Carter et al. 1994) figs 14-16.

Superfamily CYRTINOIDEA Frederiks, 1911 Family CYRTINIDAE Frederiks, 1911 MATERIAL EXAMINED. — BM-2003-6 (one articulated specimen); BM-2003-7 (three articulated specimens, one dorsal valve); BM-2003-10 (three articulated speci- Genus Cyrtina Davidson, 1858 mens, two ventral valves); Couvin 6158p. (14 articulated specimens).

TYPE SPECIES. — Calceola heteroclita Defrance, 1828 , by subsequent designation (Oehlert 1887b: 40). DESCRIPTION Shell medium- to large-sized for the genus; gen- Cyrtina douvillei Rigaux, 1908 erally wider than long (width/length ratio: 0.99- (Figs 59; 60; Table 4) 1.50), ventribiconvex; hinge line generally shorter than greatest width (hinge line width/width ratio:

Cyrtina Douvillei Rigaux, 1908: 20 , 21, pl. 1, fig. 9. — 0.85-1); cardinal extremities in some specimens

Maillieux 1909a: 10; 1909b: 120, 122, 136, 137; 1909c: mucronate.

Mottequin B.

Ventral valve hemipyramidal, more or less strongly as well as in the Frasnian part of the Lambermont curved in lateral profile; flanks sloping strongly Formation. Th ey will be described in detail when towards lateral commissure; sulcus originating at suffi cient material becomes available. beak, wide (sulcus width/width ratio: 0.53-0.59), subangular, deep, sharply delimited; tongue 1.20- 2.14 times wider than high, perpendicular to com- STRATIGRAPHIC SUMMARY missural plane or bent dorsally, subtriangular to suboval in outline; shoulder lines slightly concave to The geographic and stratigraphic distributions of subrectilinear; shoulder and apical angles identical, the brachiopod species across the Namur-Dinant between 80-97°; beak little to moderately curved, Basin are presented in Figures 61-63 . sometimes twisted; interarea triangular, very high, well-defined, pro-apsacline to apsacline; delthyrium narrow, closed by a thin pseudodeltidium pierced PALAEOGEOGRAPHIC AFFINITIES by a wide foramen. Dorsal valve wider than long (width/dorsal valve Relationships between the Namur-Dinant Basin length ratio: 1.47-1.69), semi-oval to subtrapezoidal and adjacent areas have been well known for a long in outline; fold originating at beak, high, clearly time. Maillieux (1909b) has already reported numerdelimited by two wide grooves; interarea flat, ortho- ous analogies between various fossil groups from cline, linear. the Frasnian of the Namur-Dinant Basin and the Flanks with 5 or 6 and 4 or 5 rounded to sub- Boulonnais (Northern France), which is its western angular costae on ventral and dorsal valves, respec- prolongation. More recently, similar observations tively; grooves similar but narrower; sulcus and fold have been made by Coen-Aubert (1994) on the smooth; growth lamellae prominent, sometimes Frasnian rugose corals. Several species described very crowded, irregularly spaced. in this paper are recognized in the Boulonnais or Ventral interior with short dental plates con- are close to those from that area (e.g., Athyris cf. verging towards the plane of symmetry to fuse to murchisoni ). Th e Grands Breux Formation is cor- a septum and forming a spondylium; teeth small related with the Ferques Formation by Brice et al. and rounded; tichorhinum with median partition; (1979: fig. 1), Lafuste & Tourneur (1988: fig. 3) lateral apical cavities not filled in. and Coen-Aubert (1994: table 1). More particularly, Dorsal interior with bilobed cardinal process the Boussu-en-Fagne Member corresponds to the bearing small protuberances on their internal face; two upper members of the Ferques Formation, i.e. dental socket moderately deep with internal crests the La Parisienne and Gris members. The species particularly well-developed; shelly deposits in the in common are, among others, Cyrtospirifer verposterocentral part of the valve; brachial apparatus neuili (Murchison, 1840), Athyris oehlerti Rigaux , not observed in the sectioned specimen. 1908, Douvillina dutertrei (Murchison, 1840) and

Productella subaculeata (Murchison, 1840) . On the DISCUSSION other hand, no atrypid species from the Boulonnais These specimens are identical to those identified as (Godefroid 1988) has been reported in the Frasnian Cyrtina douvillei by Brice (1988: 358, 359). units investigated in this work. The Hydrequent

Formation, which caps the Ferques Formation, DISTRIBUTION corresponds notably to the Neuville, Aisemont, Les Cyrtina douvillei is recognized within the Grands Valisettes and Matagne formations of the Namur- Breux Formation (Boussu-en-Fagne Member) on Dinant Basin, but it is poorly fossiliferous (Brice the southern border of the Dinant Synclinorium. 1988: table 2). Specimens assigned to Cyrtina and distinct from The brachiopods from the Late Frasnian and Early C. douvillei have been collected within the Neuville, Famennian from the Namur-Dinant Basin display Aisemont, Les Valisettes and Barvaux formations similarities with those reported and/or illustrated

Upper Devonian brachiopods from Belgium

(e.g., Klähn 1912; Wulff 1923; Paeckelmann 1942) athyridid species Pachyplaxoides postgyralea , which in the Aachen Basin ( Germany) which is the eastern is also recognized in the Holy Cross Mountains prolongation of the Belgian basin. More recently, ( Poland), but this taxon is absent from the Namur- Sartenaer & Hartung (1992) reported the pres- Dinant Basin. ence of Navalicria compacta Sartenaer, 1989 in the As the Frasnian was characterized by the highest Aachen Basin, a rhynchonellid characteristic of the sea-levels of the Devonian (Johnson et al. 1985) Neuville and the Aisemont formations. In the Eifel permitting exchanges between distant basins, it (Reichle quarry), the atrypid fauna collected within could be anticipated that the brachiopods investhe Ooser Plattenkalk and described by Godefroid & tigated would show also similarities at the generic Hauser (2003) is very close to that of the Neuville level with more distant areas such as Poland (e.g., Formation. Grunt (in Grunt & Racki 1998: 370, Baliński 1979, 2002), Iowa (Stainbrook 1945), figs 7, 8) described from the same locality a new Hunan (e.g., Ma et al. 2002; see also the strong

Mottequin B.

resemblance between Athyris oehlerti and A. super- Givetian), Upper Kellwasser (latest Frasnian) and vitatta Tien, 1938 illustrated by these authors) and Hangenberg events (latest Famennian). The Late the Russian platform (e.g., Liashenko 1959). Frasnian crisis, culminating during the Upper Kellwasser Event, caused the disappearance of more or less 70 to 80% of the marine species (Jablonski

CONSEQUENCES OF THE LATE 1994). It is ranked among the five most important FRASNIAN BIOLOGICAL CRISIS mass extinctions of the Phanerozoic (Sepkoski1986). ON THE BRACHIOPODS In the course of the Frasnian, the shallow waters of OF THE NAMURDINANT BASIN the platforms and continental slopes were frequently oxygen-depleted as shown by numerous occurrences

The time interval between the upper part of mid- of black shales (Algeo et al. 1995) but, according to Givetian and the Devonian/Carboniferous bound- McGhee (1996), most of them were not related to ary includes a complex sequence of events; the most extinctions and their lateral extent is not comparable signifi cant ones are the Taghanic (latest Middle to those of the Kellwasser and Hangenberg horizons.

Upper Devonian brachiopods from Belgium

However, accordingto some authors (e.g., Brett & Matagne Formation on the southern border of the Baird 1995; House 2002), these black shale horizons Dinant Synclinorium whereas in the Philippeville are related to minor extinctions but they have been Anticlinorium, it would be situated in the basal considerably less studied than those of the Kellwasser part of the Les Valisettes Formation (Bultynck et al. and Hangenberg horizons. Th e Lower and Upper 1998). On the southeastern margin of the Dinant Kellwasser events, occurring respectively within the Synclinorium, the fissile and azoic shales of the Upper Palmatolepis rhenana Zone and the P. lingui- middle part of the Les Valisettes Formation could formis Zone,correspond to two distinct rises of anoxic correspond to the LKW. Moreover, the middle part waters onto the platform (Schindler 1993) during of the intermediate shales of the Aisemont Formahighstand periods (Joachimski & Buggish 1993). tion, i.e. the dysaerobic-anaerobic unit of Poty & In the Namur-Dinant Basin, the Lower Kellwasser Chevalier (2007), could be the record of the LKW Event (LKW) is placed in the lowermost part of the according to these authors. Th e Upper Kellwasser

Mottequin B.

Event (UKW) corresponds to the Matagne Formation families ( Araksalosiidae , Monticuliferidae, Proin the Philippeville Anticlinorium (Bultynck et al. ductellidae, Sentosiidae ) recognized in the Frasnian 1998).Th e black shale horizons recognized at the top persisted in the Famennian ( Brunton et al. 2000: of the Frasnian part of the Lambermont Formation fig. 236). Nevertheless, the extinction of Devonoin the Vesdre area and on the northern margin of productinae is recorded in the Late Frasnian. In the Dinant Synclinorium have been correlated with the course of the Famennian, the Productidina the UKW by Herbosch et al. (1996). initiated an evolutionary radiation that reached a The Taghanic Event, which caused extinctions peak in the Carboniferous. Among the suborder among the ammonoids, trilobites and corals while Productidina , five species have been recognized in radiolarians and styliolinids proliferated (Aboussalam late Mid- to Late Frasnian of the Namur-Dinant 2003), is probably responsible for the disappearance Basin. Further research and additional material of stringocephalids (García-Alcalde in Brice et al. should allow precise identification of their genera 2000). Th e transgressions which occurred during and species. Early Famennian Productidina have this interval would have allowed exchanges between not been studied due to the poor preservation of various palaeobiogeographic units.Th ese would have the material available.

resulted in increased competition between the organ- Chonetidina did not suffer losses at the family isms of analogous communities with, as a corollary, level because the three families occurring in the the progressive disappearance of provincialism her- Frasnian, namely the Chonostrophiidae , Anopliidae alding the cosmopolitanism of the Frasnian faunas and Chonetidae , crossed the Frasnian-Famennian to come (Johnson 1979). Th e Frasnian was not a boundary (Racheboeuf 2000: fig. 237). In Poland major period of diversification for the atrypids (Cop- (Cracow area), a species of? Rhyssochonetes (Baliński per 1998) and the pentamerids (Godefroid in Brice 2002: 291) has been listed among the surviving taxa, et al. 2000), which became extinct at the end of the just after the Frasnian/Famennian boundary.Chone- Frasnian but this stage recorded a rapid and major tidina from the Namur-Dinant Basin have not been period of diversification of cyrtospiriferids. Among studied in this paper although they occur in all the the orders investigated, long-ranging (e.g., Athyris , investigated lithostratigraphic units. Nevertheless, Schizophoria , Warrenella ) and typical Frasnian genera Retichonetes armatus (Bouchard-Chantereaux in de (e.g., Cariniferella , Acutatheca ) have been recognized. Verneuil, 1845) was reported within the Matagne The major brachiopod turnover occurs at the top of Formation by Maillieux (1936: 18; 1941b: 7) and the P. rhenana Zone in the Namur-Dinant Basin, in Racheboeuf (1988: 399).

parallel with the deterioration of the oxygenation conditions preceding the UKW. Only an impover- ORTHIDA ished fauna has been recorded in the P. linguiformis During the Frasnian, the Orthida were represented Zone comprising only productids ( Chonetidina ), by two superfamilies, the Dalmanelloidea and Enrhynchonellids and lingulids. Post-extinction bra- teletoidea. On the four families of the Dalmanelchiopod recovery was rapid in the basal Famennian loidea, only the Rhipidomellidae persisted in the but, despite their great abundance, the brachiopod Famennian.Among the two families included in the diversity was quite low. New cosmopolitan genera Enteletoidea , only the Schizophoriidae crossed the appeared at this time especially among the spiriferids, Frasnian/Famennian boundary (Williams & Harper athyridids and rhynchonellids concomitantly with new 2000: fig. 522). According to Stigall Rode (2005: species of pre-existing orthid and orthotetid genera 164), the persistence of Schizophoria in the Famennian (Mottequin 2005b). could be explained partly by the fact that it retained deep-water lineages whereas those from the shallows PRODUCTIDA were severely affected by the Late Frasnian crisis. In The suborder Productidina does not seem to have the Namur-Dinant Basin, Schizophoria gr. striatula suffered heavy losses at the family level in relation is the last Frasnian orthid to disappear just below to the Late Frasnian mass extinction. The four the UKW or in the lowermost part of the Matagne

Upper Devonian brachiopods from Belgium

Formation although specimens may be reworked (Godefroid& Helsen 1998). Th is is also the case for material as supposed for some Spiriferida collected the other orders investigated in this paper (Mottein the same level, either at the top of the Frasnian quin 2005b). On the southern border of the Dinant part of the Lambermont Formation (northern flank Synclinorium, the last Frasnian spiriferids ( Warrenella of the Dinant Synclinorium and Vesdre area). The (W.) aquaealbae ) became extinct in the lowermost other Frasnian species described in this paper disap- part of the Matagne Formation (basal part of the pear well below the base of the Famennian. Early Upper Palmatolepis rhenana Zone ), i.e. lower than Famennian rocks yielded species of genera previously the LKW as it was defined by Bultynck et al. (1998: known ( Aulacella , Schizophoria ). 53). Nevertheless, in this area, the last spiriferids disappear generally in the upper part of the Neuville ATHYRIDIDA Formation (top of the Lower P. rhenana Zone ). The consequences of the Late Frasnian mass extinction In the Philippeville Anticlinorium (BM-2002-8), on this order have been previously discussed, notably cyrtospiriferids, Emanuella sp. as well as fragments by Baliński(1995), Rzhonsnitskaya & Modzalevskaya only identified at the generic level ( Warrenella (W.), (1996), Grunt & Racki (1998), Alvarez & Modzal- Thomasaria , Acutatheca and Adolfia ) were recovered evskaya (2001) and Alvarez (2003). In the Namur- in the upper beds of the Les Valisettes Formations Dinant Basin, Cleiothyridina davidsoni is qualified (top of the of Upper P. rhenana Zone ). In this area, as a Lazarus taxon because its last Frasnian occur- broken valves of unidentified species of Acutatheca , rence is recognized within the Neuville Formation Thomasaria and cyrtospiriferids occur in the basal (Lower Palmatolepis rhenana Zone ) at the southern part of the Matagne Formation ( P. linguiformis flank of the Dinant Synclinorium and it reappears Zone) within coquina beds (reworking?). in the basal part of the Famenne Group (“Senzeille We have only a partial view of the diversity of Formation”, Early (?)/Middle P. triangularis zones) the Frasnian Spiriferinida of southern Belgium; in the same area.The significant development of the they seem to be represented only by the cosmo- Helenathyridinae ( Biernatella and Neptunathyris ) politan genus Cyrtina , at least on the basis of the in the upper part of the Neuville Formation on the available data. Maillieux (1941a) did not report southern flank of the Dinant Synclinorium is note- it in the Famennian rocks of the Namur-Dinant worthy. Indeed, these smooth athyridids constitute Basin. In the latter, Cyrtina would re-occur in the a large part of the benthos in the distal part of the Middle Viséan (Livian regional Substage) according Namur-Dinant Basin, just before the LKW but they to the last comprehensive list of Lower Carbonibecame extinct by the end of the Frasnian.Moreover, ferous brachiopod species drawn up by Demanet Neptunathyris buxi is known also in the lower part of (1958: 96). the Matagne Formation (Upper P. rhenana Zone ). The presence of Dicamara plutonis n. sp. within the Grands Breux Formation (Boussu-en-Fagne Mem- CONCLUSIONS ber; P. hassi to Lower P. rhenana zones) correspond to one of the more recent occurrences of this genus This paper deals with some representatives of the that is reputed to disappear in the Mid-Devonian brachiopod faunas occurring in the interval spanaccording to Alvarez & Rong (2002: 1572). The ning, in terms of standard conodont zonation, the Early Famennian ( P. triangularis Zone ) is character- Palmatolepis hassi and P. triangularis zones (Late ized by the rapid expansion of the Cleiothyridininae Middle Frasnian to Early Famennian) in the Namur- ( Crinisarina ) (Mottequin 2005b). Dinant Basin ( Belgium). This area, which is the historical type area of the Frasnian and Famennian SPIRIFERIDA AND SPIRIFERINIDA stages, was located on the southeastern margin of In the Namur-Dinant Basin, the decimation of Laurussia during the Devonian and Carboniferous. spiriferid brachiopods took place in two succes- A major turnover among the brachiopods is recorded sive phases as previously reported for the Atrypida in the Upper P. rhenana Zone , just before the Upper

Mottequin B.

Kellwasser Event. During this major and widespread kollkstii devonskikh brachiopod Vostochno-Evropeydysoxic/anoxic event ( P. linguiformis Zone ), only a skoy platformy (materaly A. H. Liashenko). Byulleten’ small and poorly diversified brachiopod fauna was KF VNIGNI 2: 1-58.

ALGEO T. J., BERNER R. A., MAYNARD J. B., SCHECKLER able to develop in the widespread argillaceous en- S. E. 1995. — Late Devonian oceanic anoxic events vironments developed in the Namur-Dinant Basin. and biotic crises: “rooted” in the evolution of vascular Post-extinction brachiopod recovery occurred just land plants. GSA Today 5: 45, 64-66. after the Frasnian/Famennian boundary. New cos- ALVAREZ F. 2003. — Convergence in the evolution of mopolitan genera appeared at this time especially Palaeozoic and Mesozoic brachiopods. Journal of the

Royal Society of New Zealand 33: 189-211. among the spiriferids, athyridids and rhynchonel- ALVAREZ F. & BRUNTON C. H. C. 2005. — On the lids concomitantly with new species of pre-existing name-bearing type of Athyris concentrica (von Buch, orthid and orthotetid genera (Mottequin 2005b). At 1834). Lethaia 38: 86-87. present, only one surviving athyridid species (Lazarus ALVAREZ F., BRUNTON C. H. C. & STRUVE W. 1996. — taxon) is definitely recognized in the Early Famen- On Athyris (Brachiopoda) and its type species “ Terebratula ” concentrica von Buch. Senckenbergiana lethaea nian. Nevertheless, some productids, craniids and 76 (1/2): 65-105. lingulids may have crossed the Frasnian/Famennian ALVAREZ F. & MODZALEVSKAYA T. L. 2001. — Trends boundary but this still needs confirmation. in athyridide diversity dynamics, in BRUNTON C. H.

C., COCKS L. R. & LONG S. (eds), Brachiopods past and present. The Systematics Association Special Volume

Series 63: 212-223. Acknowledgements ALVAREZ F.&RONG JIA- YU 2002. — Order Athyridida, This paper forms part of my Ph.D. thesis supervised in KAESLER R. L. (ed.), Treatise on Invertebrate Paleontoby Jacques Godefroid (Brussels) and Édouard Poty logy, part H, Brachiopoda (revised), part 4. Geological (Liège) which was supported by the “Fonds pour la Society of America, Lawrence: 1475-1583. Formation à la Recherche dans l’Industrie et dans ALVAREZ F., RONG JIA- YU & BOUCOT A. J. 1998 —

The classification of athyridid brachiopods. Journal l’Agriculture” (FRIA) and by the Belgian Science of Paleontology 72: 827-855. Policy. Tiffany Adrain (Iowa City), Denise Brice AMSDEN T. W. 1968. — Articulate brachiopods of the St. (Lille), Jean-Claude Horrenberger (Strasbourg), Clair Limestone (Silurian), Arkansas, and the Clarita Thierry Oudoire (Lille) and Jann Thompson (Wash- Formation (Silurian), Oklahoma. The Paleontological ington D.C.) kindly gave access to collections under Society Memoir 1: 1-117.

ASSELBERGHS E. 1912. — Description d’une faune frastheir care.Marie Legrand-Blain (Gradignan) advised nienne inférieure du bord nord du bassin de Namur. on systematics of the Productidina.Wilfried Miseur Bulletin de la Société belge de Géologie, de Paléontologie (Brussels) did the photographic work. Reviews by et d’Hydrologie, Mémoires 26: 1-47. Denise Brice (Lille) and Patrick Racheboeuf (Brest) ASSELBERGHS E. 1923. — La faune de la Grauwacke de and the editorial revision by Annemarie Ohler Rouillon (base du Dévonien moyen). Mémoires du

Musée royal d’Histoire naturelle de Belgique 33: 1-76. (Paris) helped to improve the manuscript. George ASSELBERGHS E. 1936. — Le Dévonien du bord nord Sevastopulo (Dublin) advised on linguistic aspects du Bassin de Namur. Mémoires de l’Institut géologique of the paper. I am deeply grateful to all of them. de l’université de Louvain 10: 229-327.

ASSELBERGHS E. & MAILLIEUX E. 1925. — Comptes rendus de la cinquième session extraordinaire de la REFERENCES Société géologique et minéralogique de Bretagne tenue dans le terrain dévonien de l’Ardenne, entre ABOUSSALAM Z. S. 2003. — Das “Taghanic-Event” im Charleville et Gembloux du 13 au 19 avril 1925. höheren Mitteldevon von West-Europa und Marokko. Bulletin de la Société géologique et minéralogique de Münstersche Forschungen zur Geologie und Paläontolo- Bretagne 6: 117-187. gie 97: 1-332. BALIŃSKI A. 1977. — Biernatella – a new Devonian ALEKSEEVA R. E. 1969. — [New and unusual Lower double-spired brachiopod. Acta Palaeontologica Polo- Devonian spiriferids (suborder Athyrididina]. Dok- nica 22: 175-186. lady Akademii Nauk SSSR 187 (5): 1157-1159 (in BALIŃSKI A. 1979. — Brachiopods and conodonts from Russian). the Frasnian of the Dębnik Anticline, southern Poland. ALEKSEEVA R. E. & OLENEVA N. V. 2000. — Katalog Palaeontologia Polonica 39: 1-95.

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Submitted on 21 May 2007; accepted on 28 May 2008.

Mottequin B.

Figures

FIG. 61. — Distribution of the studied Frasnian brachiopods at the southern border of the Dinant Synclinorium. See Figure 62 for the stratigraphic ranges of the Famennian species from that area. The species with an asterisk has only been cited in the text. Abbreviations: CZ, conodont zones; ja., Palmatolepis jamieae Zone; PM, Petit-Mont Member.

FIG. 62. — Distribution of the studied Frasnian brachiopods in the Philippeville Anticlinorium and those of the Famennian at the southern flank of the Dinant Synclinorium. The species with an asterisk have only been cited in the text. Abbreviations: Cz, conodont zones; L., Lower; lin., Palmatolepis linguiformis Zone; Mat., Matagne Formation; PM, Petit-Mont Member; trian., P. triangularis Zone.