geol. mag. 152 6 , 2015, pp. 1009–1024. doi:10.1017

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Geol. Mag. 152 (6 ), 2015, pp. 1009–1024. c© Cambridge University Press 2015 1009doi:10.1017/S0016756814000697

Famennian rhynchonellides (Brachiopoda) from deep-water faciesof the Ougarta Basin (Saoura Valley, Algeria)

B E R NA R D M OT T E Q U I N†∗, FAT I M A Z O H R A M A LT I‡, M A DA N I B E N YO U C E F § ,C AT H E R I N E C R Ô N I E R ¶, L O U I S A S A M A R║, C A R I N E R A N D O N∗∗

& D E N I S E B R I C E†††Institut Royal des Sciences Naturelles de Belgique, D. O. Terre et Histoire de la Vie, rue Vautier 29, B 1000 Brussels,

Belgium‡Université de Béchar, Faculté des Sciences et de la Technologie, BP 417, Béchar 08000, Algeria§Université de Mascara, Département des Sciences de la Terre, BP 305, Mascara 29000, Algeria

¶FRE 3298, Géosystèmes, Université Lille 1, Sciences de la Terre, 59655 Villeneuve d’Ascq Cedex, France║SONATRACH, Centre de Recherche et Développement, Boumerdès 35000, Algeria∗∗UMR 7207 CR2P Centre de Recherche sur la Paléobiodiversité et les Paléoenvironnements, Université Pierre et

Marie Curie, Laboratoire de Micropaléontologie, 75252 Paris Cedex 05, France††Université Catholique de Lille et Groupe ISA 48 boulevard Vauban, 59046 Lille Cedex, France

(Received 9 April 2014; accepted 24 October 2014; first published online 10 April 2015)

Abstract – In the Saoura Valley (Ougarta Basin, Saharan Algeria), the lower–upper Famennian part ofthe essentially shally Marhouma Formation is characterized by deep-water facies and includes horizonsrich in ammonoids (goniatites and clymeniids) and blind to eye-reduced phacopide trilobites. They arealso rich in small-sized and smooth rhynchonellide brachiopods, investigated here for the first time inorder to detail their post-Kellwasser recovery. Rhynchonellides clearly predominate in the brachiopodassemblages (representing 90 % of the whole assemblage, with 10 species) composed otherwiseof athyridides, orthides and spiriferides. Rhynchonellides are mostly represented by relatively flatleiorhynchids and rozmanariids consistent with poor oxygenation on the sea floor. One new speciesis described (Evanidisinurostrum saouraense sp. nov.); four genera, previously known only fromthe south-eastern margin of Laurussia, are reported for the first time from the northern margin ofGondwana: the leiorhynchid Sphaeridiorhynchus and the rozmanariids Leptoterorhynchus, Pugnariaand Novaplatirostrum.

Keywords: Upper Devonian, Marhouma Formation, North Gondwana, brachiopods, Rhynchonellida,palaeoecology.

1. Introduction

Upper Devonian deposits were first recognized inthe Saoura Valley (Algeria, Ougarta Basin) by Haug(1903) on the basis of cephalopods (goniatites andclymeniids), which are particularly abundant in somelevels (see also Menchikoff, 1930). The first compre-hensive study on the geology of this area was by Gau-tier (1906) and refined by Alimen et al. (1952), whodescribed several important sections. The richness inammonoids enabled Petter (1959) and Göddertz (1987)to discriminate the zonation established by Wedekind(1908, 1926) in Germany and subsequently modifiedby several authors (e.g. Korn, Klug & Reisdorf, 2000;Korn, 2004).

Lower and Middle Devonian brachiopod faunas fromthe Ougarta were first described by Le Maître (1952)and revised in part by Boumendjel et al. (1997), OualiMehadji et al. (2004), and Brice et al. (2011) but, un-til recently, Upper Devonian brachiopods were poorlyknown (e.g. Brice, Legrand-Blain & Nicollin, 2005,

∗Author for correspondence: [email protected]

2007). Brachiopods are of interest because they flour-ished along the northern margin of Gondwana in neriticfacies sometimes unfavourable to ammonoids and con-odonts, and are therefore valuable tools for biostrati-graphy in such palaeoenvironmental contexts, espe-cially regarding the Devonian–Carboniferous boundary(Mottequin, Brice & Legrand-Blain, 2014).

This paper is the first comprehensive study of thediverse and small rhynchonellide brachiopods fromthe Famennian (Marhouma Formation) Saoura Valleywhere they are the dominant element in the brachiopodassemblages, although they also occur in benthic com-munities (cf. Crônier et al. 2013 for trilobites). Theirdiversity and palaeoecology are also discussed.

2. Geological setting and material

The material described in this paper comes fromthe Saoura Valley in the eastern part of the Ou-garta Basin (Algerian Sahara), about 350 km SSW ofBéchar (Fig. 1). The rhynchonellides were collectedby F. Z. Malti in 2006–2008 during fieldwork for herPhD thesis on the Marhouma Formation, a unit first

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1010 B. M OT T E Q U I N A N D OT H E R S

Figure 1. (a, b) Geographic locations of the Saoura Valley in the Ougarta Basin in Algeria. Abbreviations: b. – Béchar; m. – Morocco.(c) Location of studied sections (CA – Cheffar El Ahmar; St – Gara Diba; Ou – Ouarourout; Ze – Tamtert–Zereg).

described by Poueyto (unpub. internal report, Sociéténationale de Recherche et d’Exploitation de Pétroleen Algérie (SN-REPAL), 1965) and Bastien et al. (un-pub. internal report, SN-REPAL, 1965) (e.g. Ouali Me-hadji et al. 2012 for more details), following a NNW–SSE transect along the Saoura Valley and more par-ticularly in Ouarourout (30° 10′ 30′′ N; 2° 14′ 30′′ W),Gara Diba (30° 7′ 38′′ N; 2° 12′ 30′′ W), Cheffar El Ah-mar (29° 57′ 25′′ N; 2° 7′ 15′′ W) and Tamtert-Zereg(29° 54′ 30′′ N; 1° 49′ 30′′ W) (Figs 1–3). She gatheredabundant and diverse marine macro- and microfaunas,including brachiopods, ammonoids (goniatites and cly-meniids identified by D. Korn), trilobites, conodonts,and spores.

Subdivisions of the Famennian Stage are those usedby Becker, El Hassani & Tahiri (2013, fig. 2) in align-ments between the conodont and ammonoid zonations(see also Clausen, Weddige & Ziegler, 1993; Korn,2002; Crônier et al. 2013). The acronyms do I to doVI refer to successive Frasnian–Famennian cephalo-pod zones (Wedekind, 1908) with ‘do’ meaning UpperDevonian.

F.Z. Malti (unpub. PhD thesis, Oran University andAbou Bekr Belkaid University, 2012) discriminatedfour successive members in the Marhouma Formation(Figs 2–3), the latter totalling about 260 m according to

Ouali Mehadji et al. (2012); it is overlain by sandstonesof the uppermost Famennian Ourarourout Formation(Fabre, Kazi-Tani & Moussine-Pouchkine, 2005).

Member 1 consists of calcareous shales devoid ofrhynchonellides; it is Late Frasnian in age (do Iβ) onthe basis of goniatites (Manticoceras sp.) and entomo-zoaceans (Casier, 1983).

Member 2 includes silty shales with ‘griotte’ nod-ules; it is divided into two submembers (a and b). Theterm ‘griotte’ is applied to nodular to pseudonodularlimestones rich in ammonoid cephalopods (Benhamouet al. 2004). Submember a comprises silty shales withnumerous specimens of Evanidisinurostrum saour-aense sp. nov. in the Gara Diba section and a fewSphaeridiorhynchus sp. in the Cheffar El Ahmar sec-tion in thin limestone lenses rich in horizontal burrows.The age of this submember cannot be precisely de-termined due to the lack of conodonts and goniatites;the rhynchonellides suggest a possible early Famen-nian age. Submember b consists of alternating shalesand ‘griotte’ limestone levels, sometimes in a succes-sion of centimetre-thick beds. A limestone bed of thissubmember in the Gara Diba section has produced theconodont Palmatolepis minuta subtilis in the intervalspanning the P. triangularis to P. trachytera zones (doII). An interval of ‘griotte’ limestone in the Cheffar El



Famennian rhynchonellides from the Saoura Valley 1011

Figure 2. Distribution of rhynchonellides, ammonoids, conodonts and palynomorphs within the Marhouma Formation in the SaouraValley (Ougarta Basin, Algeria). Abbreviations: G. – Grandispora; P. – Palmatolepis; U. – Umbellasphaericum devonicum.

Ahmar section produced Leptoterorhynchus sp. asso-ciated with Maenoceras; this rhynchonellide was pre-viously known from the middle Famennian.

Member 3 includes the ‘griotte’ limestone whichis divided into three submembers (a, b, c); one‘griotte’ level produced several conodonts indicat-

ive of the middle–upper Palmatolepis expansa Zone(upper–uppermost Famennian). This submember in theTamtert–Zereg section is late Famennian on the basisof Platyclymenia Zone ammonoids. It has a diverserhynchonellide fauna: Centrorhynchus sp., Leptoter-orhynchus sp., Phacoiderhynchus aff. antiatlasicus




Figure 3. Distribution of rhynchonellides within the Marhouma Formation in the Cheffar El Ahmar, Gara Diba, Ouarourout andTamtert–Zereg sections.

Sartenaer, 2000, Hadyrhyncha cf. hadyensis Havlícek,1979 and Novaplatirostrum sp.

Member 4 consists of shales with some ‘griotte’levels, but also sandy and silty deposits of turbiditicorigin; these are particularly thick at Ouarourout andTamtert–Zereg. It has three submembers (a, b, c), butonly the oldest submember produced rhynchonellides(Pugnaria sp.).

3. Systematic palaeontology

The material described and figured here is housed at theMuseum of the Central University of Algiers (MUA)and was investigated by Denise Brice. The supraspe-cific classification follows Savage et al. (2002) andSavage (2007) for the Order Rhynchonellida (exceptif stated otherwise); where used, open nomenclaturefollows Feldman’s (1994) rules. Measurements of spe-cimens assigned to Evanidisinurostrum saouraense sp.nov. are presented in Appendix 1 (see online supple-mentary data at http://journals.cambridge.org/geo).

Abbreviations. L, shell length; lc, lateral costae; mc,median costae; T, shell thickness; W, shell width; Ws,sulcus width.

Order RHYNCHONELLIDA Kuhn, 1949Superfamily RHYNCHOTREMATOIDEA Schuchert, 1913

Family TRIGONIRHYNCHIIDAE Schmidt, 1965Subfamily TRIGONIRHYNCHIINAE Schmidt, 1965

Genus Centrorhynchus Sartenaer, 1970

Type species. Camarotoechia baitalensis Reed, 1922,Famennian, Pamir.

Centrorhynchus sp.Figure 4a–b

Material. One incomplete articulated specimen (Ze2/8) from the Tamtert–Zereg section, Marhouma Form-ation, Member 3, Submember c.

Description. Shell small (16 mm in width), probablywidest at mid-length, subpentagonal in outline (aa =106°). Valves covered by angular, simple costae arisingat beaks; four median and eight lateral costae in ventralvalve (not known in dorsal valve).Ventral valve almostflat with obsolete, poorly defined sulcus; beak suberect.Dorsal valve (preserved only posteriorly) convex.

Remarks. This specimen is assigned to Centrorhynchuson the basis of its external morphology but additionalmaterial is required for a species-level identification.Centrorhynchus is a cosmopolitan genus assigned to

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the Trigonirhynchiinae by Nicollin & Brice (2004); itoccurs first in the lower Famennian (Balinski, 1995). InNorth Africa, the genus has been recognized in the east-ern part of the southern margin of the Tindouf Basin byBrice (in Gourvennec, Bitam & Robardet, 1997) andon its northern flank (Brice in Brice, Legrand-Blain& Nicollin, 2005) and doubtfully in the uppermost Fa-mennian of Tafilalt by Brice (in Brice, Legrand-Blain &Nicollin, 2005 and in Becker et al. 2013). According toammonoids from the Tamtert–Zereg section, our spe-cimen is of late Famennian age (do IV or IV/V, upperPlatyclymenia Zone).

Subfamily HEMITOECHIINAE Savage, 1996Genus Paurogastroderhynchus Sartenaer, 1970

Type species. Camarotoechia (?) nalivkini Abramian,1957, Famennian, Armenia.

Paurogastroderhynchus presaharensis Brice in Brice,Legrand-Blain & Nicollin, 2005

Figure 4c–d

2005 Paurogastroderhynchus presaharensis: Brice inBrice et al., p. 15–16, pl. 2, figs 1–7, 25a–b.

Material. One incomplete specimen from Tamtert–Zereg section (Ze 2/8), Marhouma Formation, Member3, Submember c.

Description. Shell large (33.6 mm in width), dors-ibiconvex, wider than long, rounded subpentagonal inoutline. Valves covered by simple, narrow and angularcostae near beaks, becoming wider and subangularanteriorly and flattened on flanks; five median andeight ventral lateral costae (not observed on dorsalvalve due to poor preservation); intercostal groovesnarrower than costae. Ventral valve slightly convex inumbonal area; apical angle 112°; tongue not preserved;traces of short, divergent dental plates. Traces of dorsalmedian septum.

Remarks. This single specimen is assigned to Pauro-gastroderhynchus presaharensis on the basis of itsexternal features (shape and ornamentation). Thisspecies was reported previously from the upperFamennian deposits of Algeria (Timimoun region;Sartenaer, 1975), Ahnet area (material from theKhenig Sandstones collected by M. Legrand-Blain;D. Brice, unpub. data) and of the Western Anti-Atlasin Morocco (Kheneg Lakahal section; Brice in Brice,Legrand-Blain & Nicollin, 2005) where it is abundant.It is also represented by some remains from the upperFamennian Hassi Rharouar section, SE of Tinfouchy,and the Zemoul area (Morocco) (Brice in Brice,Legrand-Blain & Nicollin, 2005). The specimenfrom Ze 2/8 of the Tamtert–Zereg section is of lateFamennian age (upper Platyclymenia Zone).

Superfamily CAMAROTOECHIOIDEA Schuchert, 1929Family LEIORHYNCHIDAE Stainbrook, 1945

Subfamily LEIORHYNCHINAE Stainbrook, 1945Genus Tenuisinurostrum Sartenaer, 1967

Figure 4. Rhynchonellide brachiopods from the MarhoumaFormation, Upper Devonian (Famennian), Saoura Valley, Al-geria (see Fig. 1). (a, b) Centrorhynchus sp., MUA/1094/020:incomplete specimen in ventral and posterior views. (c, d)Paurogastroderhynchus presaharensis Brice in Brice, Legrand-Blain & Nicollin, 2005, MUA/1094/021: incomplete speci-men in ventral and lateral views. (e–g) Tenuisinurostrum? sp.,MUA/1094/022: crushed specimen in ventral, dorsal and lateralviews. (h–l) Sphaeridiorhynchus sp., MUA/1094/023: completespecimen in ventral, dorsal, lateral, posterior and anterior views.Scale bars: 5 mm.

Type species. Camarophoria crenulata Gosselet, 1877,lower Famennian, southern Belgium.

Tenuisinurostrum? sp.Figure 4e–g

Material. One incomplete, partly deformed specimenfrom Gara Diba, Marhouma Formation, Member 2,Submember b.




Description. Shell of medium size (L ?, W = 19 mm,T = 8.2 mm), suboval in outline, weakly dorsibicon-vex; lateral commissures acute; anterior commissureuniplicate; anterior margin flat. Shell with rounded, ir-regular ribs present in anterior region, devoid of lateralribs, with a weak ventral median groove originating inthe umbonal region. Ventral valve with suberect beak;sulcus arising at about mid-length, poorly defined, flat-bottomed at front, shallow (?); tongue very low; apicalangle 125°; possible traces of a dental plate left ofthe beak. Dorsal valve with greatest convexity slightlyposterior to mid-length; fold weakly developed, low,flat-topped at front, originating at about mid-length;trace of a short median septum.

Remarks. Although poorly preserved, this single speci-men is tentatively assigned to Tenuisinurostrum mainlyon the basis of its suboval outline, its irregular ribsand its low tongue. It displays a trace of a rudiment-ary dental plate, though such structures are not repor-ted in this genus (see Sartenaer, 1967). Externally,it is closer to T. crenulatum rather than to T. sub-crenulatum Biernat, 1970, which is more dorsibiconvexand wider than long. Paromoeopygma Sartenaer, 1968(Pugnacoidea) possesses dental plates and displays asimilar outline in ventral view but differs by its moreinflated dorsal valve, its higher tongue and absence ofa dorsal median septum. Further material is thereforerequired for confident identification.

The type species of Tenuisinurostrum was describedfrom the lower Palmatolepis crepida Zone (Sartenaer,1984) in southern Belgium and northern France and T.subcrenulatum Biernat, 1970 is known from the P. tri-angularis to P. marginifera zones in the Holy CrossMountains of Poland (Biernat, 1970, 1983; Sarten-aer, 1985), though the latter needs to be revaluatedfrom the taxonomic viewpoint (Sartenaer, 1984, 1985,1987). Our badly preserved specimen of Tenuisinur-ostrum? sp. is of early Famennian age (do II) on thebasis of Palmatolepis minuta subtilis (P. triangularisto P. trachytera zones). Tenuisinurostrum is not knownwith certainty from North Africa (cf. Sartenaer, 1984,1987).

Genus Sphaeridiorhynchus Sartenaer, Pushkin &Kotlyar, 1997

Type species. Sphaeridiorhynchus kuzmichiensisSartenaer, Pushkin & Kotlyar, 1997, lower Famennian,Belarus.

Sphaeridiorhynchus sp.Figure 4h–l

Material. Eight specimens including one juvenile fromthe Cheffar El Hamar (CA 1a) section, MarhoumaFormation, Member 2, Submember a.

Description. Shell of medium size (up to 20.7 mmin width), wider than long, widest near mid-length,biconvex or weakly dorsibiconvex, suborbicular to sub-pentagonal in outline; anterior commissure uniplicate,anterior margin straight. Shell smooth, but two speci-

mens display median ribs arising in the anterior regionof the shell (mc = 0/3 and 5/?). Ventral valve inflated;beak erect; sulcus weak, arising in umbonal region or atabout mid-length, flat-bottomed; tongue nearly perpen-dicular to commissure plane, low, trapezoidal. Dorsalvalve strongly inflated with greatest convexity slightlyposterior to mid-length; fold weak, arising in umbonalarea, flat-topped at front; septum lacking.

Remarks. These specimens are assigned to Sphaeri-diorhynchus on the basis of their markedly biconvex,smooth and medium-sized shell with suborbicular tosubpentagonal outline, and by the weak fold and sul-cus. They are clearly less globular than specimens ofits type species, but their specific assignment requiresadditional material.

Until now, Sphaeridiorhynchus was known onlyfrom the lower Famennian succession of Belarus(Pripyat’ Depression) and Ukraine (Dnepr–Donets De-pression) (Sartenaer, Pushkin & Kotlyar, 1997), whereit is represented by S. kuzmichiensis. Discovery ofSphaeridiorhynchus sp. in the lower Famennian suc-cession of the Saoura Valley thus represents the firstoccurrence of this genus outside the Eastern EuropeanPlatform.

Superfamily PUGNACOIDEA Rzhonsnitskaia, 1956Family PUGNACIDAE Rzhonsnitskaia, 1956Genus Evanidisinurostrum Sartenaer, 1987

Type species. Pseudoleiorhynchus? zemoulensis Drot,1964, lower Famennian, plains of Drah-el-Kelba, pre-Saharan Morocco.

Evanidisinurostrum saouraense sp. nov.Figure 5a–t

Derivatio nominis. In reference to the Saoura Valley(Ougarta Basin, Algeria).

Holotype. A complete steinkern, MUA/1094/016.

Locus typicus. Gara Diba section (30° 7′ 38′′ N;2° 12′ 30′′ W), Saoura Valley, eastern part of the Ou-garta Basin, Algerian Sahara.

Stratum typicum. Marhouma Formation (lower Famen-nian), Member 2, Submember a.

Studied material. About 400 specimens (steinkerns),125 well preserved (of which 42 were measured), 27decorticated, 21 incomplete, 227 juveniles and numer-ous isolated valves and fragments from the Gara Dibasection.

Diagnosis. A small species of Evanidisinurostrum (upto c. 17 mm in width) with shell subpentagonal inoutline, wider than long, dorsibiconvex, widest nearmid-length or more anteriorly. Fold and sulcus start-ing posterior to mid-length. Tongue moderately high,subtrapezoidal in outline, perpendicular or almost per-pendicular to the commissural plane. Most of shellssmooth with rounded median costae arising at or pos-terior to mid-length (mc = 2–3/1–2, usually 2/1); lateralcostae usually absent or rare (0–1/0–2).




Figure 5. Rhynchonellide brachiopods from the Marhouma Formation, Upper Devonian (Famennian), Saoura Valley, Algeria (seeFig. 1). (a–t) Evanidisinurostrum saouraense sp. nov. from the Marhouma Formation, Upper Devonian (Famennian), Saoura Valley,Algeria (see Fig. 1). (a–e) holotype, MUA/1094/016: complete specimen in ventral, dorsal, lateral, posterior and anterior views. (f–j)MUA/1094/018: complete specimen in ventral, dorsal, lateral, posterior and anterior views. (k–o) MUA/1094/017: complete specimenin ventral, dorsal, lateral, posterior and anterior views. (p–t) MUA/1094/19: complete juvenile specimen in ventral, dorsal, lateral,posterior and anterior views. (u–y) Hadyrhyncha cf. hadyensis Havlícek, 1979, MUA/1094/024: complete specimen in ventral, dorsal,lateral, posterior and anterior views. (z–ii) Leptoterorhynchus sp.: (z–dd) MUA/1094/025: complete specimen in ventral, dorsal, lateral,posterior and anterior views; (ee–ii) MUA/1094/026: complete specimen in ventral, dorsal, lateral, posterior and anterior views. Scalebar: 5 mm.

Description. Shell small (up to 17.2 mm in width),dorsibiconvex, wider than long, widest near mid-lengthor more anteriorly, subpentagonal in outline; anteriorcommissure uniplicate, anterior margin, in ventralview, indented by a strong median costa. Most shellsare smooth on the flanks with rounded median costaearising at or posterior to mid-length (mc = 2–3/1–

2, usually 2/1); lateral costae usually absent or rare(0–1/0–2). Ventral valve gently inflated in both pos-terior and lateral profiles with greatest convexity pos-terior to mid-length; beak small; apical angle between110° and 120°; sulcus wide, deep, originating at aboutmid-valve, with prominent median costa; tongue widerthan long, almost perpendicular to commissural plane,




trapezoidal; dental plates absent; muscle field im-pressed, triangular in outline. Dorsal valve evenly con-vex in both lateral and posterior profiles, highest atfront, impressed; fold originating at mid-length, relat-ively flat-topped at front (apart from costae); no medianseptum; myophragm short.

Dimensions in mm (n = 42 for all; see Appen-dix 1 in online supplementary data, available athttp://journals.cambridge.org/geo). Width range 9.75–17.2, average 12.5; length range 8.75–13.55, average10.6; thickness range 4.2–7.1, average 5.5; width ofsulcus range 4–13.1, average 8.18; width/length ratiorange 1.01–1.37, average 1.18; width/thickness ratiorange 1.78–2.71, average 2.27; sulcus width/width ra-tio range 0.41–0.82, average 0.64.

Remarks. As all the specimens are steinkerns, it was notpossible to investigate the interiors by serial sections.These specimens are assigned to Evanidisinurostrumrather than Perrarisinurostrum Sartenaer, 1984 andTenuisinurostrum Sartenaer, 1967 (see Sartenaer, 1987for discussion of the differences between these threegenera) as they have similarities with E. zemoulense(Drot, 1964) such as the outline, the similar develop-ment of fold and sulcus and the absence of dental platesand median septum. Nevertheless, the specimens fromthe Saoura Valley can be discriminated from Drot’s(1964) species by their smaller size, their maximumwidth near the mid-length or anteriorly, their differentW/L ratio, their larger apical angle and their differentmedian ornament.

Evanidisinurostrum was previously known with cer-tainty from the lower Famennian IIβ by its type speciesin the Zemoul and south of Akka in the Dra Plainsof pre-Saharan Morocco (Drot, 1964; Sartenaer, 1987)and on the southern margin of the Tindouf Basin (Al-geria; Drot, 1964). Furthermore, Sartenaer (1987, p.135) suggested that Tenuisinurostrum subcrenulatumBiernat, 1970 from the Holy Cross Mountains in Po-land (middle Palmatolepis triangularis to lower P. mar-ginifera zones) may belong to Evanidisinurostrum, butthis needs confirmation.

Family ROZMANARIIDAE Havlícek, 1982Genus Hadyrhyncha Havlícek, 1979

Type species. Hadyrhyncha hadyensis Havlícek, 1979,upper Famennian, Moravia, Czech Republic.

Hadyrhyncha cf. hadyensis Havlícek, 1979Figure 5u–y

cf. 1979 Hadyrhyncha hadyensis: Havlícek, p. 99, pl.2, figs 6–9, text-fig. 8.

Material. Two specimens, of which one is incomplete,from the Marhouma Formation, Member 3, Submem-ber c, of the Tamtert–Zereg section (Ze 2/8, Ze 2/8b).

Description. Shell small (up to 15.7 mm in width),wider than long, widest near mid-length, transverselyelliptic in outline, slightly ventribiconvex; anteriorcommissure unisulcate; anterior margin straight. Low,

rounded median costae originating near umbones(mc = 3/4), the central one bifurcating both in the sul-cus and on the fold; lateral costae badly preserved (lc= 6/5–8), corresponding to the division of 2 or 3 costaepresent in the posterior region. Ventral valve withgreatest convexity near mid-length; beak small, erect;fold low, flat-topped at front, arising in posterior part ofthe valve; tongue low, subtrapezoidal, perpendicular tocommissural plane. Dorsal valve with greatest convex-ity posterior to mid-length; sulcus arising near umbo,well-defined, flat-bottomed at front.

Remarks. These specimens share numerous externalsimilarities with Hadyrhyncha hadyensis, such as theirtransversally elliptic outline, the similar developmentof their dorsal sulcus and ventral fold, their sharpcommissures and their ornament. However, as the in-ternal morphology of the Algerian material remainsunknown, they are tentatively compared to Havlícek’s(1979) species. They differ from Hadyrhyncha meridi-onalis Sartenaer, 1998a from Morocco by their smallersize, their narrower apical angle, their ornament andtheir different L/W, T/W and T/L ratios. They are sep-arated from Hadyrhyncha sp. from Poland (Halamski &Balinski, 2009) by their smaller size and less transverseoutline.

Our specimens are associated with ammonoids ofthe upper Famennian Platyclymenia Zone (do IV orIV/V). Hadyrhyncha hadyensis is known from the up-per or uppermost Famennian (do V or do VI accordingto Havlícek, 1979) of Moravia (Czech Republic) andfrom the middle part of do V in south-easternThuringia(Germany) (Bartzsch & Weyer, 1986). H. meridionalisSartenaer, 1998a occurs in the upper Famennian (doV) of southern Morocco (Dra Valley, Maider, Ta-filalt, Zemoul); the genus is also reported from Po-land (Holy Cross Mountains) in the upper Palmatolepismarginifera to upper Siphonodella praesulcata zones(Halamski & Balinski, 2009).

Genus Leptoterorhynchus Sartenaer, 1998b

Type species. Rozmanaria magna Biernat & Racki,1986, middle Famennian, Holy Cross Mountains, Po-land.

Leptoterorhynchus sp.Figure 5z–ii

Material. Three specimens from the Marhouma Form-ation, Member 3, Submember c of the Tamtert–Zereg(Ze 2/8) section and two from the Marhouma Forma-tion, Member 2, Submember b of the Cheffar El Ahmar(CA 1c, CA2) section.

Description. Shell small (up to 13.1 mm in width),smooth, ventribiconvex, wider than long, widest aboutmid-length and transversally elliptical in outline; an-terior commissure unisulcate; anterior margin straight.Ventral valve with greatest convexity near mid-length;beak small, erect; fold poorly developed, low, round-topped at front; tongue high, bent ventrally. Dorsalvalve with greatest convexity near valve mid-length;

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sulcus wide, originating in posterior part of the valve,shallow, flat-bottomed at front; tongue low, rounded,perpendicular to commissural plane or bent dorsally;internal features not observed; fold poorly developed,low, round-topped at front; no median septum.Remarks. These specimens are assigned to Leptotero-rhynchus as they share many external similarities withthe type species L. magnus (Biernat & Racki, 1986)such as the outline, the slightly developed and smoothsulcus and fold, and the sharp commissure. They prob-ably belong to a new species which is not named be-cause of insufficient material. However, they can be dis-criminated from the type species by their smaller size,their narrower apical angle, their different W/L, W/Tand T/L ratios, their not-so-well-defined sulcus and foldand the absence of a median depression on the fold.

Leptoterorhynchus magnus (Biernat & Racki, 1986)is known only from the middle Famennian deposits ofthe southern Holy Cross Mountains, Poland accordingto Halamski & Balinski (2009). An unidentified specieswas reported in the Famennian (UD IIIC–IVB interval)of north-western Sauerland in Germany by Sartenaer(1998b). In the Saoura Valley, Leptoterorhynchus sp. isknown from the upper Famennian Tamtert–Zereg sec-tion on the basis of Platyclymenia annulata and fromthe middle Famennian (on the basis of Maenoceras)Cheffar El Ahmar section.

Genus Novaplatirostrum Sartenaer, 1997

Type species. Novaplatirostrum sauerlandense Sarten-aer, 1997, upper Famennian, north-western Sauerland,Germany.

Novaplatirostrum sp.Figure 6a–j

Material. Five specimens from the Tamtert–Zereg sec-tion (Ze 2/8, 2/8b, 2/9) and one from “Plat” betweenOuarourout and Gara Diba; Marhouma Formation,Member 3, Submember c.

Description. Shell small (up to 16.2 mm in width),flatly biconvex, suborbicular in outline; anterior com-missure uniplicate; anterior margin slightly rounded,with sharp commissures (juveniles uniplicate, flattlybiconvex, transversely subelliptical in outline). Lowrounded median costae (3/2 or 4/3) near anterior com-missure; no lateral costae. Ventral valve with greatestconvexity in umbonal area; sulcus absent or wide(Ws/W: 0.68; n = 1), very shallow, starting from mid-length, flat-bottomed; tongue low, not perpendicularto commissural plane; maximal apical angle of 127°.Dorsal valve with greatest convexity posterior to mid-valve; fold obsolete, flat-bottomed.

Remarks. The Algerian specimens are externally closeto Novaplatirostrum sauerlandense Sartenaer, 1997,especially paratypes F and G illustrated by Sartenaer(1997, pl. 1, figs 31–40) but they are smaller and flatterthan the latter and devoid of lateral costae. Our speci-mens display less median costae, have an obsolete foldand a smaller apical angle.

In the Saoura Valley, Novaplatirostrum sp. is knownfrom the Tamtert–Zereg section where it is associatedwith goniatites and clymeniids of late Famennian age inlevels Ze 2/8 and Ze 2/8b (upper Platyclymenia Zone)and in level Ze 2/9 (Gonioclymenia Zone). The age ofthe latter could be younger on the basis of a conodontfauna characteristic of the early upper Palmatolepisexpansa Zone. N. sauerlandense occurs in deep-waterenvironments in the German upper Famennian depos-its (southern Sauerland and south-eastern Thuringia;Sartenaer, 1997) where it is associated with goniatites,but also in the Holy Cross Mountains in Poland in levelsof probable do V–VI age (Halamski & Balinski, 2009;A. Halamski, pers. comm., 2014).

Genus Pugnaria Biernat & Racki, 1986

Type species. Pugnaria plana Biernat & Racki, 1986,middle Famennian, Holy Cross Mountains, Poland.

Pugnaria sp.Figure 6h–o

Material. Three specimens (one incomplete adult andtwo juveniles) from the Tamtert–Zereg (Ze 2/8) sec-tion (Marhouma Formation, Member 3, Submember c)and one adult from the Ouarourout (Ou 1/15) section(Marhouma Formation, Member 4, Submember a).

Description. Shell small (19.2 mm in width), widerthan long, flatly dorsibiconvex, transversely elliptical inoutline; anterior commissure rectimarginate; anteriormargin slightly rounded. Shell smooth, growth linesthin, irregularly distributed, observed on ventral valve.Ventral valve with greatest convexity posterior to mid-length; beak small, curved; sulcus lacking. Dorsal valvewith greatest convexity posterior to mid-length; foldlacking.

Remarks. Despite lack of knowledge about the internalfeatures, our specimens are externally very close toPugnaria plana Biernat & Racki, 1986 but are smal-ler, devoid of a fold and display a wider apical angle.Additional material is required for accurate specificidentification.

Pugnaria sp. occurs in do III–do IV in the Tamtert–Zereg section (Ze 2/8) and is also known from the Ou-arourout section (Ou1/15) where it is associated withammonoids of probable do V age. In the Kowala sec-tion (Holy Cross Mountains, Poland), the range of P.plana in terms of conodont zones spans the Palmato-lepis marginifera to Siphonodella praesulcata zones(Halamski & Balinski, 2009).

Genus Phacoiderhynchus Sartenaer, 2000

Type species. Phacoiderhynchus antiatlasicus Sarten-aer, 2000, middle Famennian, Maider, Morocco.

Phacoiderhynchus aff. antiatlasicus Sartenaer, 2000Figure 6p–x

aff. 2000 Phacoiderhynchus antiatlasicus n. gen., n.sp.: Sartenaer, p. 79–84, pl. 1, figs 1–30, pl. 2, figs31–65.




Figure 6. Rhynchonellide brachiopods from the Marhouma Formation, Upper Devonian (Famennian), Saoura Valley, Algeria (seeFig. 1). (a–j) Novaplatirostrum sp.: (a–e) MUA/1024/027: complete specimen in ventral, dorsal, lateral, posterior and anterior views;(f–j) MUA/1094/028: complete specimen in ventral, dorsal, lateral, posterior and anterior views. (k–o) Pugnaria sp., MUA/1094/029:complete specimen in ventral, dorsal, lateral, posterior and anterior views. (p–x) Phacoiderhynchus aff. antiatlasicus Sartenaer, 2000:(p–s) MUA/1094/030: almost complete specimen in ventral, dorsal, lateral and posterior views; (t–x) MUA/1094/031: incompletespecimen in ventral, dorsal, lateral, posterior and anterior views. Scale bars: 5 mm.

Material. Eleven incomplete specimens from theTamtert–Zereg section (Ze 2/7: one adult and onejuvenile; Ze 2/8: six adults, one juvenile; Ze2/8b: two adults), Marhouma Formation, Member 3,Submember c.

Description. Shell of medium-size (up to 29 mm inwidth), wider than long, flatly biconvex, transversallyelliptical in outline, widest at mid-length; commissuresharp; anterior commissure uniplicate; anterior marginflattened. Shells mostly smooth with rounded and wide

median costae sometimes divided near the frontal mar-gin (mc: 3–4/2–4); lateral costae not preserved. Vent-ral valve with greatest convexity close to mid-length;beak very small; sulcus shallow, flat-bottomed at front;tongue very low, not perpendicular to commissuralplane. Dorsal valve with maximum convexity near mid-valve; fold scarcely developed, originating anterior tomid-length.

Remarks. These incomplete specimens are tentativelyassigned to Phacoiderhynchus antiatlasicus Sartenaer,




2000 on the basis of their external morphology, al-though they can be distinguished by their smaller size,their sulcus and fold starting a little more anteriorlyand their less numerous median costae (the lateral onesare absent or not preserved). The specimens from theSaoura Valley have a narrower apical angle (134–145°versus 147–157°) and may represent a new species, butspecific assignment requires additional material.

Phacoiderhynchus aff. antiatlasicus is known fromthe upper Famennian Tamtert–Zereg section on thebasis of ammonoids (upper Platyclymenia Zone). P.antiatlasicus occurs with certainty in middle–upper Fa-mennian IIIB to IVB deposits from southern Morocco(Maïder and eastern Draa Plains) according to Sarten-aer (2000), and probably in the Immouzer du Kandararea south of Fès (central Morocco; see Brice et al.1984).

4. Discussion

4.a. Stratigraphic summary and general comments onbrachiopod diversity during Famennian time

The stratigraphic range of the rhynchonellides withinthe Famennian part of the Marhouma Formation acrossthe Saoura Valley (Ouarourout, Gara Diba, CheffarAmar and Tamtert–Zereg sections) is presented inFigures 2 and 3.

As the Frasnian–Famennian (F–F) boundary, andthus the position of the Upper Kellwasser Event (Pal-matolepis linguiformis Zone), is not precisely estab-lished within the Marhouma Formation, it is difficultto discuss in detail the post-Kellwasser recovery of thebrachiopods in this part of the Ougarta Basin. Note thatin regions where environmental conditions were morefavourable to brachiopods, such as the Namur–DinantBasin in southern Belgium and the Polish epicontin-ental basin, their recovery started in the Palmatolepistriangularis Zone very soon after the F–F boundary,in which athyridides, rhynchonellides and spiriferidesplayed a significant role (Balinski 1996, 2002; Mot-tequin 2008a, b). On a worldwide scale, brachiopoddiversity declined significantly during Frasnian timewith a significant reduction in generic diversity; thiswas accentuated during Famennian time, possibly dueto increasing faunal cosmopolitanism related to majorsea-level changes and other parameters that suppressedfaunal barriers and allowed interchange between re-mote areas (e.g. Copper, 1998; Brice et al. 2000). Fromthe rhynchonellide viewpoint the Famennian Stage ismarked by great generic diversity, contrasting withother brachiopod orders (e.g. orthides); this is similarto the situation reported during Emsian time (Curry &Brunton, 2007) and the occurrence of numerous cos-mopolitan genera, including some of those identifiedin the Marhouma Formation (see discussion in Section4.c). This Devonian stage is also characterized by thepredominance of the rhynchonellides within brachio-pod faunas (Curry & Brunton, 2007).

4.b. Palaeoecology and taphonomy

Brachiopods obtained from several horizons of the Fa-mennian part of the Marhouma Formation include,in order of abundance, rhynchonellides (439 spe-cimens), orthides (small Aulacella; 45 specimens),athyridides (medium-size Composita; 4 specimens)and spiriferides (only one small specimen; affinitiesnot apparent due to poor preservation). Rhynchonel-lides therefore represent 90 % of the whole brachiopodassemblage, with 10 species. For comparison, 197 am-monoids (goniatites and clymeniids; D. Korn and F.Z.Malti, unpub. data) and 19 trilobites (Crônier et al.2013), including the material from two other sectionsof the Saoura Valley (Béchir and Idhir; see locationin Crônier et al. 2013), have been collected in thesame levels of the Marhouma Formation. Within theselithostratigraphic units, the greatest rhynchonellide di-versity is at the top of Submember c of Member 3,which corresponds to the top of the early upper P. ex-pansa Zone, i.e. at the base of the upper Famenniandeposits (Fig. 2). Development of the ‘griotte’ lime-stones took place during a sea-level rise (F.Z. Malti,unpub. data) that probably facilitated faunal exchangesbetween distant basins.

The low-diversity brachiopod fauna is clearly an insitu rhynchonellide-dominated brachiopod communitywith much lower diversity of other orders; the spe-cimens are not disarticulated, sorted or broken. Nostrophomenides or productides occurred in associationwith the rhynchonellides. Deep-water conditions areindicated by cephalopods (goniatites and clymeniids)and trilobites; the latter comprise blind to reduced-eyed phacopides (Crônier et al. 2013). One of themost striking features of this rhynchonellide fauna isthe predominance of smooth, thin-shelled and smallforms (except some scarce large Trigonirhynchiidae),also characterized by flattened shells. Moreover, amongthese rhynchonellides, leiorhynchids (Camarotoech-ioidea) and rozmanariids (Pugnacoidea) are by far themost abundant elements as is generally the case in latePalaeozoic dysaerobic communities, though the Pug-nacoidea are represented by other families (cf. Bowen,Rhoads & McAlester, 1974; Biernat & Racki, 1986;Racki, 1989; Alexander, 1994; Mottequin & Legrand-Blain, 2010). We also draw attention to the occurrenceof a monospecific assemblage with Evanidisinurostrumsaouraense in which immature specimens are particu-larly abundant. Almost monospecific rhynchonellideassemblages of leiorhynchids have been reported else-where, notably in the Frasnian deposits of southernBelgium (e.g. Sartenaer, 1974) and Poland (Racki,1993; Racki & Szulczewski in Sartenaer, Racki &Szulczewski, 1998). Overrepresentation of juvenilesof E. saouraense is probably related to the combina-tion of several biological (e.g. nutrient availability) andphysicochemical controlling factors (including wateroxygenation and depth) (see Pérez-Huerta & Sheldon,2006) that induced significant losses among immaturespecimens.




The state of preservation of our material does notpermit precision as regards the size of the foramenfor each species, but adults of the identified generaare characterized by a reduced foramen (cf. data inSavage et al. 2002). We are therefore inclined to re-gard the rhynchonellides as lying unattached on the seafloor. Another significant feature is the aequibiconvex-ity of the shells, notably for Pugnaria, Novaplatisin-urostrum and Phacoiderhynchus. According to Walker(1974) this is indicative of soft substrates; it was in-terpreted in this way by Biernat & Racki (1986) forFamennian rhynchonellides. Relatively more biconvexshells such as those of Sphaeridiorhynchus may accordwith a firmer substrate (see Racki & Szulczewski inSartenaer, Racki & Szulczewski, 1998). According toRacki (1998), the deep-water rhynchonellide biofaciesdisplays apparent continuity in spite of generic lossesacross the Frasnian–Famennian boundary. One of themajor changes in this biofacies is the huge developmentof the rozmanariids during Famennian time. They madetheir appearance during Pragian time (cf. Savage et al.2002; Savage, 2007), before disappearing at the endof the Devonian period (Mottequin, Brice & Legrand-Blain, 2014).

4.c. Comparison with other rhynchonellide faunas fromNorth Africa and biogeographic affinities

Despite the increased number of publications, Famen-nian brachiopod assemblages from North Africa, andtherefore from the northern margin of Gondwana,are still poorly documented. This is due in part tomost papers being generally focused on a particulargroup, as with this paper. Taxonomic data are availablefor most of the important brachiopod orders presentin the Famennian deposits of Algeria and Morocco(see below for Libyan data): productides (e.g. Brous-miche, 1975; Legrand-Blain, 1995a, b; Legrand-Blainin Brice, Legrand-Blain & Nicollin, 2005), spiriferides(e.g. Drot, 1964; Nicollin & Brice, 2000; Brice &Nicollin in Brice, Legrand-Blain & Nicollin, 2005),and rhynchonellides.

Five rhynchonellide genera, namely Centro-rhynchus, Evanidisinurostrum, Hadyrhyncha, Pauro-gastroderhynchus and Phacoiderhynchus, occurringwithin the Marhouma Formation were previously repor-ted from Famennian successions on the northern mar-gin of Gondwana (Morocco and Algeria). Attention isdrawn to two salient absences from the rhynchonellidesof the Marhuma Formation: (1) TetragonorhynchusSartenaer, 1999, reported from the upper Famennian(UD V) deep-water facies from southern Morocco(Maïder) (Sartenaer, 1999); and (2) DzieduszyckiaSiemiradzki, 1909 represented by two species in theFamennian deposits of the Middle Atlas of Morocco(Balinski & Biernat, 2003). Dzieduszyckia has beenconsidered to be typical of Upper Devonian vent-seepenvironments (Campbell & Bottjer, 1995), but Balinski& Biernat (2003) have challenged this judgement on thebasis of geochemical analyses of rocks associated with

these Moroccan occurrences, insisting that no particu-lar environmental conditions can be discerned.

Famennian brachiopods from western Libya weredescribed by Havlícek (1984), Havlícek & Röhlich(1987), and Mergl & Massa (1992). They came fromshallow-water environments characterized by silici-clastic deposits with benthic communities dominatedby lingulide and rhynchonellide brachiopods (Mergl& Massa, 2000). The Libyan rhynchonellides includespecies of Cupularostrum Sartenaer, 1961 and Liby-aerhynchus Mergl & Massa, 1992 that are coarselyribbed, and thus markedly different from those occur-ring in contemporaneous deep-water environments ofMorocco and Algeria. As rightly reported by Mergl &Massa (1992) and Mergl, Massa & Plauchut (2001), thestratigraphic range of Cupularostrum, originally sug-gested as Givetian, should be reconsidered.

Four genera known from deep-water environmentsare recognized for the first time in North Africa:Pugnaria, Leptoterorhynchus, Sphaeridiorhynchus andNovaplatirostrum. They were previously known onlyfrom the south-eastern margin of Laurussia and wereviewed as being restricted geographically: Leptoter-orhynchus (Germany and Poland; Sartenaer, 1998b),Novaplatirostrum (Germany and Poland; Sartenaer,1997; Halamski & Balinski, 2009), Pugnaria (Po-land; Biernat & Racki, 1986), and Sphaeridiorhynchus(Belarus and Ukraine; Sartenaer, Pushkin & Kotl-yar, 1997). Tenuisinurostrum, previously reported fromwestern and possibly eastern Europe (cf. Sartenaer,1987), is here reported from the Famennian MarhoumaFormation, but needs confirmation on the basis of well-preserved material.

Halamski & Balinski (2013) recently stressed thegreat similarities between Middle Devonian benthicfaunas from the northern and southern shores of theVariscan Sea, which accords with a narrow oceaniczone between Laurussia and Gondwana (McKerrowet al. 2000) rather than an extensive Rheic Ocean(e.g. Torsvik & Cocks, 2011, 2013; Stampfli et al.2013). Legrand-Blain (1995a) also noted clear af-finities between the productides, rhynchonellides andspiriferides of North Africa and Southern Europe withthose from Armenia and the Urals across the Devonian–Carboniferous boundary. Close affinities between theFamennian rhynchonellides from central Algeria andthose from western and eastern Europe is consistentwith the hypothesis of prolonged proximity betweenboth continental masses, but island arcs between Laur-ussia and Gondwana (Stampfli et al. 2013; Torsvik &Cocks, 2013) could have facilitated dissemination andinteraction of the brachiopod faunas on both sides ofthe Rheic Ocean.

5. Conclusions

In the Saoura Valley (Ougarta Basin, Algeria), theFamennian part of the Marhouma Formation (mem-bers 2–4) contains rhynchonellide-dominated brachi-opod assemblages in which a generally low diversity




of small athyridides (Composita), orthides (Aula-cella) and spiriferides occur. The rhynchonellides, lei-orhynchids and rozmanariids are the most abundantand the most diverse elements, represented by small,thin-shelled species. These brachiopods are generallyfound in association with cephalopods and blind oreye-reduced phacopide trilobites (Crônier et al. 2013),interpreted to have lived in deep, oxygen-depletedenvironments. Such rhynchonellide assemblages areinterpreted as being characteristic of dysaerobic en-vironments found on epicontinental platforms duringLate Devonian time, especially in Laurussia and NorthGondwana. The leiorhynchid Sphaeridiorhynchus andthe rozmanariids Leptoterorhynchus, Pugnaria andNovaplatirostrum are reported for the first time out-side Europe; their occurrence on the margin of Gond-wana reinforces the biostratigraphic value of these fourgenera and underscores the global character of the Fa-mennian benthic faunas.

Acknowledgements. We are greatly indebted to DieterKorn for having provided identifications of the cephalopodscited in the text, to Thierry Hubin for technical assistanceand to Adam T. Halamski, Andrzej Balinski and John Talentfor reviewing the manuscript. This paper is a contributionto the International Geoscience Programme (IGCP) Project596, Climate change and biodiversity patterns in the MidPalaeozoic.

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