Geol. Mag. 152 (6), 2015, pp. 1009–1024. c Cambridge University Press 2015 1009 doi:10.1017/S0016756814000697 Famennian rhynchonellides (Brachiopoda) from deep-water facies of the Basin ( Valley, )

∗ BERNARD MOTTEQUIN† , FATIMA ZOHRA MALTI‡, MADANI BENYOUCEF§, ∗∗ CATHERINE CRÔNIER¶, LOUISA SAMAR║, CARINE RANDON & DENISE BRICE†† †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 of the essentially shally Formation is characterized by deep-water facies and includes horizons rich in ammonoids (goniatites and clymeniids) and blind to eye-reduced phacopide trilobites. They are also rich in small-sized and smooth rhynchonellide brachiopods, investigated here for the first time in order to detail their post-Kellwasser recovery. Rhynchonellides clearly predominate in the brachiopod assemblages (representing 90 % of the whole assemblage, with 10 species) composed otherwise of athyridides, orthides and spiriferides. Rhynchonellides are mostly represented by relatively flat leiorhynchids and rozmanariids consistent with poor oxygenation on the sea floor. One new species is described (Evanidisinurostrum saouraense sp. nov.); four genera, previously known only from the south-eastern margin of Laurussia, are reported for the first time from the northern margin of Gondwana: the leiorhynchid Sphaeridiorhynchus and the rozmanariids Leptoterorhynchus, Pugnaria and Novaplatirostrum. Keywords: Upper Devonian, Marhouma Formation, North Gondwana, brachiopods, Rhynchonellida, palaeoecology.

1. Introduction 2007). Brachiopods are of interest because they flour- ished along the northern margin of Gondwana in neritic Upper Devonian deposits were first recognized in facies sometimes unfavourable to ammonoids and con- the Saoura Valley (Algeria, Ougarta Basin) by Haug odonts, and are therefore valuable tools for biostrati- (1903) on the basis of cephalopods (goniatites and graphy in such palaeoenvironmental contexts, espe- clymeniids), which are particularly abundant in some cially regarding the Devonian–Carboniferous boundary levels (see also Menchikoff, 1930). The first compre- (Mottequin, Brice & Legrand-Blain, 2014). hensive study on the geology of this area was by Gau- This paper is the first comprehensive study of the tier (1906) and refined by Alimen et al. (1952), who diverse and small rhynchonellide brachiopods from described several important sections. The richness in the Famennian (Marhouma Formation) Saoura Valley ammonoids enabled Petter (1959) and Göddertz (1987) where they are the dominant element in the brachiopod to discriminate the zonation established by Wedekind assemblages, although they also occur in benthic com- (1908, 1926) in Germany and subsequently modified munities (cf. Crônier et al. 2013 for trilobites). Their by several authors (e.g. Korn, Klug & Reisdorf, 2000; diversity and palaeoecology are also discussed. Korn, 2004). Lower and Middle Devonian brachiopod faunas from the Ougarta were first described by Le Maître (1952) 2. Geological setting and material and revised in part by Boumendjel et al. (1997), Ouali Mehadji et al. (2004), and Brice et al. (2011) but, un- The material described in this paper comes from til recently, Upper Devonian brachiopods were poorly the Saoura Valley in the eastern part of the Ou- known (e.g. Brice, Legrand-Blain & Nicollin, 2005, garta Basin (Algerian Sahara), about 350 km SSW of Béchar (Fig. 1). The rhynchonellides were collected by F. Z. Malti in 2006–2008 during fieldwork for her ∗ Author for correspondence: [email protected] PhD thesis on the Marhouma Formation, a unit first

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

described by Poueyto (unpub. internal report, Société Ouali Mehadji et al.(2012); it is overlain by sandstones nationale de Recherche et d’Exploitation de Pétrole of the uppermost Famennian Ourarourout Formation en Algérie (SN-REPAL), 1965) and Bastien et al. (un- (Fabre, Kazi-Tani & Moussine-Pouchkine, 2005). pub. internal report, SN-REPAL, 1965) (e.g. Ouali Me- Member 1 consists of calcareous shales devoid of hadji et al. 2012 for more details), following a NNW– rhynchonellides; it is Late Frasnian in age (do Iβ)on SSE transect along the Saoura Valley and more par- the basis of goniatites (Manticoceras sp.) and entomo- ticularly in Ouarourout (30° 10 30 N; 2° 14 30 W), zoaceans (Casier, 1983). Gara Diba (30° 7 38 N; 2° 12 30 W), Cheffar El Ah- Member 2 includes silty shales with ‘griotte’ nod- mar (29° 57 25 N; 2° 7 15 W) and Tamtert-Zereg ules; it is divided into two submembers (a and b). The (29° 54 30 N; 1° 49 30 W) (Figs 1–3). She gathered term ‘griotte’ is applied to nodular to pseudonodular abundant and diverse marine macro- and microfaunas, limestones rich in ammonoid cephalopods (Benhamou including brachiopods, ammonoids (goniatites and cly- et al. 2004). Submember a comprises silty shales with meniids identified by D. Korn), trilobites, conodonts, numerous specimens of Evanidisinurostrum saour- and spores. aense sp. nov. in the Gara Diba section and a few Subdivisions of the Famennian Stage are those used Sphaeridiorhynchus sp. in the Cheffar El Ahmar sec- by Becker, El Hassani & Tahiri (2013, fig. 2) in align- tion in thin limestone lenses rich in horizontal burrows. ments between the conodont and ammonoid zonations The age of this submember cannot be precisely de- (see also Clausen, Weddige & Ziegler, 1993; Korn, termined due to the lack of conodonts and goniatites; 2002; Crônier et al. 2013). The acronyms do I to do the rhynchonellides suggest a possible early Famen- VI refer to successive Frasnian–Famennian cephalo- nian age. Submember b consists of alternating shales pod zones (Wedekind, 1908) with ‘do’ meaning Upper and ‘griotte’ limestone levels, sometimes in a succes- Devonian. sion of centimetre-thick beds. A limestone bed of this F.Z. Malti (unpub. PhD thesis, Oran University and submember in the Gara Diba section has produced the Abou Bekr Belkaid University, 2012) discriminated conodont Palmatolepis minuta subtilis in the interval four successive members in the Marhouma Formation spanning the P. triangularis to P. trachytera zones (do (Figs 2–3), the latter totalling about 260 m according to II). An interval of ‘griotte’ limestone in the Cheffar El

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

Ahmar section produced Leptoterorhynchus sp. asso- ive of the middle–upper Palmatolepis expansa Zone ciated with Maenoceras; this rhynchonellide was pre- (upper–uppermost Famennian). This submember in the viously known from the middle Famennian. Tamtert–Zereg section is late Famennian on the basis Member 3 includes the ‘griotte’ limestone which of Platyclymenia Zone ammonoids. It has a diverse is divided into three submembers (a, b, c); one rhynchonellide fauna: Centrorhynchus sp., Leptoter- ‘griotte’ level produced several conodonts indicat- orhynchus sp., Phacoiderhynchus aff. antiatlasicus

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

Sartenaer, 2000, Hadyrhyncha cf. hadyensis Havlícek,ˇ Order RHYNCHONELLIDA Kuhn, 1949 1979 and Novaplatirostrum sp. Superfamily RHYNCHOTREMATOIDEA Schuchert, 1913 Member 4 consists of shales with some ‘griotte’ Family TRIGONIRHYNCHIIDAE Schmidt, 1965 levels, but also sandy and silty deposits of turbiditic Subfamily TRIGONIRHYNCHIINAE Schmidt, 1965 origin; these are particularly thick at Ouarourout and Genus Centrorhynchus Sartenaer, 1970 Tamtert–Zereg. It has three submembers (a, b, c), but Type species. Camarotoechia baitalensis Reed, 1922, only the oldest submember produced rhynchonellides Famennian, Pamir. (Pugnaria sp.). Centrorhynchus sp. Figure 4a–b

3. Systematic palaeontology Material. One incomplete articulated specimen (Ze 2/8) from the Tamtert–Zereg section, Marhouma Form- The material described and figured here is housed at the ation, Member 3, Submember c. Museum of the Central University of Algiers (MUA) and was investigated by Denise Brice. The supraspe- Description. Shell small (16 mm in width), probably cific classification follows Savage et al.(2002) and widest at mid-length, subpentagonal in outline (aa = Savage (2007) for the Order Rhynchonellida (except 106°). Valves covered by angular, simple costae arising if stated otherwise); where used, open nomenclature at beaks; four median and eight lateral costae in ventral follows Feldman’s (1994) rules. Measurements of spe- valve (not known in dorsal valve).Ventral valve almost cimens assigned to Evanidisinurostrum saouraense sp. flat with obsolete, poorly defined sulcus; beak suberect. nov. are presented in Appendix 1 (see online supple- Dorsal valve (preserved only posteriorly) convex. mentary data at http://journals.cambridge.org/geo). Remarks. This specimen is assigned to Centrorhynchus Abbreviations. L, shell length; lc, lateral costae; mc, on the basis of its external morphology but additional median costae; T, shell thickness; W, shell width; Ws, material is required for a species-level identification. sulcus width. Centrorhynchus is a cosmopolitan genus assigned to

http://journals.cambridge.org Downloaded: 01 Aug 2016 IP address: 157.193.5.232 Famennian rhynchonellides from the Saoura Valley 1013 the Trigonirhynchiinae by Nicollin & Brice (2004); it occurs first in the lower Famennian (Balinski,´ 1995). In North Africa, the genus has been recognized in the east- ern part of the southern margin of the Basin by Brice (in Gourvennec, Bitam & Robardet, 1997) and on 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 to ammonoids from the Tamtert–Zereg section, our spe- cimen is of late Famennian age (do IV or IV/V, upper Platyclymenia Zone). Subfamily HEMITOECHIINAE Savage, 1996 Genus 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 in Brice 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, Member 3, Submember c. Description. Shell large (33.6 mm in width), dors- ibiconvex, wider than long, rounded subpentagonal in outline. Valves covered by simple, narrow and angular costae near beaks, becoming wider and subangular anteriorly and flattened on flanks; five median and eight ventral lateral costae (not observed on dorsal valve due to poor preservation); intercostal grooves narrower than costae. Ventral valve slightly convex in umbonal area; apical angle 112°; tongue not preserved; traces of short, divergent dental plates. Traces of dorsal median septum. Remarks. This single specimen is assigned to Pauro- Figure 4. Rhynchonellide brachiopods from the Marhouma gastroderhynchus presaharensis on the basis of its Formation, Upper Devonian (Famennian), Saoura Valley, Al- external features (shape and ornamentation). This geria (see Fig. 1). (a, b) Centrorhynchus sp., MUA/1094/020: species was reported previously from the upper incomplete specimen in ventral and posterior views. (c, d) Famennian deposits of Algeria (Timimoun region; Paurogastroderhynchus presaharensis Brice in Brice, Legrand- Sartenaer, 1975), Ahnet area (material from the Blain & Nicollin, 2005, MUA/1094/021: incomplete speci- Khenig Sandstones collected by M. Legrand-Blain; men in ventral and lateral views. (e–g) Tenuisinurostrum? sp., D. Brice, unpub. data) and of the Western Anti-Atlas MUA/1094/022: crushed specimen in ventral, dorsal and lateral views. (h–l) Sphaeridiorhynchus sp., MUA/1094/023: complete in Morocco (Kheneg Lakahal section; Brice in Brice, specimen in ventral, dorsal, lateral, posterior and anterior views. Legrand-Blain & Nicollin, 2005) where it is abundant. Scale bars: 5 mm. It is also represented by some remains from the upper Famennian Hassi Rharouar section, SE of Tinfouchy, and the Zemoul area (Morocco) (Brice in Brice, Type species. Camarophoria crenulata Gosselet, 1877, Legrand-Blain & Nicollin, 2005). The specimen lower Famennian, southern Belgium. from Ze 2/8 of the Tamtert–Zereg section is of late Famennian age (upper Platyclymenia Zone). Tenuisinurostrum? sp. Figure 4e–g Superfamily CAMAROTOECHIOIDEA Schuchert, 1929 Family LEIORHYNCHIDAE Stainbrook, 1945 Material. One incomplete, partly deformed specimen Subfamily LEIORHYNCHINAE Stainbrook, 1945 from Gara Diba, Marhouma Formation, Member 2, Genus Tenuisinurostrum Sartenaer, 1967 Submember b.

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Description. Shell of medium size (L ?, W = 19 mm, mens display median ribs arising in the anterior region T = 8.2 mm), suboval in outline, weakly dorsibicon- of the shell (mc = 0/3 and 5/?). Ventral valve inflated; vex; lateral commissures acute; anterior commissure beak erect; sulcus weak, arising in umbonal region or at uniplicate; anterior margin flat. Shell with rounded, ir- about mid-length, flat-bottomed; tongue nearly perpen- regular ribs present in anterior region, devoid of lateral dicular to commissure plane, low, trapezoidal. Dorsal ribs, with a weak ventral median groove originating in valve strongly inflated with greatest convexity slightly the umbonal region. Ventral valve with suberect beak; posterior to mid-length; fold weak, arising in umbonal sulcus arising at about mid-length, poorly defined, flat- area, flat-topped at front; septum lacking. bottomed at front, shallow (?); tongue very low; apical Remarks. These specimens are assigned to Sphaeri- angle 125°; possible traces of a dental plate left of diorhynchus on the basis of their markedly biconvex, the beak. Dorsal valve with greatest convexity slightly smooth and medium-sized shell with suborbicular to posterior to mid-length; fold weakly developed, low, subpentagonal outline, and by the weak fold and sul- flat-topped at front, originating at about mid-length; cus. They are clearly less globular than specimens of trace of a short median septum. its type species, but their specific assignment requires Remarks. Although poorly preserved, this single speci- additional material. men is tentatively assigned to Tenuisinurostrum mainly Until now, Sphaeridiorhynchus was known only on the basis of its suboval outline, its irregular ribs from the lower Famennian succession of Belarus and its low tongue. It displays a trace of a rudiment- (Pripyat’ Depression) and Ukraine (Dnepr–Donets De- ary dental plate, though such structures are not repor- pression) (Sartenaer, Pushkin & Kotlyar, 1997), where ted in this genus (see Sartenaer, 1967). Externally, it is represented by S. kuzmichiensis. Discovery of it is closer to T. crenulatum rather than to T. sub- Sphaeridiorhynchus sp. in the lower Famennian suc- crenulatum Biernat, 1970, which is more dorsibiconvex cession of the Saoura Valley thus represents the first and wider than long. Paromoeopygma Sartenaer, 1968 occurrence of this genus outside the Eastern European (Pugnacoidea) possesses dental plates and displays a Platform. similar outline in ventral view but differs by its more Superfamily PUGNACOIDEA Rzhonsnitskaia, 1956 inflated dorsal valve, its higher tongue and absence of Family PUGNACIDAE Rzhonsnitskaia, 1956 a dorsal median septum. Further material is therefore Genus Evanidisinurostrum Sartenaer, 1987 required for confident identification. The type species of Tenuisinurostrum was described Type species. Pseudoleiorhynchus? zemoulensis Drot, from the lower Palmatolepis crepida Zone (Sartenaer, 1964, lower Famennian, plains of Drah-el-Kelba, pre- 1984) in southern Belgium and northern France and T. Saharan Morocco. subcrenulatum Biernat, 1970 is known from the P. tri- angularis to P. marginifera zones in the Holy Cross Evanidisinurostrum saouraense sp. nov. Mountains of Poland (Biernat, 1970, 1983; Sarten- Figure 5a–t aer, 1985), though the latter needs to be revaluated Derivatio nominis. In reference to the Saoura Valley from the taxonomic viewpoint (Sartenaer, 1984, 1985, (Ougarta Basin, Algeria). 1987). Our badly preserved specimen of Tenuisinur- ostrum? sp. is of early Famennian age (do II) on the Holotype. A complete steinkern, MUA/1094/016. basis of Palmatolepis minuta subtilis (P. triangularis Locus typicus. Gara Diba section (30° 7 38 N; to P.trachytera zones). Tenuisinurostrum is not known 2° 12 30 W), Saoura Valley, eastern part of the Ou- with certainty from North Africa (cf. Sartenaer, 1984, garta Basin, Algerian Sahara. 1987). Stratum typicum. Marhouma Formation (lower Famen- Genus Sphaeridiorhynchus Sartenaer, Pushkin & nian), Member 2, Submember a. Kotlyar, 1997 Studied material. About 400 specimens (steinkerns), Type species. Sphaeridiorhynchus kuzmichiensis 125 well preserved (of which 42 were measured), 27 Sartenaer, Pushkin & Kotlyar, 1997, lower Famennian, decorticated, 21 incomplete, 227 juveniles and numer- Belarus. ous isolated valves and fragments from the Gara Diba Sphaeridiorhynchus sp. section. Figure 4h–l Diagnosis. A small species of Evanidisinurostrum (up to c. 17 mm in width) with shell subpentagonal in Material. Eight specimens including one juvenile from outline, wider than long, dorsibiconvex, widest near the Cheffar El Hamar (CA 1a) section, Marhouma mid-length or more anteriorly. Fold and sulcus start- Formation, Member 2, Submember a. ing posterior to mid-length. Tongue moderately high, Description. Shell of medium size (up to 20.7 mm subtrapezoidal in outline, perpendicular or almost per- in width), wider than long, widest near mid-length, pendicular to the commissural plane. Most of shells biconvex or weakly dorsibiconvex, suborbicular to sub- smooth with rounded median costae arising at or pos- pentagonal in outline; anterior commissure uniplicate, terior to mid-length (mc = 2–3/1–2, usually 2/1); lateral anterior margin straight. Shell smooth, but two speci- costae usually absent or rare (0–1/0–2).

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Figure 5. Rhynchonellide brachiopods from the Marhouma Formation, Upper Devonian (Famennian), Saoura Valley, Algeria (see Fig. 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 specimen in 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,ˇ 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. Scale bar: 5 mm.

Description. Shell small (up to 17.2 mm in width), 2, usually 2/1); lateral costae usually absent or rare dorsibiconvex, wider than long, widest near mid-length (0–1/0–2). Ventral valve gently inflated in both pos- or more anteriorly, subpentagonal in outline; anterior terior and lateral profiles with greatest convexity pos- commissure uniplicate, anterior margin, in ventral terior to mid-length; beak small; apical angle between view, indented by a strong median costa. Most shells 110° and 120°; sulcus wide, deep, originating at about are smooth on the flanks with rounded median costae mid-valve, with prominent median costa; tongue wider arising at or posterior to mid-length (mc = 2–3/1– than long, almost perpendicular to commissural plane,

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trapezoidal; dental plates absent; muscle field im- rounded median costae originating near umbones pressed, triangular in outline. Dorsal valve evenly con- (mc = 3/4), the central one bifurcating both in the sul- vex in both lateral and posterior profiles, highest at cus and on the fold; lateral costae badly preserved (lc front, impressed; fold originating at mid-length, relat- = 6/5–8), corresponding to the division of 2 or 3 costae ively flat-topped at front (apart from costae); no median present in the posterior region. Ventral valve with septum; myophragm short. greatest convexity near mid-length; beak small, erect; Dimensions in mm (n = 42 for all; see Appen- fold low, flat-topped at front, arising in posterior part of dix 1 in online supplementary data, available at the valve; tongue low, subtrapezoidal, perpendicular to http://journals.cambridge.org/geo). Width range 9.75– commissural plane. Dorsal valve with greatest convex- 17.2, average 12.5; length range 8.75–13.55, average ity posterior to mid-length; sulcus arising near umbo, 10.6; thickness range 4.2–7.1, average 5.5; width of well-defined, flat-bottomed at front. sulcus range 4–13.1, average 8.18; width/length ratio Remarks. These specimens share numerous external range 1.01–1.37, average 1.18; width/thickness ratio similarities with Hadyrhyncha hadyensis, such as their range 1.78–2.71, average 2.27; sulcus width/width ra- transversally elliptic outline, the similar development tio range 0.41–0.82, average 0.64. of their dorsal sulcus and ventral fold, their sharp Remarks. As all the specimens are steinkerns, it was not commissures and their ornament. However, as the in- possible to investigate the interiors by serial sections. ternal morphology of the Algerian material remains These specimens are assigned to Evanidisinurostrum unknown, they are tentatively compared to Havlícek’sˇ rather than Perrarisinurostrum Sartenaer, 1984 and (1979) species. They differ from Hadyrhyncha meridi- Tenuisinurostrum Sartenaer, 1967 (see Sartenaer, 1987 onalis Sartenaer, 1998a from Morocco by their smaller for discussion of the differences between these three size, their narrower apical angle, their ornament and genera) as they have similarities with E. zemoulense their different L/W, T/W and T/L ratios. They are sep- (Drot, 1964) such as the outline, the similar develop- arated from Hadyrhyncha sp. from Poland (Halamski & ment of fold and sulcus and the absence of dental plates Balinski,´ 2009) by their smaller size and less transverse and median septum. Nevertheless, the specimens from outline. the Saoura Valley can be discriminated from Drot’s Our specimens are associated with ammonoids of (1964) species by their smaller size, their maximum the upper Famennian Platyclymenia Zone (do IV or width near the mid-length or anteriorly, their different IV/V). Hadyrhyncha hadyensis is known from the up- W/L ratio, their larger apical angle and their different per or uppermost Famennian (do V or do VI according median ornament. to Havlícek,ˇ 1979) of Moravia (Czech Republic) and Evanidisinurostrum was previously known with cer- from the middle part of do V in south-easternThuringia tainty from the lower Famennian IIβ by its type species (Germany) (Bartzsch & Weyer, 1986). H. meridionalis in the Zemoul and south of Akka in the Dra Plains Sartenaer, 1998a occurs in the upper Famennian (do of pre-Saharan Morocco (Drot, 1964; Sartenaer, 1987) V) of southern Morocco (Dra Valley, Maider, Ta- and on the southern margin of the Tindouf Basin (Al- filalt, Zemoul); the genus is also reported from Po- geria; Drot, 1964). Furthermore, Sartenaer (1987,p. land (Holy Cross Mountains) in the upper Palmatolepis 135) suggested that Tenuisinurostrum subcrenulatum marginifera to upper Siphonodella praesulcata zones Biernat, 1970 from the Holy Cross Mountains in Po- (Halamski & Balinski,´ 2009). land (middle Palmatolepis triangularis to lower P. m a r - Genus Leptoterorhynchus Sartenaer, 1998b ginifera zones) may belong to Evanidisinurostrum,but this needs confirmation. Type species. Rozmanaria magna Biernat & Racki, 1986, middle Famennian, Holy Cross Mountains, Po- Family ROZMANARIIDAE Havlícek,ˇ 1982 land. Genus Hadyrhyncha Havlícek,ˇ 1979 Leptoterorhynchus sp. Type species. Hadyrhyncha hadyensis Havlícek,ˇ 1979, Figure 5z–ii upper Famennian, Moravia, Czech Republic. Material. Three specimens from the Marhouma Form- Hadyrhyncha cf. hadyensis Havlícek,ˇ 1979 ation, Member 3, Submember c of the Tamtert–Zereg Figure 5u–y (Ze 2/8) section and two from the Marhouma Forma- tion, Member 2, Submember b of the Cheffar El Ahmar cf. 1979 Hadyrhyncha hadyensis: Havlícek,ˇ p. 99, pl. (CA 1c, CA2) section. 2, figs 6–9, text-fig. 8. Description. Shell small (up to 13.1 mm in width), Material. Two specimens, of which one is incomplete, smooth, ventribiconvex, wider than long, widest about from the Marhouma Formation, Member 3, Submem- mid-length and transversally elliptical in outline; an- ber c, of the Tamtert–Zereg section (Ze 2/8, Ze 2/8b). terior commissure unisulcate; anterior margin straight. Description. Shell small (up to 15.7 mm in width), Ventral valve with greatest convexity near mid-length; wider than long, widest near mid-length, transversely beak small, erect; fold poorly developed, low, round- elliptic in outline, slightly ventribiconvex; anterior topped at front; tongue high, bent ventrally. Dorsal commissure unisulcate; anterior margin straight. Low, valve with greatest convexity near valve mid-length;

http://journals.cambridge.org Downloaded: 01 Aug 2016 IP address: 157.193.5.232 Famennian rhynchonellides from the Saoura Valley 1017 sulcus wide, originating in posterior part of the valve, In the Saoura Valley, Novaplatirostrum sp. is known shallow, flat-bottomed at front; tongue low, rounded, from the Tamtert–Zereg section where it is associated perpendicular to commissural plane or bent dorsally; with goniatites and clymeniids of late Famennian age in internal features not observed; fold poorly developed, levels Ze 2/8 and Ze 2/8b (upper Platyclymenia Zone) low, round-topped at front; no median septum. and in level Ze 2/9 (Gonioclymenia Zone). The age of Remarks. These specimens are assigned to Leptotero- the latter could be younger on the basis of a conodont rhynchus as they share many external similarities with fauna characteristic of the early upper Palmatolepis the type species L. magnus (Biernat & Racki, 1986) expansa Zone. N. sauerlandense occurs in deep-water such as the outline, the slightly developed and smooth environments in the German upper Famennian depos- sulcus and fold, and the sharp commissure. They prob- its (southern Sauerland and south-eastern Thuringia; ably belong to a new species which is not named be- Sartenaer, 1997) where it is associated with goniatites, cause of insufficient material. However, they can be dis- but also in the Holy Cross Mountains in Poland in levels criminated from the type species by their smaller size, of probable do V–VI age (Halamski & Balinski,´ 2009; their narrower apical angle, their different W/L, W/T A. Halamski, pers. comm., 2014). and T/L ratios, their not-so-well-defined sulcus and fold Genus Pugnaria Biernat & Racki, 1986 and the absence of a median depression on the fold. Type species. Pugnaria plana Biernat & Racki, 1986, Leptoterorhynchus magnus (Biernat & Racki, 1986) middle Famennian, Holy Cross Mountains, Poland. is known only from the middle Famennian deposits of the southern Holy Cross Mountains, Poland according Pugnaria sp. to Halamski & Balinski´ (2009). An unidentified species Figure 6h–o was reported in the Famennian (UD IIIC–IVB interval) Material. Three specimens (one incomplete adult and of north-western Sauerland in Germany by Sartenaer two juveniles) from the Tamtert–Zereg (Ze 2/8) sec- (1998b). In the Saoura Valley, Leptoterorhynchus sp. is tion (Marhouma Formation, Member 3, Submember c) known from the upper Famennian Tamtert–Zereg sec- and one adult from the Ouarourout (Ou 1/15) section tion on the basis of Platyclymenia annulata and from (Marhouma Formation, Member 4, Submember a). the middle Famennian (on the basis of Maenoceras) Cheffar El Ahmar section. Description. Shell small (19.2 mm in width), wider than long, flatly dorsibiconvex, transversely elliptical in Genus Novaplatirostrum Sartenaer, 1997 outline; anterior commissure rectimarginate; anterior Type species. Novaplatirostrum sauerlandense Sarten- margin slightly rounded. Shell smooth, growth lines aer, 1997, upper Famennian, north-western Sauerland, thin, irregularly distributed, observed on ventral valve. Germany. Ventral valve with greatest convexity posterior to mid- Novaplatirostrum sp. length; beak small, curved; sulcus lacking. Dorsal valve Figure 6a–j with greatest convexity posterior to mid-length; fold lacking. Material. Five specimens from the Tamtert–Zereg sec- tion (Ze 2/8, 2/8b, 2/9) and one from “Plat” between Remarks. Despite lack of knowledge about the internal Ouarourout and Gara Diba; Marhouma Formation, features, our specimens are externally very close to Member 3, Submember c. Pugnaria plana Biernat & Racki, 1986 but are smal- ler, devoid of a fold and display a wider apical angle. Description. Shell small (up to 16.2 mm in width), Additional material is required for accurate specific flatly biconvex, suborbicular in outline; anterior com- identification. missure uniplicate; anterior margin slightly rounded, with sharp commissures (juveniles uniplicate, flattly Pugnaria sp. occurs in do III–do IV in the Tamtert– biconvex, transversely subelliptical in outline). Low Zereg section (Ze 2/8) and is also known from the Ou- rounded median costae (3/2 or 4/3) near anterior com- arourout section (Ou1/15) where it is associated with missure; no lateral costae. Ventral valve with greatest ammonoids of probable do V age. In the Kowala sec- convexity in umbonal area; sulcus absent or wide tion (Holy Cross Mountains, Poland), the range of P. (Ws/W: 0.68; n = 1), very shallow, starting from mid- plana in terms of conodont zones spans the Palmato- length, flat-bottomed; tongue low, not perpendicular lepis marginifera to Siphonodella praesulcata zones to commissural plane; maximal apical angle of 127°. (Halamski & Balinski,´ 2009). Dorsal valve with greatest convexity posterior to mid- Genus Phacoiderhynchus Sartenaer, 2000 valve; fold obsolete, flat-bottomed. Type species. Phacoiderhynchus antiatlasicus Sarten- Remarks. The Algerian specimens are externally close aer, 2000, middle Famennian, Maider, Morocco. to Novaplatirostrum sauerlandense Sartenaer, 1997, especially paratypes F and G illustrated by Sartenaer Phacoiderhynchus aff. antiatlasicus Sartenaer, 2000 (1997, pl. 1, figs 31–40) but they are smaller and flatter Figure 6p–x than the latter and devoid of lateral costae. Our speci- aff. 2000 Phacoiderhynchus antiatlasicus n. gen., n. mens display less median costae, have an obsolete fold sp.: Sartenaer, p. 79–84, pl. 1, figs 1–30, pl. 2, figs and a smaller apical angle. 31–65.

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Figure 6. Rhynchonellide brachiopods from the Marhouma Formation, Upper Devonian (Famennian), Saoura Valley, Algeria (see Fig. 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: incomplete specimen in ventral, dorsal, lateral, posterior and anterior views. Scale bars: 5 mm.

Material. Eleven incomplete specimens from the median costae sometimes divided near the frontal mar- Tamtert–Zereg section (Ze 2/7: one adult and one gin (mc: 3–4/2–4); lateral costae not preserved. Vent- juvenile; Ze 2/8: six adults, one juvenile; Ze ral valve with greatest convexity close to mid-length; 2/8b: two adults), Marhouma Formation, Member 3, beak very small; sulcus shallow, flat-bottomed at front; Submember c. tongue very low, not perpendicular to commissural plane. Dorsal valve with maximum convexity near mid- Description. Shell of medium-size (up to 29 mm in valve; fold scarcely developed, originating anterior to width), wider than long, flatly biconvex, transversally mid-length. elliptical in outline, widest at mid-length; commissure sharp; anterior commissure uniplicate; anterior margin Remarks. These incomplete specimens are tentatively flattened. Shells mostly smooth with rounded and wide assigned to Phacoiderhynchus antiatlasicus Sartenaer,

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2000 on the basis of their external morphology, al- 4.b. Palaeoecology and taphonomy though they can be distinguished by their smaller size, Brachiopods obtained from several horizons of the Fa- their sulcus and fold starting a little more anteriorly mennian part of the Marhouma Formation include, and their less numerous median costae (the lateral ones in order of abundance, rhynchonellides (439 spe- are absent or not preserved). The specimens from the cimens), orthides (small Aulacella; 45 specimens), Saoura Valley have a narrower apical angle (134–145° athyridides (medium-size Composita; 4 specimens) versus 147–157°) and may represent a new species, but and spiriferides (only one small specimen; affinities specific assignment requires additional material. not apparent due to poor preservation). Rhynchonel- Phacoiderhynchus aff. antiatlasicus is known from lides therefore represent 90 % of the whole brachiopod the upper Famennian Tamtert–Zereg section on the assemblage, with 10 species. For comparison, 197 am- basis of ammonoids (upper Platyclymenia Zone). P. monoids (goniatites and clymeniids; D. Korn and F.Z. antiatlasicus occurs with certainty in middle–upper Fa- Malti, unpub. data) and 19 trilobites (Crônier et al. mennian IIIB to IVB deposits from southern Morocco 2013), including the material from two other sections (Maïder and eastern Draa Plains) according to Sarten- of the Saoura Valley (Béchir and Idhir; see location aer (2000), and probably in the Immouzer du Kandar in Crônier et al. 2013), have been collected in the area south of Fès (central Morocco; see Brice et al. same levels of the Marhouma Formation. Within these 1984). lithostratigraphic 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- 4. Discussion pansa Zone, i.e. at the base of the upper Famennian deposits (Fig. 2). Development of the ‘griotte’ lime- 4.a. Stratigraphic summary and general comments on stones took place during a sea-level rise (F.Z. Malti, brachiopod diversity during Famennian time unpub. data) that probably facilitated faunal exchanges The stratigraphic range of the rhynchonellides within between distant basins. the Famennian part of the Marhouma Formation across The low-diversity brachiopod fauna is clearly an in the Saoura Valley (Ouarourout, Gara Diba, Cheffar situ rhynchonellide-dominated brachiopod community Amar and Tamtert–Zereg sections) is presented in with much lower diversity of other orders; the spe- Figures 2 and 3. cimens are not disarticulated, sorted or broken. No As the Frasnian–Famennian (F–F) boundary, and strophomenides or productides occurred in association thus the position of the Upper Kellwasser Event (Pal- with the rhynchonellides. Deep-water conditions are matolepis linguiformis Zone), is not precisely estab- indicated by cephalopods (goniatites and clymeniids) lished within the Marhouma Formation, it is difficult and trilobites; the latter comprise blind to reduced- to discuss in detail the post-Kellwasser recovery of the eyed phacopides (Crônier et al. 2013). One of the brachiopods in this part of the Ougarta Basin. Note that most striking features of this rhynchonellide fauna is in regions where environmental conditions were more the predominance of smooth, thin-shelled and small favourable to brachiopods, such as the Namur–Dinant forms (except some scarce large Trigonirhynchiidae), Basin in southern Belgium and the Polish epicontin- also characterized by flattened shells. Moreover, among ental basin, their recovery started in the Palmatolepis these rhynchonellides, leiorhynchids (Camarotoech- triangularis Zone very soon after the F–F boundary, ioidea) and rozmanariids (Pugnacoidea) are by far the in which athyridides, rhynchonellides and spiriferides most abundant elements as is generally the case in late played a significant role (Balinski´ 1996, 2002; Mot- Palaeozoic dysaerobic communities, though the Pug- tequin 2008a, b). On a worldwide scale, brachiopod nacoidea are represented by other families (cf. Bowen, diversity declined significantly during Frasnian time Rhoads & McAlester, 1974; Biernat & Racki, 1986; with a significant reduction in generic diversity; this Racki, 1989; Alexander, 1994; Mottequin & Legrand- was accentuated during Famennian time, possibly due Blain, 2010). We also draw attention to the occurrence to increasing faunal cosmopolitanism related to major of a monospecific assemblage with Evanidisinurostrum sea-level changes and other parameters that suppressed saouraense in which immature specimens are particu- faunal barriers and allowed interchange between re- larly abundant. Almost monospecific rhynchonellide mote areas (e.g. Copper, 1998; Brice et al. 2000). From assemblages of leiorhynchids have been reported else- the rhynchonellide viewpoint the Famennian Stage is where, notably in the Frasnian deposits of southern marked by great generic diversity, contrasting with Belgium (e.g. Sartenaer, 1974) and Poland (Racki, other brachiopod orders (e.g. orthides); this is similar 1993; Racki & Szulczewski in Sartenaer, Racki & to the situation reported during Emsian time (Curry & Szulczewski, 1998). Overrepresentation of juveniles Brunton, 2007) and the occurrence of numerous cos- of E. saouraense is probably related to the combina- mopolitan genera, including some of those identified tion of several biological (e.g. nutrient availability) and in the Marhouma Formation (see discussion in Section physicochemical controlling factors (including water 4.c). This Devonian stage is also characterized by the oxygenation and depth) (see Pérez-Huerta & Sheldon, predominance of the rhynchonellides within brachio- 2006) that induced significant losses among immature pod faunas (Curry & Brunton, 2007). specimens.

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The state of preservation of our material does not these Moroccan occurrences, insisting that no particu- permit precision as regards the size of the foramen lar environmental conditions can be discerned. for each species, but adults of the identified genera Famennian brachiopods from western were are characterized by a reduced foramen (cf. data in described by Havlícekˇ (1984), Havlícekˇ & Röhlich Savage et al. 2002). We are therefore inclined to re- (1987), and Mergl & Massa (1992). They came from gard the rhynchonellides as lying unattached on the sea shallow-water environments characterized by silici- floor. Another significant feature is the aequibiconvex- clastic deposits with benthic communities dominated ity of the shells, notably for Pugnaria, Novaplatisin- by lingulide and rhynchonellide brachiopods (Mergl urostrum and Phacoiderhynchus. According to Walker & Massa, 2000). The Libyan rhynchonellides include (1974) this is indicative of soft substrates; it was in- species of Cupularostrum Sartenaer, 1961 and Liby- terpreted in this way by Biernat & Racki (1986)for aerhynchus Mergl & Massa, 1992 that are coarsely Famennian rhynchonellides. Relatively more biconvex ribbed, and thus markedly different from those occur- shells such as those of Sphaeridiorhynchus may accord ring in contemporaneous deep-water environments of with a firmer substrate (see Racki & Szulczewski in Morocco and Algeria. As rightly reported by Mergl & Sartenaer, Racki & Szulczewski, 1998). According to Massa (1992) and Mergl, Massa & Plauchut (2001), the Racki (1998), the deep-water rhynchonellide biofacies stratigraphic range of Cupularostrum, originally sug- displays apparent continuity in spite of generic losses gested as Givetian, should be reconsidered. across the Frasnian–Famennian boundary. One of the Four genera known from deep-water environments major changes in this biofacies is the huge development are recognized for the first time in North Africa: of the rozmanariids during Famennian time. They made Pugnaria, Leptoterorhynchus, Sphaeridiorhynchus and their appearance during Pragian time (cf. Savage et al. Novaplatirostrum. They were previously known only 2002; Savage, 2007), before disappearing at the end from the south-eastern margin of Laurussia and were of the Devonian period (Mottequin, Brice & Legrand- viewed as being restricted geographically: Leptoter- Blain, 2014). orhynchus (Germany and Poland; Sartenaer, 1998b), Novaplatirostrum (Germany and Poland; Sartenaer, 1997; Halamski & Balinski,´ 2009), Pugnaria (Po- 4.c. Comparison with other rhynchonellide faunas from land; Biernat & Racki, 1986), and Sphaeridiorhynchus North Africa and biogeographic affinities (Belarus and Ukraine; Sartenaer, Pushkin & Kotl- Despite the increased number of publications, Famen- yar, 1997). Tenuisinurostrum, previously reported from nian brachiopod assemblages from North Africa, and western and possibly eastern Europe (cf. Sartenaer, therefore from the northern margin of Gondwana, 1987), is here reported from the Famennian Marhouma are still poorly documented. This is due in part to Formation, but needs confirmation on the basis of well- most papers being generally focused on a particular preserved material. group, as with this paper. Taxonomic data are available Halamski & Balinski´ (2013) recently stressed the for most of the important brachiopod orders present great similarities between Middle Devonian benthic in the Famennian deposits of Algeria and Morocco faunas from the northern and southern shores of the (see below for Libyan data): productides (e.g. Brous- Variscan Sea, which accords with a narrow oceanic miche, 1975; Legrand-Blain, 1995a, b; Legrand-Blain zone between Laurussia and Gondwana (McKerrow in Brice, Legrand-Blain & Nicollin, 2005), spiriferides et al. 2000) rather than an extensive Rheic Ocean (e.g. Drot, 1964; Nicollin & Brice, 2000; Brice & (e.g. Torsvik & Cocks, 2011, 2013; Stampfli et al. Nicollin in Brice, Legrand-Blain & Nicollin, 2005), 2013). Legrand-Blain (1995a) also noted clear af- and rhynchonellides. finities between the productides, rhynchonellides and Five rhynchonellide genera, namely Centro- spiriferides of North Africa and Southern Europe with rhynchus, Evanidisinurostrum, Hadyrhyncha, Pauro- those from Armenia and the Urals across the Devonian– gastroderhynchus and Phacoiderhynchus, occurring Carboniferous boundary. Close affinities between the within the Marhouma Formation were previously repor- Famennian rhynchonellides from central Algeria and ted from Famennian successions on the northern mar- those from western and eastern Europe is consistent gin of Gondwana (Morocco and Algeria). Attention is with the hypothesis of prolonged proximity between drawn to two salient absences from the rhynchonellides both continental masses, but island arcs between Laur- of the Marhuma Formation: (1) Tetragonorhynchus ussia and Gondwana (Stampfli et al. 2013; Torsvik & Sartenaer, 1999, reported from the upper Famennian Cocks, 2013) could have facilitated dissemination and (UD V) deep-water facies from southern Morocco interaction of the brachiopod faunas on both sides of (Maïder) (Sartenaer, 1999); and (2) Dzieduszyckia the Rheic Ocean. Siemiradzki, 1909 represented by two species in the Famennian deposits of the Middle Atlas of Morocco 5. Conclusions (Balinski´ & Biernat, 2003). Dzieduszyckia has been considered to be typical of Upper Devonian vent-seep In the Saoura Valley (Ougarta Basin, Algeria), the environments (Campbell & Bottjer, 1995), but Balinski´ Famennian part of the Marhouma Formation (mem- & Biernat (2003) have challenged this judgement on the bers 2–4) contains rhynchonellide-dominated brachi- basis of geochemical analyses of rocks associated with opod assemblages in which a generally low diversity

http://journals.cambridge.org Downloaded: 01 Aug 2016 IP address: 157.193.5.232 Famennian rhynchonellides from the Saoura Valley 1021 of small athyridides (Composita), orthides (Aula- BARTZSCH,K.&WEYER, D. 1986. Biostratigraphie der cella) and spiriferides occur. The rhynchonellides, lei- Devon/Karbon Grenze im Bohlen-Profil bei Saalfeld orhynchids and rozmanariids are the most abundant (Thüringen, DDR). Zeitschrift für Geologische Wis- and the most diverse elements, represented by small, senschaften 14, 147–52. BECKER,R.T.,ElHASSANI,A.&TAHIRI, A. (eds) 2013. thin-shelled species. These brachiopods are generally International Field Symposium ‘The Devonian and found in association with cephalopods and blind or Lower Carboniferous of northern Gondwana’. Docu- eye-reduced phacopide trilobites (Crônier et al. 2013), ments de l’Institut Scientifique, Rabat 27, 1–150. interpreted to have lived in deep, oxygen-depleted BECKER,R.T.,HARTENFELS,S.,ABOUSSALAM,Z.S., environments. Such rhynchonellide assemblages are TRAGELEHN,H.,BRICE,D.&ElHASSANI, A. 2013. The interpreted as being characteristic of dysaerobic en- Devonian–Carboniferous boundary at Lalla Mimouna vironments found on epicontinental platforms during (northern Maider) – A progress report. Documents de l’Institut scientifique de Rabat 27, 109–17. Late Devonian time, especially in Laurussia and North BENHAMOU, M., ABBACHE,A.,ELMI,S.,MEKHALI, Gondwana. The leiorhynchid Sphaeridiorhynchus and L., RACHEBŒUF,P.R.,OUALI-MAHADJI,A.& the rozmanariids Leptoterorhynchus, Pugnaria and BOUMENDJEL, K. 2004. Les calcaires ‘griottes’ et faciès Novaplatirostrum are reported for the first time out- associés du Dévonien supérieur des environs de Beni- side Europe; their occurrence on the margin of Gond- Abbes au Djebel Heche (Saoura, Algérie): environne- wana reinforces the biostratigraphic value of these four ments et implications paléogéographiques. Bulletin du genera and underscores the global character of the Fa- Service Géologique d’Algérie 15, 27–49. BIERNAT, G. 1970. Lower Famennian brachiopods from Holy mennian benthic faunas. Cross Mountains, Poland. Acta Palaeontologica Polon- ica 15, 33–62. Acknowledgements. We are greatly indebted to Dieter BIERNAT, G. 1983. On the Famennian brachiopods from Korn for having provided identifications of the cephalopods Jabłonna, Góry Swi˛´ etokrzyskie Mts., Poland. Biuletyn cited in the text, to Thierry Hubin for technical assistance Instytutu Geologicznego 345, 137–54. and to Adam T. Halamski, Andrzej Balinski´ and John Talent BIERNAT,G.&RACKI, G. 1986. A rhynchonellid-dominated for reviewing the manuscript. This paper is a contribution late Famennian brachiopod assemblage from the Holy to the International Geoscience Programme (IGCP) Project Cross Mountains (Poland). Acta Palaeontologica Polon- 596, Climate change and biodiversity patterns in the Mid ica 31, 85–109. Palaeozoic. 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