Annales Societatis Geologorum Poloniae (2013), vol. 83: 243–307.

MIDDLE DEVO NIAN BRACHIO PODS FROM THE SOUTH ERN MADER (EASTERN ANTI- ATLAS, MO ROCCO)

Adam T. HALAM SKI & Andrzej BALIÑSKI

Insti tute of Paleo biology, Polish Academy of Sciences, Twarda 51/55, PL-00- 818 Warszawa, Poland; e-mails: [email protected], [email protected]

Ha lam ski, A. T. & Bali ñski, A., 2013. Mid dle De vo nian brachio pods from the southern MaÎder (eastern Anti-Atlas, Mo rocco). An nales So cie ta tis Ge olo go rum Po lo niae, 83: 243–307.

Ab stract: Sixty-two Middle De vonian brachiopod species are described on the basis of >1300 speci mens from the Tabou ma khloõf Forma tion (and sub or di nately proba bly also from the top of the El Ot fal For mation) at Ma- dÀne el Mrakib (middle to late Eifelian or early Givetian?), Afer dou el Mrakib (late Eifelian to middle Givetian), and Guelb el Ma harch (early Giv etian) on the south ern edge of the MaÎder Syn cline (eastern Anti-Atlas, Mo - rocco). Rep re sen ta tives of Cra niida (2 taxa), Stro phomen ida (5), Pro duc tida (2), Or tho tet ida (2), Or thida (5), Pentame rida (5), Rhynchonel lida (9), Atrypida (14), Athyridida (7), Spiriferida (9), and Spirif er inida (2) are pres ent. The fauna is domi nated quan ti ta tively by the Atrypida (24% of taxa, about the half of speci mens, and the com mon est spe cies Atry paria dis persa making up about 13% of the mate rial); the Rhyn chonel lida (16% of taxa, about one- sixth of speci mens) are the sec ond larg est or der. A new ge nus of the fam ily Pugnaci dae (or der Rhyn- chonel lida), Pau linaer hynchia, is pro posed with the type spe cies P. pau li nae gen. et sp. nov. from Maharch; it is clos est to Pug nax, from which it dif fers in dis tinct costa tion, lack of a dor sal sep tum and sep talium, and ru di men- tary den tal plates. Desquama tia (D.) deserti sp. nov., a large and finely co state rep re sen ta tive of the ge nus, is described also from Ma harch. An tirhyn chonella and Glosshy po thy ri dina are re ported for the first time or con - firmed to be pres ent in the Giv etian. Forty (possi bly up to 46) spe cies (71 or pos si bly up to 82% of the taxa iden ti fied at the spe cies level) are pres ent also in ei ther Eifel (Ger many) or the Holy Cross Mountains (Po land). Such a high ra tio of spe cies in com mon at tests to un con strained fau nal ex changes among ben thic fau nas be tween the north ern and south ern shores of the Varis can Sea dur ing the Mid dle De vo nian. This fa vours the palaeo geo - graphic hy pothe sis of a nar row Variscan Sea.

Key words: sys te mat ics, Brachio poda, Mo rocco, De vo nian, Eife lian, Giv etian, Varis can Sea, pa laeo bio ge o- g ra phy.

Manu script re ceived 11 March 2013, ac cepted 27 December 2013

IN TRO DUC TION His tory of re search

The rich ness of Palaeozoic fau nas in the Mo roc can Geological stud ies in the eastern Anti-At las began only Anti-Atlas is famed. Devo nian brachi o pods, the major fau- in the 1930s. As far as it was pos sible to estab lish, the first nal com po nent in sev eral geo log i cal units, are es pe cially nu - palaeontologist to de scribe De vo nian brachi o pods from mer ous and di ver si fied. How ever, de spite sev eral stud ies by MaÎder was Dorothée Le Maître. On the ba sis of Geor ges pre vi ous au thors, Mid dle De vo nian brachi o pods still are re- Choubert’s collec tions, she reported a small faunule from latively poorly known, owing to, among other factors, col- “Merakib” in the MaÎder Syncline (Le Maître, 1939). Al- lections having been assem bled in large part for purposes of though the bulk of the fauna is Early Devo nian in age, a part geolog i cal mapping. The aim of the present pa per is (i) to of it might belong to the Middle Devo nian. The de tailed pres ent a de tailed system atic account of a middle Eifelian to study of this fauna, announced in the above-mentioned pa- middle Givetian brachio pod fauna from three nearby local i - per, has never ap peared. ties in the southern MaÎder; (ii) to de fine its palaeobiogeo- The sys tem atic study of De vo nian brachi o pods from graphic af fini ties, which lie, as will be shown, within the MaÎder is mainly the work of geol o gist Henri Hollard, who Rhenish subprovince; (iii) last, to discuss the im pact of the collected exten sively in Morocco for over twenty years new data upon questions of the width of the Variscan Sea (Hollard, 1962, 1963, 1974, 1985) and of palaeontologist and of the sepa ra tion between Rhenish- and Ameri can-type Jeannine Drot. In a series of papers dealing with Si lurian brachi o pod fau nas in the Mid dle De vo nian of north west ern and De vo nian brachi o pods of the pre-Sa ha ran Mo rocco Af rica. (Anti-Atlas and northern flank of the Tindouf Syncline), 244 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 245

Drot (1964, 1966, 1971) described several taxa of Middle scriptions: Jebel Issoumour, Jbel Issimour; includ ing the Devo nian rhynchonellides and spiriferides from the MaÎder solitary hill called Jebel Abiir), Jebel (el) Otfal, Jebel (includ ing Aferdou el Mrakib). Middle Devo nian stropho- Maharch, Jebel (el) Mrakib, and Jebel OufatÀne (Fig. 1C). menides, productides, orthotetides, orthides, pentamerides, The present paper is based on the mate rial from three out- athyridides, and atrypides have not been under detailed re- crops, sit u ated in the south ern part of these ranges (geo logic search. map sheet Todrha-Ma’der, du Dresnay et al., 1988), namely Struve (1995) il lustrated and briefly com mented upon MadÀne el Mrakib, Aferdou el Mrakib and Guelb (el) sev eral Mid dle De vo nian pentamerides, atrypides, and Maharch. spiriferides from the northern MaÎder (Jebel Issomour). He In the Mid dle Devo nian (Fig. 1A), the present Anti-At- also listed sev eral other taxa (identi fied at ge nus level) and las region formed a part of the pas sive north-western margin con cluded that the af fili a tion of the fauna was re fer able to of Gondwana (Piqué et al., 1993) at a palaeolatitude of about the Rhenish subprovince. Single Givetian brachio pod spe- 30–40° S (Scotese and McKerrow, 1990; Matte, 2001; Go- cies from MaÎder were described by Godefroid (2000), lonka et al., 2006; Torsvik and Cocks, 2011; Stampfli et al., Schemm-Gregory (2009), and Franchi et al. (2012). 2013). In general, the eastern part of the Anti-Atlas is charac - Last but not least, it is appro pri ate to note some major ter ised during the entire Devo nian by sedi ments with rel a- results regard ing the geol ogy of the study area. The age of tively low content of clastic and terrigenous compo nents (as the coral limestone (calcaire coralligÀne), now corre spond - compared to the west ern part; Piqué and Michard, 1989). ing to a part of the Taboumakhloõf For ma tion (Hollard, The subdi vi sion of the eastern Anti-At las into four deposi- 1974), was estab lished as Givetian by Hollard (1962; con- tional realms (from west to east, the MaÎder Platform, the sidered as Eifelian by Delépine, 1951). The strati graphic de- MaÎder Ba sin, the Tafilalt Plat form, the Tafilalt Ba sin) re - scription of the MaÎder was presented by Hollard, first in sulted from bathymetric differ en ti a tion of the basin, due to brief (Hollard, 1963) and then in detail (Hollard, 1981). A early Variscan block faulting in the late Emsian (Wendt, sedimentological study of the three sites, from which the 1985, 1988; Belka et al., 1997). Aferdou el Mrakib and fauna described in the present paper comes, was done by Guelb el Maharch are the two larg est bioconstructions, sev- Kaufmann (1998). Cono dont-based stratig raphy of the eral of which devel oped in a wide neritic facies belt, surrou- Eifelian of the eastern Anti-At las was pre sented by Belka et nding the central basinal part of the MaÎder Basin (Wendt, al. (1997). The Devo nian palaeo geogra phy of the Anti-At- 1993; Kaufmann, 1995). It may be noted that simi lar early las was dealt with by Wendt (1985, 1988, 1993), by Belka et Givetian reefs and mud mounds devel oped on both sides of al. (1997), and by Döring and Kazmierczak (2001). the Variscan Ocean: on the northern borders of Gondwana further west (pres ent-day Western Sa hara: Dumestre and Illing, 1967; Wendt and Kaufmann, 2004) and east (Wendt GEO LOGIC SET TING et al., 1997), as well as on the southern borders of Eurame- rica (present-day Sauerland: Krebs, 1967; May, 2003; pres- The MaÎder (spelt also Ma’der, Mader, Maader, and ent-day Eifel: Schmid et al., 2001; present-day New York: MaÎdÀre) is a Variscan syncline, sit u ated in the east ern part Heckel, 1972; see also Krause, 2004). of the broadly named Anti-Atlas (i.e., includ ing Jebel The middle part of the Mid dle Devo nian in the southern Sarhro and Jebel Ougnate; Joly et al., 1949, p. 30; Choubert MaÎder (Fig. 2) was divided into two major units (forma - and Mar¸ais, 1956; Joly, 1962; Michard, 1976, p. 41; Piqué, tions) by Hollard (1974). The ages given below, mod ified 1994, p. 26), itself a transi tional zone between the stable do- accord ing to the results of Kaufmann (1998) in respect to main of the West Afri can craton in the south and the Varis- Hollard (1974), are as follows: can chain in the north (Piqué et al., 1993; see also Ennih and 1. upper Eifelian (aus tra lis Zone) shal low-water crino- Liégeois, 2008; Soulaimani and Burkhardt, 2008). The idal grainstones (top of the El Otfal Forma tion) cropping Arabic noun maÎder denotes a fine-grained allu vial plain out on the eastern flank of Aferdou; (Dresch et al., 1952, p. 171; Joly, 1962, p. 546) and in this 2. upper Eifelian to lower Givetian (kockelianus to Lo- case describes the eroded Carbon ifer ous core of the syn- wer varcus zones) mound debris and off-mound strata crop- cline, cov ered by Qua ter nary sed iments (Daya el MaÎder, ping out on western, northern, and partly eastern flanks of Joly, 1962, p. 416, 537; corre spond ing to Mader el Kebir in Aferdou (Taboumakhloõf Forma tion; about 45 m in thick- Fig. 1C). This plain is bordered all round by low moun tain ness); this is the TM 554 lo cal ity of Drot (1971) and Hollard ranges that are predom i nantly Devo nian, listed clockwise, (1974, 1981, p. 28). This forma tion includes the lower Gi- begin ning from the northwest: Jebel Issomour (vari ant tran- vetian Ivdelinia pulchra lumachelle (thickness about 3 m;

Fig. 1. Geo log i cal set ting of fauna stud ied. A. Sit ua tion of study area and selected other im por tant brachi opod fau nas on a palaeogeo - graphic re con struc tion for the Mid dle De vo nian [mod i fied af ter Scotese and McKerrow (1990), Golonka et al. (2006), and Murphy et al. (2011)]. B. Sche matic geo logic map of north west ern Af rica (much sim pli fied af ter Choubert and Faure-Muret, 1988), showing ar eas men - tioned in the text. (Due to map scale in some cases “Mid dle De vonian” stands for both Mid dle and Up per De vonian, and “Lower De vo- nian” for all three Devo nian series.). C. Geo logic map of the MaÎder [modi fied and sim pli fied af ter du Dresnay et al. (1988), Hollard (1974, 1985), and Destombes and Hollard (1986)]. Names of the three sites studied in the pres ent pa per are in cap i tal bold face. Geo log i cal struc tures, moun tain ranges, and other top o graphic fea tures in cap i tals. Wa ters (seas in A and B, wa dis in C) in ital ics. Towns and vil lages in Ro man type face 246 A. T. HALAMSKI & A. BALIÑSKI

Fig. 2. Stratigraphic sections of upper Emsian to lower Givetian strata at the three local ities of the MaÎder Syncline (eastern Anti-Atlas) studied, show ing the ap prox i mate po sition of the ho ri zons, from which brachi o pod fauna has been col lected (black ver ti cal ar rowed lines). Strati graphic and lith o logic data af ter Hollard (1974) and Kaufmann (1998)

Tessitore et al., 2013) cropping out in some points of the multiplicata (>120 speci mens). Atrypa (Planatrypa?) con- east ern flank. fusa is also common (>30 speci mens). The fauna studied co mes from three lo cali ties, situ ated Aferdou el Mrakib (variant transcrip tion: Aferdou n’ within 20 km of each other. A descrip tion of them is given Merakib; in the system atic part abridged as Aferdou) is a flat- in the next chapter. topped mountain, about 900 m in diam e ter and up to 130 m high (Kaufmann, 1998, pls 1, 2), situ ated on the northern flank of the Jebel Mrakib at about 30°46'N 4°37'W. The gen- MA TE RIAL AND METH ODS eral geol ogy was presented by Hollard (1974, p. 24–28, 48– 49) and a de tailed sedimentological ac count of this and the De scrip tion of lo cal i ties and gen eral char ac ter is tics follow ing out crop was given by Kaufmann (1998) and is fol- of their fauna lowed here. Colo nial rugose corals and stromatoporoids are MadÀne el Mrakib denotes a contin u ous section of (at dom inant builders of this largest reef of the east ern Anti-At - least) Lower to Mid dle Devo nian strata sit u ated at ap proxi - las. Rich brachio pod collec tions (833 speci mens in total) mately 30°44'N 4°41'W (ca. 2 km NE from the cop per mine). were assem bled, mostly from the Taboumakhloõf Formation The authors used rich col lec tions (>200 speci mens) made (Fig. 2). The most frequent brachio pods are atrypides, by U. Jansen (SMF) in 1998 from the Drotops beds (Fig. 2; which consti tute the domi nant order with 12 taxa and over Eifelian, ac cording to Struve, 1990; a Drotops ho ri zon is 400 spec i mens, in par tic u lar, the com mon est spe cies Atry- lower Givetian in age at Aferdou, accord ing to Kaufmann, paria dispersa repre sented by 125 speci mens, Atrypa (Pla- 1998; however, most proba bly there is more than one Dro- natrypa?) confusa with >90 speci mens, and Kerpina vineta tops ho ri zon in the Mid dle De vo nian) char ac ter ised by com - with 77 speci mens. The rhynchonellide Kransia parallele- mon occur rence of the above-mentioned phacopid trilo bite. pipeda (91 speci mens) and the lumachelle-forming penta- Another collec tion (172 spec imens) from the same section meride Ivdelinia pulchra (123 spec imens) are also frequent. was made in Febru ary and March 2000 from an exca va tion It should be noted that brachio pods were collected mostly as at 30°44'57"N 4°41'42"W (“kegelförmiger Berg, Drotop- weath ered spec imens from the scree on the flanks of Afer- sschurf”) by the late Volker Ebbighausen. It has been do- dou el Mrakib. Accord ingly, in most cases, the detailed stra- nated to the Nat u ral History Museum in Berlin and could be tigraphic posi tion of single taxa cannot be given. Except for studied by the present au thors, although only at a very ad- pentamerides, col lected from the lower Givetian lumachelle vanced stage of prepa ra tion of the manuscript. Better out- (Tessitore et al., 2013), it was im possi ble to make any dis- crops of the same level were described from the northern tinction between speci mens, eroded from the Eifelian and MaÎder by Struve (1995). These strata are char acter ised by Givetian parts of the Taboumakhloõf For mation. The lat ter the fre quent oc cur rence of the pentameride Glyptogypa might be more numer ous, but it should be stressed that the MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 247

sam ple from Aferdou does contain taxa, reported up to now as microornamentation is never preserved, while the inte ri ors being both solely from the Eifelian (e.g., Plectospira ferita and are al ways bro ken and/or recrystallised. For ex am ple, in ter - Phragmophora schnuri) and solely from the Givetian (e.g., pret ing the in ter nal struc tures of Paulinaerhynchia paulinae Kransia subcordiformis and Spinatrypa trigonella). gen. et sp. nov. re quired the section ing of five speci mens. Guelb el Maharch (vari ant transcrip tion: Guelb Mharch; The pres er va tion of brachi o pods com ing from MadÀne el in the sys tematic part abridged as Maharch) is a sol itary Mrakib is quite sim i lar to those from Aferdou, at least so far dome-like hill, about 45 m high (Kaufmann, 1998, pl. 4: 1) as the ex teri ors are con cerned (se rial sec tions were not made and situ ated on the plain (about 30°47'N 4°33'W) near on speci mens from this local ity). Mate rial from the ex cava - Foum el MaÎder, which is the gorge of Oued Chouiref, con- tions, made by the late V. Ebbighausen, is partic u larly well stitut ing the border between Jebel Maharch in the east and pre served. Jebel Mrakib in the west. It con sists solely of mas sive lime - stone of early Givetian (Fig. 2; hemiansatus to Lower var- Re pos i to ries cus zones; Kaufmann, 1998) age. Tab u lates and cri noids are quite com mon, while colo nial rugosans are ab sent (Kauf- The described mate rial (>1300 speci mens) is kept in the mann, 1998). Charac ter is tic (i.e., most frequent and occur - Insti tute of Paleobiology of the Polish Academy of Sciences ring only at this local ity) brachio pods of Guelb el Maharch (War saw, Po land) un der the col lec tion num ber ZPAL Bp 68 (106 speci mens at the authors’ disposal) include Paulinaer- (68/1, Aferdou and 68/2, Maharch; the au thors’ and B³a¿ej hynchia paulinae gen. et sp. nov. (21 speci mens) and Des- Berkowski’s collec tions), in the Senckenbergische Natur- quamatia (D.) deserti sp. nov. (40 speci mens). Given the forschende Gesellschaft in Frankfurt am Main (MadÀne el geologic setting de scribed above, all the mate rial col lected Mrakib; Ulrich Jansen’s col lec tion; in sti tu tional ab bre vi a- at this lo cal ity comes from the upper part of the Tabouma- tion SMF), and in the Museum für Naturkunde in Berlin khloõf Forma tion and is undoubt edly early Givetian in age (MadÀne el Mrakib, Volker Ebbighausen’s collec tion; insti - (Kaufmann, 1998). tu tional ab bre vi a tion MB). Ad di tional ma te rial from the in - Out of the twenty species found at MadÀne el Mrakib, sti tu tions listed be low also was used: BGR, Bundesanstalt eight (40%) have not been found at the other two sites. für Geowissenschaften und Rohstoffe, Berlin-Spandau, Ger- However, the gen eral aspect of brachi o pod faunas at Ma- many; GFCL, Faculté libre des Sci ences et Tech nol o gies, dÀne el Mrakib and at Aferdou is not very differ ent, the co- Lille, France; HLMD, Hessisches Landes-Mu seum, Darm- mmon species being in part the same (Xystostrophia umbra- stadt, Ger many; MCZ, Mu seum of Com par a tive Zo ol ogy, culum, Atrypa confusa). Taxa com mon at MadÀne and very Harvard Univer sity, Cam bridge, Mass., USA; MGUWr, Mu- rare or absent at Aferdou include Glyptogypa multiplicata zeum Geologiczne Uniwersytetu Wroc³awskiego, Wroc³aw, and Quiringites arensentiae; their presence is proba bly due Poland; and MNHN, Muséum national d’Histoire naturelle, to older strata cropping out at the former lo cal ity. Out of Paris, France. eleven species found at Maharch, five (45%) are known Spec imens of de scribed spe cies from other sec tions in only from this site. The numer i cal ra tio is quite the same as north-west ern Af rica, rep re sent ing the spe cies de scribed in the previ ous case, yet the gen eral as pect of the Maharch here, have been reported, if rele vant, un der “Addi tional ma- fauna is very differ ent from that at Aferdou. At Maharch, terial” (Figs 3A, H, I–K, 5J–M, 7DD, EE, 19I–M, 25M–Q). the fauna is domi nated by Paulinaerhynchia paulinae and However, the mate rial from the northern MaÎder is quoted Desquamatia deserti, both of which do occur at Aferdou, only seldom, as it will be the object of a sepa rate study. In but are repre sented only by sin gle speci mens. This dissim i- ex cep tional cases, speci mens from outside NW Af rica have lar ity cor re sponds to dif fer ent bathymetric sit u a tions: mod - been figured or discussed (Figs 4T–V, 14A–E, P–T, erate water depths (but be low the photic zone) in the mid- 16V–Z, 19Y–DD, 33B); they are listed under the head ing ramp zone for Aferdou (and prob ably MadÀne) and a deeper “Other ma te rial ex am ined”. How ever, de scrip tions, unless outer ramp set ting for Maharch (Kaufmann, 1998). As in oth er wise stated, are based solely on spec imens from the the case of cnidari ans and sponges (Kaufmann, 1998), the three lo cal i ties in ves ti gated. brachio pod fauna at Aferdou is much richer (49 taxa, com - pared to 11 in Maharch). One should, however, bear in mind Spe cial tax o nomic ques tions that in the former section, brachio pods have been collected from a much longer stratigraphic inter val and from a larger Since a monograph of Middle Devo nian brachio pod out crop. fau nas from cen tral Eu rope, in clud ing the sys tem atic re vi - The col lec tion and prep a ra tion of brachi o pods from sion of taxa on the basis of type and topotypic collec tions, is Aferdou was rel atively easy, owing to their abundance in now in progress (Halamski, 2004a, 2009), several problems the scree. Only in some cases, a small amount of mechan ical are discussed only briefly in the pres ent account, with refer - prep a ra tion was re quired. The pres er va tion of this ma te rial ence to the above. Given that on the one hand it seemed ad- is quite variable: many spec imens are cracked and their ex - visable to the authors to pub lish the present de scrip tion as ternal surfaces are abraded, owing to aeolian corrasion; no- soon as pos sible and on the other there was a de lay in pub- netheless, a lot of very good speci mens have been collected lishing subse quent parts of the Holy Cross Mountains bra- as well, in some cases even with microornamentation (Figs chiopod mono graph (first part by Halamski, 2009), the re- 6B, 12L, 34L, 35R, S). Inter nal structures are rela tively well sult is that some tax o nomic de cisions out of ne ces sity were preserved. On the other hand, the limestone at Maharch is an tic i pated, in par tic u lar those con cern ing the representati- very hard, mak ing me chan i cal prep a ra tion dif fi cult. Shell ves of Kransia. 248 A. T. HALAMSKI & A. BALIÑSKI

When ever pos sible, synony mies are lim ited to the origi - two groups, differ ing in char ac ter and the den sity of orna menta - nal pub li ca tion and a sin gle (ex cep tion ally more) ref er ence tion. The first group is char ac ter ised by thick (i.e., thicker than fur- to papers, in which a given species was most thoroughly rows between them) and rela tively scarce radial ribs, whereas the discussed and where a full list of refer ences is to be found. sec ond one has fine (i.e., nar rower than fur rows between them) and nu merous ribs. If a nu meri cal char ac ter is needed, the quo tient This rule suffers three ex ceptions: pa pers dealing with North of the to tal rib numer and the max i mal shell di ame ter may be Af ri can mate rial always have been cited fully; taxa not dealt taken; its val ues are be tween 6.5 and 11.3 in the first group and be - with in the above-mentioned monograph (Halamski, 2009) tween 16.6 and 26.3 in the sec ond one. The two groups are de - have been given complete synony mies; and addi tions to the scribed be low as sep a rate taxa, but their vari abil ity should be already published first part of the monograph also have checked on a larger sample. been taken into ac count. Forma lised signs in synonymic lists follow Halamski (2009; further refer ences therein). Deliella deliae Halamski, 2004 [2004b] In the authors’ opinion, the value of the pres ent de scrip - Fig. 3Q2, R2, S2 tion of a Rhenish-type fauna from Africa is largely in an ex- ten sive pho to graphic doc u men ta tion of the ma te rial, tes ti fy- v* 2004b Deliella deliae n. gen. et sp. – Halamski: pp. 184– ing that the use of Euro pean taxa names is not the fancy of a 188, figs 1A–F, 2C–D, 3B–D. lumping-style taxon omy, but the only solu tion that could be v. 2009 Deliella deliae Halamski, 2004 – Halamski, p. 49, pl. 1, figs 9, 13 [ubi syn.]. taken after a metic u lous study of the brachio pods. In some cases, this has re quired pho to graph ing ma te rial of oth er wise Ma te rial: MB.B.9384b at tached to the ven tral valve of Spino- poorly il lus trated taxa orig i nat ing from Eu rope. Im ages were cyrtia cf. elburzensis (MB.B.9384c), from MadÀne el Mrakib (coll. V. Ebbighausen). taken mostly using a Fujifilm Finepix S2 pro cam era with a Additional ma te rial: MNHN.F.A48107c at tached to the oper- Nikon micro 60 mm lens; in some cases a Canon 350D cam- culum of Calceola sp. (MNHN.F.A48107a), lo cal ity 348A sensu era with a Canon EF 100 mm macro lens was used. Enlarge - Sougy (1964; Zemmour Noir, Mau ri ta nia). ments were photo graphed under a Nikon SMZ 1500 binoc u - De scrip tion: Dorsal valve coni cal, el liptic in outline, 4.1 mm lar micro scope, equipped with a Nikon D800 dig ital cam era. long. Apex excentric, sit uated 1.4 mm from the poste rior mar gin Spec imens were coated with am mo nium chloride before be- (34% of the max i mal di am e ter). Ra dial or na men ta tion of ca. 108 ing photo graphed. Some photo graphs were as sem bled from a straight, fine ribs (to tal numer of ribs es ti mated on the ba sis of series of up to five frames, using the Helicon Focus pro- count on the pre served fourth of the shell). gramme. In ter nal struc tures were ex am ined, if needed, Re marks: The finely ribbed rep re sen ta tive of Deliella from north- ern Af rica shows no sig nif i ca tive dif fer ences from D. deliae from through the standard method of serial sections and ace tate the Holy Cross Mountains. peels; drawings were prepared with a camera lucida. Dis tri bu tion: Holy Cross Mountains, MaÎder, and Mau ri ta nia; up - per Eifelian to lower Givetian. De scrip tions Phylum Brachiopoda Duméril, 1805 Deliella aff. rhenana (Dahmer, 1930) Subphylum Craniiformea Popov et al., 1993 Fig. 3O, P, Q1, R1, S1 Class Craniata Williams et al., 1996 aff. 1930 Philhedra rhenana n. sp. – Dahmer: pp. 92–93, pl. 6, Order Craniida Waagen, 1885 fig. 2. Family Craniidae Menke, 1828 v 2009 Deliella sp. – Halamski, p. 49, pl. 9, fig. 8. aff. 2012 Deliella rhenana (Dahmer, 1930) – Franke, p. 135, pl. Ge nus Deliella Halamski, 2004 [2004b] 1, figs 1–3 [ubi syn.]. Type spe cies: Deliella deliae Halamski, 2004 [2004b]; Ma te rial: Two in di vid uals, MB.B.9382a at tached to the dor sal Ska³y, northern region of the Holy Cross Mountains; lower valve of Xystostrophia umbraculum (MB.B.9832b) and MB.B. part of the Ska³y beds, upper Eifelian 9384a at tached to the ven tral valve of Spinocyrtia cf. elburzensis Re marks: The mate rial of Deliella from northern Africa is scarce (MB.B.9384c), both from MadÀne el Mrakib (coll. V. Ebbig- (five in di vid uals), none the less it may be quite easily di vided into hausen).

Fig. 3. Middle De vo nian Craniida and Strophomenida from north west ern Af rica. A–L. Leptagonia analogaeformis Biernat, 1966. A. Artic ulated shell MNHN.F.A48100 in ventral view. MaÎder, local ity TM 810/1, coll. H. Hollard. B–D. Artic ulated shell ZPAL Bp 68/1/8/1 from Aferdou in ven tral, dor sal, and lat eral views. E. Dor sal valve ZPAL Bp 68/1/8/3 from Aferdou. F. Spec i men ZPAL Bp 68/1/8/2 from Aferdou in dor sal view. G. Spec i men ZPAL Bp 68/1/8/4 from Aferdou in ven tral view. H. Spec i men MNHN.F.A48106 from local ity 346N sensu Sougy (1964) (Zemmour Noir, Mau ri ta nia). I–K. Dor sal valve SMF 94838 from Jebel Abiir, close-up of cardinalia, in ter nal and ex ter nal views. L. Spec i men ZPAL Bp 68/1/8/5 from Aferdou in ven tral view. M–N, U. Parastrophonella anaglypha (Kayser, 1871). Ven tral valve SMF 94853 from MadÀne el Mrakib. Ex ter nal (N) and in ter nal (M) views, and mus cle area (U). O–S. Deliella aff. rhenana (Dahmer, 1930). All speci mens from MadÀne el Mrakib. O, P. Shell MB.B.9382a attached onto the dor sal valve of Xystostrophia umbraculum (von Schlotheim, 1820) MB.B.9832b in ven tral (O) and lat eral (P) views. Q, S. Shell MNHN. F.A48107b attached onto an operculum of Calceola sp. MNHN.F.A48107a in general (Q1) and en larged (S1) views. R. Shell MB.B. 9384a (R1) at tached on the ven tral valve of Spinocyrtia cf. elburzensis MB.B.9384c (see also Fig. 34E). Q–S. Deliella deliae Halamski, 2004. All spec i mens from MadÀne el Mrakib. Q, S. Shell MNHN.F.A48107c at tached onto an operculum of Calceola sp. MNHN.F. A48107a in gen eral (Q2) and enlarged (S2) views. R. Shell MB.B.9384b (R2) attached on the ven tral valve of Spinocyrtia cf. elburzensis MB.B.9384c (see also Pl. 34E). T. Protodouvillininae gen. et sp. indet. Frag mentary ven tral valve ZPAL Bp 68/1/12/1 from Aferdou MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 249 250 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 251

Additional ma te rial: One in di vid ual MNHN.F.A48107b at tached (1964, pl. 44, p. 11); ar tic ulated shell MNHN.F.A48106 from the to the operculum of Calceola sp. (MNHN.F.A48107a), lo cal ity lo cal ity 346N; two ven tral valves MNHN.F.A48105 from the lo- 348A sensu Sougy (1964; Zemmour Noir, Mau ri ta nia). cal ity 2428; all the above from the Zemmour Noir, Mau ri ta nia, Other ma te rial ex am ined: SMF 66200 attached to Protodou- coll. J. Sougy; iso lated dor sal valve SMF 94838, Jebel Abiir, villina interstrialis, Eifel, Middle Devo nian, illus trated by Ha- north ern MaÎder, coll. U. Jansen; artic u lated shell MNHN.F.A lamski (2009, p. 1, fig. 8). 48100 from the lo cal ity TM 810/1, MaÎder?, coll. H. Hollard. De scrip tion: Dorsal valve coni cal, el lip tic in out line, up to 7.1 mm De scrip tion: Shell up to ca. 30 mm in width, trans verse (width-to- long and 6.1 mm wide. Apex excentric, its distance from the pos te - length ra tio ca. 1.5), subrectangular to subtrapezoidal in out line, rior margin be ing 29–35% (N = 3) of the max i mal di ame ter. Ra dial geniculate. Hinge line and an te rior commissure straight, par al lel. or na men ta tion of thick, straight costae, bi fur cat ing rather rarely, Ventral valve with weakly con vex (sometimes flattened later ally) 35–55 in number. disk and dorsally di rected trail; interarea rect angu lar, apsacline. Re marks: As observed al ready by Halamski (2009), the Eifel rep - Dorsal valve concave (flat disk and geniculation); interarea low, re sen ta tive of Deliella has coarser or na menta tion than those from hypercline. Pseudodeltidium and chilidium not pre served. Or na - the Holy Cross Mountains. The anal ysis of the ma te rial from Af - menta tion of con cen tric rugae, (6–)8–10 per cm, and ra dial costae rica con firms this sepa ra tion. The coarsely or na mented Mid dle and costellae, ca. 7–8 per 5 mm at ante rior margin. Devo nian species is most simi lar to late Emsian Deliella rhenana Inte rior (de scrip tion based on the spec i men SMF 94838 from from which it dif fers in its smaller size (the diam e ter of the Jebel Abiir). Dor sal valve: car di nal pro cess con sisting of two mas- holotype of D. rhenana is 18 mm; Franke, 2012). It may rep re sent sive lobes; muscle area rhomboidal, bi sected by a me dian sep tum a new spe cies, but the ma te rial is in suffi cient to pro vide a ba sis for that con tin ues an te ri orly for a short length; vas cu lar im prints in an - a for mal de scrip tion. te rior re gion of the valve. Ven tral valve un known. Dis tri bu tion: Eifel, MaÎder, and Mau ri ta nia, Mid dle De vo nian. Re marks: The mate rial described here agrees in both exter nal and (dor sal) in ter nal char ac ters with Leptagonia analogaeformis (Bie- rnat, 1966) from the Mid dle De vonian of the Holy Cross Moun- Subphylum Rhynchonelliformea Williams et al., 1996 tains and the Eifel (Halamski, 2009). The den sity of con cen tric Class Strophomenata Williams et al., 1996 rugae is ap par ently a vari able char ac ter with in di vid u als from the Order Strophomenida Öpik, 1934 Holy Cross Mountains and from Mau ri ta nia, hav ing sometimes as Family Rafinesquinidae Schuchert, 1893 few as 4 rugae per cm; none the less, this is not a suf fi cient rea son to subdi vide this long-lasting member of a bradytelic lin eage. Ge nus Leptagonia M’Coy, 1844 Dis tri bu tion: Middle De vo nian of the Holy Cross Mountains, Type spe cies: Producta analoga Phillips, 1836; proba bly Eifel, Sauerland, MaÎder, and Mau ri ta nia; pos si bly also In ner Bolland, Yorkshire; lower Visean Mongolia and Ja pan (Chen and Tazawa, 2003). Leptagonia analogaeformis (Biernat, 1966) Fig. 3A–L Family Douvillinidae Caster, 1939 1934 Leptaena rhomboidalis (Wilckens) – Torley, p. 128, Ge nus Protodouvillina Harper and Boucot, 1978 pl. 9, fig. 82. v 1964 Leptaena rhomboidalis (Wilckens) – Sougy, pl. 44, Type spe cies: Strophomena inaequistriata Conrad, 1842; figs 10–11. New York; Ham il ton (Mid dle De vo nian) v* 1966 Leptagonia analogaeformis n. sp. – Biernat, pp. 38– Protodouvillina interstrialis (Phillips, 1841) 42, text-figs 6, 7, pl. 3, figs 1– 16, pl. 4, figs 1–13. Fig. 4A–F, L v. 2009 Leptagonia analogaeformis (Biernat 1966) – Ha- lamski, pp. 50–52, text-figs 2, 3, pl. 1, figs 16–19, * 1841 Orthis interstrialis, Leptaena interstrialis – Phillips: 21–23, 25, pl. 2, figs 1–12, 14, 15, 17, pl. 16, figs pp. 61–62, pl. 25, fig. 103 a–e. 19–22, 29 [ubi syn.]. v. 1885 Strophomena interstrialis Phillips – Mau rer, pp. 144– Ma te rial: Five subcomplete to frag mentary spec i mens from Afer- 145, pl. 5, fig. 17. dou, ZPAL Bp 68/1/8/1; nine ar tic ulated shells SMF 85681 and v. 1964 Douvillina sp. e. g. interstrialis (Phillips, 1841) – MB (unnum bered; V. Ebbighausen’s col lec tion) from MadÀne el Sougy, p. 451. Mrakib. v. 2009 Protodouvillina interstrialis (Phillips 1841) – Ha- Ad di tional ma te rial: Artic u lated shell MNHN.F.R10276 from lamski, pp. 53–54, text-fig. 4, pl. 3, figs 1–6, 8–17, Aguelt Tighirt fig ured by Sougy (1964, pl. 44, p. 10); ar tic u lated 19–20, 30–32 [ubi syn.]. shell MNHN.F.R10277 from Amgala marls figured by Sougy Ma te rial: A single subcomplete, but decorticated ar tic ulated shell

Fig. 4. Middle De vonian Strophomenida and Productida from north western Af rica and Europe. A–F, L. Protodouvillina interstrialis (Phillips, 1841). A–E. Artic u lated shell ZPAL Bp 68/1/52/1 from Aferdou in an te rior, pos te rior, ven tral, dor sal, and lat eral views. F, L. In - te rior of the ven tral valve MB.B.9383 from MadÀne el Mrakib in gen eral (L) and en larged (F) views. G–K, M. Devonaria sp. All spec i - mens from Aferdou. G, H. Ven tral valve 68/1/11/3 embed ded in lime stone in ven tral and pos te rior views. I–K. Subcomplete ar tic ulated shell 68/1/11/1 in lat eral, ventral, and dorsal views. M. In complete artic ulated shell 68/1/11/2 in dor sal view. N–V. Radiomena irregularis (Roemer, 1844). N. In complete ven tral valve 68/1/7/3 from Aferdou embed ded in lime stone. O. In complete ar tic ulated shell ZPAL Bp 68/1/6/1 from Aferdou in ven tral view. P. In complete ar tic u lated shell 68/1/7/4 from Aferdou in pos te rior views. Q. In complete ar tic u lated shell 68/1/7/2 from Aferdou in ventral view. R. Ar tic ulated shell 68/1/7/1 from Aferdou over grown by bryozoans. S. In complete ventral valve 68/1/7/5 from Aferdou embed ded in lime stone. T, V. In ter nal (T) and ex ter nal (V) mould of a ven tral valve (part and coun ter part); holotype HLMD-Mr 8845, 8847 of Strophomena porrigata Mau rer, 1885 from Grube Hainau at Waldgirmes. U. In complete ven tral valve HLMD-Mr 8842 from Grube Hainau at Waldgirmes embed ded in lime stone 252 A. T. HALAMSKI & A. BALIÑSKI

from Aferdou, ZPAL Bp 68/1/52/1 and a sin gle ven tral valve MB. mus cle field; ra dial man tle ca nals im pres sions pre served an te ri- B.9383 from MadÀne el Mrakib (V. Ebbighausen’s col lec tion). orly. Dor sal in te rior: traces of a very poorly pre served muscle field Ad di tional ma te rial: Two ar tic u lated shells MNHN.F.A48117, and mantle canals. 48118 from the lo cal ity 402M, 1.5 km ENE from Agouelt Oudiate Re marks: Mau rer (1885) and Halamski (2009) were misled in in - el Khyan, Zemmour Noir, Mau ri ta nia; “ba sis of the Calceola ter pret ing flat-valved and more weakly or na mented in di vid u als of marls, lower Givetian”; coll. J. Sougy. R. irregularis as sep a rate species, Strophomena porrigata and Other ma te rial ex am ined: Frag men tary spec i men HLMD-Mr Zophostrophia? aaatos, respec tively. They are better consid ered 11605, 11606 from Grube Rothe Erde at Waldgirmes (Lahn Syn- as end members of vari a tion se ries of the for mer. cline) fig ured by Mau rer (1885, pl. 5, p. 17). Dis tri bu tion: R. irregularis is known from the Eifelian to Give- De scrip tion: The sin gle avail able shell is concavo-con vex, ca. 27 tian of the Eifel, the Ardennes (Godefroid, 1975, p. 76), the Lahn mm wide, ca. 22.5 mm long, and ca. 12 mm thick. Hinge line Syncline, the Holy Cross Mountains, Devonshire, Moravia, and straight. Interareas low, lon gi tu di nally stri ate. An te rior commis- Burma (Harper et al., 1967; Halamski, 2009). sure rectimarginate. Or na men ta tion parvicostellate, poorly pre served; 1(–2) cost- Ge nus Parastrophonella Bublièenko, 1956 (ell)ae per mm at ante rior margin and 3–4 striae between each pair Type spe cies: Strophomena anaglypha Kayser, 1871; of cost(ell)ae. Shell substance punctate. Ven tral in te rior: muscle area as long as one fourth of the valve Eifel, Prüm Syncline; Crinoiden-Schicht, Middle Devo nian length, bilobed, bor dered lat er ally by low rounded ridges, pos te ri - (mid dle Eifelian?) orly bi sected by a short me dian sep tum; valve floor pustulose. Parastrophonella anaglypha (Kayser, 1871) Dor sal in te rior un known. Fig. 3M, N, U Dis tri bu tion: This species was reported from several Early to Late De vo nian lo cal i ties in Eu rope and Asia (Brice, 1971) and in - v* 1871 Strophomena anaglypha – Kayser: pp. 628–629, pl. deed it seems a cosmo pol i tan taxon, even if part of these data re - 14, fig. 3 a–f. quires con fir ma tion (Halamski, 2009). v. 1885 Strophomena anaglypha Kayser – Mau rer, p. 148, pl. 5, fig. 26. v. 1961b ? Parastrophonella anaglypha (Kayser 1871) – Drot, Ge nus Radiomena Havlíèek, 1962 pp. 257–258, pl. 3, figs 1, 2–4. Type spe cies: Orthis irregularis Roemer, 1844, v. 2009 Parastrophonella anaglypha (Kayser 1871) – Ha- Gerolstein, Eifel; Mid dle De vonian lamski, pp. 60–62, text-fig. 7, pl. 5, figs 1–5, 9–13, Radiomena irregularis (Roemer, 1844) 16–20, 23–27, 30–47 [ubi syn.]. Fig. 4N–V Ma te rial: Two ven tral valves SMF 94853 and MB unnum bered (V. Ebbighausen’s col lec tion) from the Drotops beds at MadÀne el v* 1844 Orthis irregularis mihi – Roemer: p. 75, pl. 4, fig. 1 Mrakib. a–c. Other ma te rial ex am ined: HLMD-Mr 8837 from Grube Hainau at v. 1885 Strophomena irregularis Roemer – Mau rer, pp. 145– Waldgirmes (Lahn Syncline), fig ured by Mau rer (1885, pl. 5: 26). 146, pl. 5, figs 18–20. De scrip tion: The single available ventral valve is ca. 13 mm wide v. 1885 Strophomena porrigata n. s. – Mau rer, pp. 148–149, and 8.9 mm long, mucronate, and ven trally geniculate with a trail pl. 6, fig. 1. ca. 3 mm high; interarea anacline, tri an gular, low, pseudodelti- v. 2009 Radiomena irregularis (Roemer 1844) – Halamski, p. dium preserved only in the apical part; hinge line denticulate. 56, pl. 7, figs 12, 16, 19, 22, 23, pl. 16, figs 2, 25–28 Traces of poorly pre served, multicostellate or na menta tion. Diduc- [ubi syn.]. tor scars deep, ex tend ing to about 1/2 of the valve length, slightly v. 2009 Zophostrophia? aaatos sp. n. – Halamski, pp. 59–60, flabellate, their ex ter nal bor ders forming an an gle of ca. 30°. Ad- pl. 2, figs 13, 23, pl. 4, figs 1–10, pl. 16, fig. 1 [ubi syn.]. ductor scars nar row, elon gate, sep a rated by a low, rounded me dian Ma te rial: One subcomplete, three in complete spec i mens, one ex fo- ridge. Valve floor strongly pustulose out side the mus cle scars, li ated and show ing traces of dorsal in ter nal struc tures, one poorly pustules hav ing even in some cases the ap pear ance of high rows or pre served ven tral in te rior, four frag ments, all from Aferdou, ZPAL rounded spines. Bp 68/1/6/1, 68/1/7/1–7. Dis tri bu tion: This Eifelian to Givetian species is known from the Other ma te rial ex am ined: Ven tral valve em bed ded in lime stone Eifel, the Lahn Syncline, the Ardennes, the Holy Cross Mountains, HLMD-Mr 8842, Grube Hainau at Waldgirmes (Lahn Syncline), the Pyre nees, the Montagne Noire, Mo rocco, and Mauri ta nia (Har- fig ured by Mau rer (1885, pl. 5: 20); holotype of Strophomena per and Boucot, 1978; Halamski, 2009, and ref er ences therein). porrigata HLMD-Mr 8845, 8847 from Grube Hainau at Waldgir- mes (inter nal mould and exter nal cast of the same speci men, see Protodouvillininae gen. et sp. indet. Fig. 4T, V; HLMD-Mr 8846 is the orig i nal gutta-percha cast by F. Fig. 3T Maurer). De scrip tion: Shell subrectangular to subtrapezoidal in out line, ge- Ma te rial: A sin gle frag men tary spec i men em bed ded in lime stone niculate, up to 71 mm wide, width-to-length ra tio 1.65–2.09, dor - (frag ment of the ventral valve vis i ble) from Aferdou, ZPAL Bp sally geniculate. Pos te rior mar gin straight. Ven tral valve con vex in 68/1/12/1. umbonal re gion, flat in lat eral flanks, ir reg u larly un du lat ing; some- De scrip tion: The frag mentary ven tral valve is >14.4 mm wide times, however, nearly flat. On the trail, traces of poorly pre served and >7 mm long, weakly con vex, and subtriangular in an te rior parvicostellate or na menta tion with 3–4 costellae per mm. Ven tral view. The umbo is rather fine, weakly incurved, and the pos te rior interarea anacline, higher than the apsacline dorsal one. Orna men ta - margin is straight. The or na menta tion con sists of radial and sub- tion of faint rugae, 3–4 per 5 mm, and slightly wavy ra dial costae concen tric ele ments: costae, costellae, and striae form a parvicos- and costellae, 4–6 (–7) per mm at disk margin. Ven tral in te rior: tellate pattern with ca. 2 cost(ell)ae per mm at 5 mm from the mus cle field bor dered by fine ridges di ver gent at 70°; a fine and umbo and 5–7 striae between each pair of cost(ell)ae, whereas nu- low median sep tum on low elon gated cal lus in the pos te rior third merous transverse wrin kles are aligned along presumed growth of the valve; teeth not pre served; floor pustulose lat er ally from the lines, 3–4 lines of them per mm. MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 253

Re marks: This frag mentary spec i men is included within the sub- west from a small escarp ment 2 km NE from Aguelt Oudiate el family Protodouvillininae Caster, 1939 on ac count of char ac ter is- Khyam; all from Zemmour Noir, Mauri ta nia. tic or na men ta tion. How ever, choos ing be tween sev eral gen era in De scrip tion: Shell concavo-con vex, wider than long, auriculate, which it is pres ent, such as, e.g. Cymostrophia Caster, 1939 or up to 27.2 mm in width. Ven tral valve strongly con vex, umbo very Moravostrophia Havlíèek, 1962, is im possi ble. thick, beak strongly incurved. Interareas lin ear. Radial or na ment lack ing. Ir reg ular rugae present on both valves. Ven tral spines ar - Order Productida Sarytcheva and Sokolskaya, 1959 ranged in a row along the hinge line, 2 per mm (spine bases poorly Suborder Chonetidina Muir-Wood, 1959 pre served in two spec i mens), oth er wise ir reg u larly dis trib uted on Family Anopliidae Muir-Wood, 1962 the ven tral valve; no dor sal spines. Dor sal in te rior (Fig. 5C, M, N): car di nal pro cess bifid, the dor sal sur face of the lobes with two lon - Ge nus Devonaria Biernat, 1966 gi tu di nal ridges, sep a rated by a me dian fur row; den tal sock ets Type spe cies: Chonetes zeuschneri Sobolew, 1909; shal low, wide; me dian sep tum high, but short. Ven tral in te rior un - Ska³y, Holy Cross Mountains, Ska³y beds, upper Eifelian known. Re marks: The de scribed sam ple is in cluded in Poloniproductus Devonaria sp. on ac count of ir reg u lar rugae on the dor sal valve and ab sence of Fig. 4G–K, M ribs. In Productella Hall, 1867 the ven tral valve is smoother. Spinulicosta Nalivkin, 1937 and Helaspis Imbrie, 1959 have an te - Ma te rial: Four in complete spec i mens from Aferdou, ZPAL Bp rior rib bing. Devonoproductus Stainbrook, 1943 has reg u lar rugae 68/1/11/1–4. and ven tral rib bing. The max i mal size of the MaÎder spec i mens is De scrip tion: Shell up to ca. 12 mm in length, trans verse. Pos te rior slightly greater than that of the Holy Cross ones (27 mm vs. 22 margin straight. Ven tral valve moder ately to strongly con vex with mm; data for the latter af ter Biernat, 1966), but in view of sig nif i - orthocline interarea; dorsal valve concave, interarea not preserved. cant variabil ity of this species expressed even in its name it is An te rior commissure rectimarginate. Or na men ta tion of straight judged in sig nif i cant. costae and costellae, 3(–4) per mm, the latter appear ing by bifur ca - The ex am ined ma te rial of Productella subaculeata sensu Sougy tion. (1964, p. 446, pl. 44: 9) is conspecific with Poloniproductus Re marks: The available mate rial shows some varia tion in valve varians, as above. However, other Mid dle De vo nian productides con vex ity and or na men ta tion den sity; this is in ter preted as intra- may also be pres ent in the Zemmour (Productella navicella sensu spe cific vari a tion. Devonaria zeuschneri (Sobolew, 1909) from Sougy, 1964; MNHN.F.A48109, 48111, 48112 from Tighirt, lo - the late Eifelian to early Givetian of the Holy Cross Mountains and cal ity 526; Sougy, 1964, p. 388). Productella subaculeata re - D. minuta (von Buch, 1837) from the Eifelian of the Eifel [re - ported by Brousmiche (1975) from the Famennian of the Drâa ported also from the Zemmour Noir by Sougy, 1964; “Devonaria valley is unre lated to the samples described here. sp., minuta? (Buch, 1837)” from Smara by Schemm-Gregory and Dis tri bu tion: Holy Cross Mountains (Eifelian and Givetian; Ha- Jansen (2005)] have coarser or na menta tion with, re spec tively 2 lamski, 2009 and ref er ences therein) and MaÎder. and 1.6 ± 0.2 cost(ell)ae per mm (and not 5–6 as erro ne ously stated for the for mer by Halamski, 2009, p. 62; data for D. minuta after Struve, 1981b). D. minutissima Struve, 1981 [1981b] has a simi - Order Orthotetida Waagen, 1884 larly fine or na menta tion (3.8 ± 0.45 costellae per mm) but is sig- Family Chilidiopsidae Boucot, 1959 nifi cantly smaller, its max i mal width being about 6 mm (Struve, 1981b). Ge nus Xystostrophia Havlíèek, 1965 Type spe cies: Terebratulites umbraculum von Schlotheim, Suborder Productidina Waagen, 1883 1820; Gerolstein (Eifel); Middle Devo nian (most proba bly Family Monticuliferidae Muir-Wood and Cooper, 1960 Eifelian) Ge nus Poloniproductus Biernat and Lazarev, 1988 Xystrostrophia umbraculum (von Schlotheim, 1820) Type spe cies: Productella varians Biernat, 1966; Ska³y, Fig. 6B, E–K Holy Cross Mountains; Ska³y beds, upper Eifelian * 1820 Terebratulites umbraculum – von Schlotheim: p. 256. Poloniproductus varians (Biernat, 1966) ? 1940 Streptorhynchus umbraculum Schloth. – Cottreau, p. Fig. 5A–U 196. non 1950 Schellwienella umbracula Schlotheim – Termier and vp 1964 Productella subaculeata (Murchi son, 1840) – Sougy, Termier, pl. 88, figs 24, 25. p. 446, pl. 44, fig. 9. ? 1950 Indeterminé – Termier and Termier, pl. 89, figs 17– v* 1966 Productella varians n. sp. – Biernat: pp. 66–73, text- 21. figs 19–21, pl. 11, figs 1–23, pl. 12, figs 1–16, pl. 13, ? 1950 Orthotetes Chouberti nov. sp. – Termier and Termier, figs 1–10. pl. 91, figs 13–18. v 1988 Poloniproductus varians (Biernat, 1966) – Biernat v. 1952b Schellwienella umbracula (Schlotheim) – Le Maître, and Lazarev, pp. 67–68, text-fig. 2, pl. 17, figs 1–5, pl. pp. 326–327, pl. 2, figs 21–22. 18, figs 1–4, pl. 19, figs 1–3, pl. 20, figs 1–2. 1960 Schellwienella umbracula Schlotheim – Gevin, p. Ma te rial: Four subcomplete ar tic u lated shells, one frag mentary 162. spec i men, two ven tral valves embed ded in lime stone, all from v. 1964 Schuchertella sp. e. g. umbracula (Schlotheim) – Aferdou, ZPAL 68/1/10/1–7; six ar tic u lated shells (MB unnum - Sougy, pl. 49, fig. 13. bered; V. Ebbighausen’s col lec tion) and a sin gle poorly pre served 2005 Xystostrophia cf. umbraculum (von Schlotheim, dor sal in te rior SMF 94850 from MadÀne el Mrakib. 1820) – Schemm-Gregory and Jansen, p. 27. Ad di tional ma te rial: Sixty spec i mens, MNHN.F.A48110, 48113, v. 2009 Xystrostrophia umbraculum (von Schlotheim 1820) – 50411, 50412, mostly frag mentary but in clud ing some subcom- Halamski, pp. 69–71, pl. 5, figs 6–8, 14, 15, 21, 22, 28, plete ar tic u lated shells and sin gle valves from local ity 2118 sensu 29, pl. 9, figs 1–31, pl. 16, figs 3, 4, 8, 9, 13, 17, 18 [ubi Sougy (1964); two frag mentary spec i mens, MNHN.F.A48114, syn.]. 254 A. T. HALAMSKI & A. BALIÑSKI

Fig. 5. Poloniproductus varians (Biernat, 1966). Aferdou. A, B, D, E. Ar tic u lated shell ZPAL Bp 68/1/10/1 in dor sal (A, D) and ven - tral (B, E) views. C, N. In complete dor sal in te rior SMF 94850 from MadÀne el Mrakib. F–I. Ar tic ulated shell ZPAL Bp 68/1/10/2 in dor - sal, ventral, lat eral, and poste rior views. J–L. Ar tic u lated shell MNHN.F.A50411 from the local ity S2118B sensu Sougy (1964) (Zemmour Noir, Mauri ta nia) in dor sal, ven tral, and lat eral views. M. Dor sal in te rior MNHN.F.A50412 from local ity S2118B sensu Sougy (1964) (Zemmour Noir, Mau ri ta nia). O–R. Ar tic ulated shell ZPAL Bp 68/1/10/3 in dor sal, ven tral, lat eral, and poste rior views. S–T. Ar- tic u lated shell ZPAL Bp 68/1/10/2 in dor sal and ven tral views. U. Ven tral in te rior ZPAL Bp 68/1/10/7

Ma te rial: Three subcomplete to incom plete artic u lated shells from the lo cal i ties 2103T, 346I, 401A, 508G, Zemmour Noir, from Aferdou, ZPAL Bp 68/1/41/1-3; one ar tic u lated shell MB.B. Mauri ta nia, coll. J. Sougy; two ar tic ulated shells MNHN.F.A 9382b, two iso lated dor sal valves SMF 94851 and MB un num- 48121 from the lo cal ity Tf 82, Mo rocco, coll. H. Hollard; ar tic u - bered (V. Ebbighausen’s col lec tion), and a sin gle ven tral valve lated shell GFCL 2623 col lected south from Hassi el Motleh SMF 94852 from the Drotops beds at MadÀne el Mrakib. (Adrar, Mau ri ta nia) fig ured by Le Maître (1952b, pl. 2: 21–22); ar - Ad di tional ma te rial: Artic u lated shell MNHN.F.R10286 from tic u lated shell MB.B.1933 from Jebel Issomour, north ern MaÎder, the “Givetian” of Amgli Zguilma (Mau ri ta nia) fig ured by Sougy coll. V. Ebbighausen, fig ured by Halamski (2009, pl. 16: 3, 4, 8, 9, (1964); ar tic u lated shells MNHN.F.A48119, 48122, 48123, 48124 13, 17, 18). MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 255

Fig. 6. Middle De vonian Orthotetida from south ern MaÎder. A, C–D. Iridistrophia cf. undifera (Schnur, 1854 in 1853–54) from Aferdou. A. Frag mentary shell showing traces of dorsal in te rior. C, D. In complete ar tic ulated shell in ven tral and ante rior views. B, E–K. Xystostrophia umbraculum (von Schlotheim, 1820). B. Microornamentation of the ar tic u lated shell ZPAL Bp 68/1/41/2 from Aferdou. E–I. Ar tic ulated shell ZPAL Bp 68/1/41/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and ante rior views. J. Ven tral in te rior SMF 94852 from MadÀne el Mrakib. K. Dorsal in te rior SMF 94851 from MadÀne el Mrakib 256 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 257

De scrip tion: Shell up to 63.3 mm in width, moder ately to mark - line, trans verse (width-to-length ra tio 1.38–1.63), dorsibiconvex. edly trans verse (width-to-length ra tio 1.21 to 1.50), slightly resu- Poste rior mar gin straight. Dorsal valve strongly con vex. Ven tral pinate. Ven tral interarea apsacline, high, pseudodeltidium well de - valve weakly con vex, interarea high. Orna menta tion of smooth vel oped; dor sal interarea anacline, lin ear. Or na men ta tion (Fig. 6B) costae and costellae, 1–2 per mm at an te rior mar gin; in mar ginal of crenulated costae and costellae, 1–2 per mm. re gion fila, not con tin uous through the costellae. Dor sal in te rior: a Ventral inte rior: teeth stout; dental plates prolong ing as ridges short median sep tum oc cu pies up to 1/4 of the valve length. Ven - border ing muscle scars; diductor scars large, radi ally stri ate, bi- tral inte rior: traces of adduc tors wide, distally flabellate; their ex- sected by a low median septal ridge; adductor scar situ ated medi - te rior bor ders di verg ing at ca. 80o. ally, rounded; valve floor el e vated pos te ri orly to the adductor scar. Re marks: This char ac ter is tic brachi o pod is very sim i lar (and, in Dorsal inte rior: cardi nal process bifid with two ventrally di rected con se quence, con sid ered conspecific) to Iridistrophia cf. undifera lobes, pos te ri orly with four ridges vis i ble in the pedicle open ing; from the Mid dle De vo nian of the Eifel and of the Holy Cross Mou- muscle scars strongly sculp tured, sub di vided by a low me dian ridge. ntains (Halamski, 2009). The type ma te rial of I. undifera from the Re marks: It may be use ful to note that the pre sumed Givetian age Eifel is lost, where fore the use of open no mencla ture is jus ti fied. of a spec imen of X. umbraculum from Jebel Issomour in the north - Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder; Middle De- ern MaÎder (Halamski, 2009, pl. 16) was based on a pre limi nary vo nian. in ves ti ga tion of that sec tion; more over, the spec i men it self was col lected from rubble (†V. Ebbighausen, pers. comm., 2008). In any case, any speci mens of this species, for which an ac curate age Class Rhynchonellata Williams et al., 1996 deter mi na tion can be given, are from the Eifelian (see Halamski, Order Orthida Schuchert and Cooper, 1932 2009 for de tails). The Givetian age given for the Zemmour mate - Family Rhipidomellidae Schuchert, 1913 rial by Sougy (1964) might also per haps re sult from a misiden ti fi - ca tion of an Anarcestes as a Werneroceras (Dumestre and Illing, Ge nus Aulacella Schuchert and Cooper, 1931 1967), giv ing a too young age for an im por tant cor re la tion ho ri zon Type spe cies: “Orthis eifeliensis Schnur, 1853” (misspell - (see also Basse, 2012, p. 30). Con trary to the opin ion of Rache- ing for eifliensis; = Orthis prisca Schnur, 1851); Eifel, boeuf et al. (2001, p. 146), Schellwienella umbracula sensu Le Mid dle De vo nian Maître (1952b) from the Mau ri ta nian Adrar rep re sents X. um bra- culum and not a rep re sen ta tive of Iridistrophia. Aulacella prisca (Schnur, 1851) Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, Zemmour Fig. 7A–M Noir, Adrar, Turkestan, Burma (Halamski, 2009, sup ple mented); v* 1851 O[rthis] prisca nob. – Schnur: p. 13. Mid dle De vo nian. 1950 Aulacella interlineata Sowerby – Termier and Ter- mier, pl. 81, figs 19–26. Ge nus Iridistrophia Havlíèek, 1967 v. 1959 Aulacella eifeliensis (de Verneuil) – Biernat, pp. 26– 35, text-figs 9, 10, text-pl. 3, pl. 1, figs 10–15, pl. 2, Type spe cies: Orthis umbella Barrande, 1848, Barrandien, figs 1–15, pl. 3, figs 9, 10, pl. 12, figs 1, 2. Bo he mia; up per Lochkovian v. 1964 Aulacella interlineata – Sougy, p. 437, pl. 42, figs Iridistrophia cf. undifera (Schnur, 1854 in 1853–54) 18–19. Fig. 6A, C–D v. 2009 Aulacella prisca (Schnur, 1851) – Halamski, pp. 74– 77, pl. 11, figs 1–36, pl. 14, figs 3–10, 13–16, 21–27, cf. 1854 Orth[is] undifera n. sp. – Schnur: p. 217, pl. 45, fig. 29, 30, text-fig. 11 [ubi syn.]. 1a–d. Ma te rial: Thirty-one for the most part com plete spec i mens from v. 2009 Iridistrophia cf. undifera (Schnur, 1854 in 1853–54) Aferdou, ZPAL Bp 68/1/13/1–31; one poorly pre served from – Halamski, p. 71–72, pl. 8, figs 1–11, pl. 16, figs 5–7, Maharch, ZPAL Bp 68/2/13/1; a sin gle ju ve nile from MadÀne el 11, 12, 16 [ubi syn.]. Mrakib, MB un numbered (V. Ebbighausen’s col lec tion). Ma te rial: Four incom plete artic u lated shells, three fragments, Ad di tional ma te rial: Artic u lated shell MNHN.F.R10268 and in - three poorly pre served in te ri ors, one ven tral and two dorsal, all ter nal mould MNHN.F.R10269, both from the “Givetian” of Bir from Aferdou, ZPAL Bp 68/1/9/1–9. AÎdiate (Zemmour Noir, Mauri ta nia), fig ured by Sougy (1964, pl. Ad di tional ma te rial: frag mentary ven tral valve MNHN.F.A48127 42: 18–19); over twenty five ar tic u lated shells MNHN.F.A48143 from the local ity TM 809, MaÎder?, Mo rocco, coll. H. Hollard. from a small escarp ment 2 km ENE from Aguelt Oudiate el De scrip tion: Shell up to ca. 80 mm in width, semielliptic in out - Khyam (sec tion D-13, lev els 7–8), Zemmour Noir, Mau ri ta nia,

Fig. 7. Mid dle De vo nian Orthida from north west ern Af rica. A–M. Aulacella prisca (Schnur, 1851). A–E. Ar tic u lated shell ZPAL Bp 68/1/13/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F, G. Large ar tic u lated shell ZPAL Bp 68/1/13/3 from Aferdou in dor sal and ven tral views. H–L. Ar tic ulated shell ZPAL Bp 68/1/13/2 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an - te rior views. M. In complete ar tic u lated shell ZPAL Bp 68/2/13/1 from Maharch in ven tral view. N–NN. Tyersella canalicula (Schnur, 1851). N–R. Ar tic ulated shell ZPAL Bp 68/1/14/1 in dor sal, ven tral, lat eral, poste rior, and ante rior views. S. Dor sal valve ZPAL Bp 68/1/14/5. T–X. Artic u lated shell ZPAL Bp 68/1/14/2 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. Y–CC. Ar tic u lated shell ZPAL Bp 68/1/14/3 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. DD, EE. Artic u lated shell MNHN.F.A50413 from a small es - carp ment 2 km ENE from Aguelt Oudiate el Khyam; sec tion D13, bed 7–8, level 80 sensu Sougy (1964) (Zemmour Noir, Mau ri ta nia). FF–JJ. Artic u lated shell ZPAL Bp 68/1/14/4 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. KK, LL. Ven tral valve SMF 94848, in te rior (KK) and en large ment (LL) of the mus cle area. MM, NN. Ven tral valve SMF 94849, in te rior (NN) and en large ment (MM) of the mus cle area (vasculum myare arrowed). OO–XX. Phragmophora schnuri Cooper, 1955. Ar tic u lated shells from Aferdou in dorsal, ven - tral, lat eral, pos te rior, and an te rior views. OO–SS. Spec i men ZPAL Bp 68/1/16/1. TT–XX. Spec i men ZPAL Bp 68/1/16/2 258 A. T. HALAMSKI & A. BALIÑSKI

“upper Couvinian”, bed 80, coll. J. Sougy; two ar tic ulated shells, field; a pair of elon gate scars (one of them ar rowed in Fig. 7MM) one ven tral and two dorsal valves, and some fragments MNHN.F. situ ated antero-later ally to the muscle field might repre sent vas - A48099 from the lo cal ity Zl 16 (H 683), Mo rocco, coll. H. Hol- cula myaria. Dorsal inte rior: cardi nal process small; brachiophores lard. strong, flat tened, di ver gent at ca. 80–100°; den tal sock ets very De scrip tion: Shell subelliptic in out line, trans verse, up to 16.3 mm deep. Mus cle area 4.6 mm long in a spec i men 11.7 mm long, bor- in width, ventribiconvex or more sel dom aequibiconvex. Max i mal dered by two strong C-shaped ridges and sub di vided by a strong width an te ri orly to midlength. An te rior commissure straight to broa- me dian sep tum; pos te rior diductor scars nar row, an te rior ones dly and very shal lowly unisulcate. Dor sal valve with a shal low sul- broad, sep a rated by an in distinct ridge. cus, slit-like pos te ri orly, flat-bot tomed me di ally, and dis ap pear ing Re marks: The ma te rial from the “up per Couvinian” of the Zem- ante riorly. Dor sal interarea orthocline, linear; ventral one triangu- mour Noir (over fif teen ar tic ulated shells and iso lated dor sal and lar, apsacline, umbo weakly incurved. Or na menta tion of ra dial ven tral valves MNHN.F.A48142, 50413 from a small es carp ment costae and costellae, (1–)2(–3) per mm at an te rior mar gin. In te rior 2 km ENE from Aguelt Oudiate el Khyam; sec tion D-13, lev els not stud ied. 7–8; bed 80, coll. J. Sougy) is iden ti cal in shape to T. canalicula, Re marks: Vari abil ity of this wide spread Mid dle De vo nian spe - but has finer orna men ta tion (Fig. 7DD, EE). It is tenta tively con - cies was dis cussed in de tail by Biernat (1959) and Halamski sidered conspecific to the above-men tioned spe cies. (2009). Aulacella interlineata (Sowerby, 1840) is a Famennian Dis tri bu tion: This spe cies is known from the Holy Cross Moun- spe cies pres ent in Tafilalt (Halamski, un pub lished data), but the tains, Eifel, and from MaÎder; most prob a bly its strati graphic range re port from the Mid dle De vo nian of Zemmour Noir in Mau ri ta nia in cludes only the Eifelian (see Halamski, 2009 for de tails). (Sougy, 1964; most proba bly also that of Termier and Termier, Walmsley and Boucot (1975, p. 61) re port an iso lated occur rence 1950 from the same re gion) con cerns A. prisca (Halamski and of this taxon (neither de scrip tion nor fig ure) from the Emsian of Baliñski, 2009). Bathurst Is land (Arc tic Can ada); this re port is con sidered doubt ful. Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, Mau ri ta nia, Burma (Halamski, 2009, supple mented; re ported also from Bel - gium, Spain, and Iran); Eifelian and Givetian. Ge nus Phragmophora Cooper, 1955 Type spe cies: Phragmophora schnuri Cooper, 1955; Gerolstein, Eifel; cri noid beds (Eifelian) Family Dalmanellidae Schuchert and Cooper, 1931 Phragmophora schnuri Cooper, 1955 Ge nus Tyersella Philip, 1962 Fig. 7OO–XX Type spe cies: Tyersella typica Philip, 1962; Victo ria, Aus- v* 1955 Phragmophora schnuri Coo per, n. sp. – Coo per: p. tra lia; Siluro-De vo nian bound ary beds 52, pl. 12b, figs 12–26, pl. 14a, figs 1–7 [ubi syn.]. Tyersella canalicula (Schnur, 1851) v. 2009 Phragmophora schnuri Coo per, 1955 – Halamski, pp. Fig. 7N–NN 81–83, text-fig. 15, pl. 10, figs 39–48, pl.13, figs 26–48 [ubi syn.]. v* 1851 O[rthis] canalicula, n. sp. – Schnur: p. 13 Ma te rial: Two subcomplete ar tic u lated shells from Aferdou, vp 1885 Strophomena conf. pecten Linné sp. – Mau rer, pp. ZPAL Bp 68/1/16/1–2. 150–151, non pl. 6, fig. 3. De scrip tion: Shell up to 11 mm in width, wider than long, ventri- ? 1964 Isorthis sp. e. g. tetragona (Roemer, 1844) – Sougy, p. biconvex. Pos te rior mar gin straight, an te rior commissure broadly 437, non pl. 36, fig. 7. and shal lowly unisulcate. Dor sal valve mod er ately con vex, with a v. 2009 Tyersella canalicula (Schnur 1851) – Halamski, pp. median sulcus, slit-like pos te ri orly, broad, shallow, and U-shaped 77–79, text-fig. 12, pl. 12 figs 1–8, 28, 29, 31–34, pl. ante ri orly. Ventral valve subpyramidal, interarea apsacline, high. 13, figs 1–25 [ubi syn.]. Orna men ta tion of radial costae and costellae, 3–5 per mm at ante - Ma te rial: Twenty-four complete or subcomplete ar tic ulated shells, rior margin. a sin gle in complete dor sal valve from Aferdou, ZPAL Bp 68/1/14/ Re marks: The species was de scribed in de tail, in cluding its intra- 1–22; one ar tic ulated shell, one dor sal valve (MB un numbered, V. spe cific and ontogenetic vari abil ity, by Biernat (1959) and Ha- Ebbighausen’s col lec tion), and two subcomplete ven tral valves lamski (2009). SMF 94848, 94849 from MadÀne el Mrakib. Dis tri bu tion: This spe cies is known from the Eifelian of the Holy Other ma te rial ex am ined: Ven tral valve HLMD-Mr 8835, em- Cross Mountains, Eifel, and MaÎder (Halamski, 2009). bed ded in lime stone from Grube Hainau at Waldgirmes (Lahn Syncline). De scrip tion: Shell subrectangular in out line, trans verse, aequi- Family Schizophoriidae Schuchert and LeVene, 1929 biconvex to mod er ately ventribiconvex, up to 21.1 mm in width. Ge nus Schizophoria King, 1850 Mar gins rounded, antero-lat eral more so than the postero-lat eral. Type spe cies: Conchyliolithes Anomites resupinatus Dorsal valve with a flat-bot tomed, U-, V-, or slit-shaped me dian Mar tin, 1809; Derbyshire; Lower Carbon if er ous sulcus. Ventral valve trian gu lar to semielliptic in ante rior view; interarea apsacline to anacline, rather high; umbo thick, incurved. Schizophoria schnuri Struve, 1965 [1965a] Ante rior commissure weakly unisulcate; tongue subtrapezoidal to Fig. 8A–E, H–L V-shaped, low, nar row to mod er ately broad. Or na men ta tion uni - formly to un equally costellate, costae and costellae 1–3 per mm at * 1965a Schizophoria schnuri n. sp. – Struve: pp. 202–208, pl. an te rior mar gin. 19, fig. 4; pl. 20, pl. 21. Ventral inte rior: teeth stout, some times with cavi ties on the an- ? 1995 Schizophoria sp. – Struve, fig. 27. te rior part of the admedial sur face; dental plates pro long ing as v. 2012 Schizophoria (Schizophoria) schnuri Struve, 1965 – thick subparallel or slightly an te ri orly con verg ing ridges bor der ing Halamski, pp. 352–359, figs 3P–II, 4A–G, I, M, Q– the diductor scars; the latter narrow, bisected by a low to distinct EE, 5–7. median ridge, first un di vided, then with a me dian fur row, an te ri - Ma te rial: A single com plete ar tic u lated shell and four frag men- orly bi fur cat ing. Vas cula me dia sit u ated an te ri orly to the mus cle tary ones, one subcomplete ven tral valve, ZPAL Bp 68/1/15/1–6, MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 259

Fig. 8. Middle De vo nian Orthida from south ern MaÎder. A–E, H–L. Schizophoria schnuri. A. Dor sal valve ZPAL Bp 68/1/15/1 from Aferdou in dor sal view. B, D. Ven tral in te rior SMF 94860 from MadÀne el Mrakib. C, E. Ven tral in te rior SMF 94860. H–L. Ar tic u lated shell SMF 94862 from MadÀne el Mrakib in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–G. Schizophoria? sp. In complete in - ter nal mould ZPAL Bp 68/2/42/1 from Maharch in dor sal and ven tral views two complete artic u lated shells and a ventral valve from MadÀne vex. Dor sal valve with a very shal low V-shaped sulcus. On the dor - el Mrakib, SMF 94860, 94862, 94863. Forty ar tic ulated shells and sal valve and in the an te rior re gion of the ven tral valve poorly a sin gle ven tral valve from MadÀne el Mrakib, SMF 94899 and preserved costellate orna men ta tion with ca. 1–2 costellae per 2 mm MB unnum bered (V. Ebbighausen’s col lec tion). at ante rior margin. Traces of subparallel dental plates. De scrip tion: Shell subelliptic in out line, trans verse, dorsibicon- Re marks: Schizophoria? sp. dif fers from Schizophoria schnuri vex, up to 38.2 mm in width. Maxi mal width at mid-length or from Aferdou in the pres ence of a dor sal sulcus and or na menta tion slightly an teri orly. Dorsal interarea apsacline, ventral one ana- den sity. cline. An te rior commissure broadly uniplicate. Or na men ta tion of radial costae and costellae, 2–3 per mm at ante rior margin. Ventral Order Pentamerida Schuchert and Cooper, 1931 inte rior: teeth moder ately strong to stout; muscle field elongate, at - Family Gypidulidae Schuchert and Le Vene, 1929 tain ing one third to two fifth of the valve length, bor dered by a ridge, slightly me di ally in cised and sub di vided ei ther an te ri orly or Ge nus Gypidula Hall, 1863 throught its en tire length by a low me dian ridge. Dorsal in te rior Type spe cies: Gypidula typicalis Amsden, 1953 ( = un known. Pentamerus occidentalis Hall, 1858 non Hall, 1852); Inde - Re marks: Schizophoria schnuri is a long-last ing spe cies, sub di - pend ence, Iowa; Ham il ton Group (Mid dle De vo nian) vided into a se ries of chronosubspecies (Struve, 1965a; Halamski, 2012). The ma te rial from Aferdou is not iden ti fi able at the sub spe - Gypidula biplicata (Schnur, 1851) cies level. The mate rial from MadÀne el Mrakib shows intergra- Fig. 9A–E ding of morphotypes, re fer able to more than one sub spe cies, as de- limited by Struve (1965a), a situ a tion simi lar to that at Ska³y in the v* 1851 P[entamerus] biplicatus, n. sp. – Schnur: p. 8 Holy Cross Mountains (Halamski, 2012). v. 1966 Gypidula (Gypidula) cf. biplicata – Biernat, pp. 29– Dis tri bu tion: Eifel, Ardennes, Holy Cross Mountains, Moravia, 30 pl. 1, fig. 11. MaÎder, Burma (Halamski, 2012). Ma te rial: A single complete, slightly decorticated shell from Aferdou, ZPAL Bp 68/1/2/1. Schizophoria? sp. De scrip tion: The sin gle avail able shell is subpentagonal in out - Fig. 8F, G line, strongly ventribiconvex, 20.4 mm wide, 19.0 mm long, and 14.9 mm thick; max i mal width at an te rior third of the shell length. Ma te rial: A single in complete speci men from Maharch, ZPAL Bp Dorsal valve mod er ately con vex, subtriangular in an te rior view, 68/2/42/1. with a flat-bot tomed sulcus in the an te rior third; a broad me dian Description: The single available speci men is el liptic in outline, ca. plica present in the im medi ate proxim ity of the ante rior margin. 23 mm wide, >16 mm long, and >8 mm thick, weakly dorsibi- con - Ven tral valve strongly con vex; umbo thick, beak strongly in cur- 260 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 261

ved; two low plicae, sep a rated by a shal low fur row, pres ent in the Ven tral in te rior (Fig. 9Q): spondylium supported by a me dian im me di ate prox im ity of the an te rior mar gin. An te rior commissure sep tum, the lat ter thick poste ri orly, high and blade-like an te ri orly. plicosulcate, tongue subtrapezoidal, 9.3 mm wide at the base and Dorsal in te rior (Fig. 9R): mus cle area ex tend ing at most to the one 4.0 mm at the top, 6.7 mm high. Shell smooth. Inte rior not stud ied. third of the valve length, in ner hinge plates con verg ing on the Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, pos si bly Ar- valve floor, coales cent with crural bases, outer hinge plates flat- menia (Abramân, 1974); Mid dle De vo nian. tened, rather wide, sepa rated by a rounded in den ta tion from wide, flat tened outer socket ridges. Gonoglyphs on dorsal and ven tral Ge nus Glyptogypa Struve, 1992 valve floors ad jacent to muscle fields. Re marks: Halamski (un pub lished) dis tin guished G. multiplicata Type spe cies: Pentamerus galeatus var. multiplicata (Roemer, 1856) from cen tral Eu rope (Eifel, Holy Cross Moun- Roemer, 1856, Schwiersheim near Prüm, Eifel; Middle tains) and G. chouberti (Termier and Termier, 1950) from North - De vo nian (prob a bly Eifelian) ern Af rica (Mo rocco, Mau ri ta nia), on the ba sis of or na men ta tion Glyptogypa multiplicata (Roemer, 1856) den sity: in the sam ples in vesti gated, the quo tient of ventral costae Fig. 9F–R number and the shell width var ied be tween 0.81 and 1.26 for the for mer pre sumed species and be tween 1.30 and 1.59 for the lat ter. * 1856 Pentamerus galeatus var. multiplicata – Roemer: p. The value for the in ves ti gated spec i men (0.64 to 1.26, mean 0.82; 352, pl. 2, fig. 9a-c. N = 20) falls clearly within the vari a tion range of the Eu ro pean vp 1964 Gypidula sp. – Sougy, p. 455, pl. 44, fig. 4a–c; non p. sample. Given the un cer tainty on ex act strati graphic po si tion of 358, pl. 33, fig. 12. these brachi o pods, it is un clear, whether the two morphotypes rep - v. 1966 Gypidula (Ivdelinia) sp. cf. G. (Ivdelinia) multiplicata re sent sep a rate mem bers of an evo lu tion ary lin eage, co-oc cur ring – Biernat, pp. 33–34, pl. 2, figs 2–4. spe cies, or only end members of a con tin u ous vari a tion se ries. v. 1995 Glyptogypa sp., Gr. multiplicata (C. F. Roemer 1844) Glyptogypa multiplicata is proba bly the spe cies, which is the – Struve, p. 100, fig. 31. most encrusted by epizoans (bryo zoans, microconchids, favositid v. 1995 Gypidula sp. cf. chouberti Termier and Termier 1950 tabu lates) in the ma te rial studied (Fig. 9K–P). This is argu ably re - – Struve, p. 100, fig. 30. lated to its large size and strong or na menta tion, as shown by 1995 Glyptogypa cf. multiplicata (Roemer, 1856) – Go- Zapalski (2005) for co eval de posits of the Holy Cross Mountains. defroid, pp. 83–85, fig. 6a–e. Dis tri bu tion: Rhenish Slate Moun tains, Ardennes (Godefroid, Ma te rial: One hun dred and thirty-one com plete or subcomplete 1995), Holy Cross Mountains, MaÎder, and Mau ri ta nia; Mid dle (of ten de formed) ar tic ulated shells SMF 94864–94868 and MB De vo nian. unnum bered (V. Ebbighausen’s col lec tion), three ven tral, SMF 94854–94855, 94870, and six dor sal valves, SMF 94856–94859, Ge nus Ivdelinia Andronov, 1961 94869, 94871, and over thirty frag ments from the Drotops beds at Type spe cies: Gypidula ivdelensis Khodalevich, 1951, MadÀne el Mrakib; a single artic ulated shell ZPAL Bp 68/1/1/1 Ivdel, East ern Urals; Emsian–Eifelian from Aferdou. Ad di tional ma te rial: Artic u lated shells SMF 54779, 54781 from Ivdelinia (Ivdelinia) pulchra Franchi, Schemm-Gregory the Eifelian of Jebel Issomour (Fp. 2999, northern MaÎder), fig - and Klug, 2012 ured by Struve (1995); one subcomplete ar tic ulated shell and one Figs 10A–P, 11 frag ment MNHN.F.R10274 from the “Givetian” of Aguelt Ou- diate (Zemmour, Mauri ta nia), coll. J. Sougy. v. 1995 Ivdelinia sp., cf. formosa (Schnur, 1851) – Struve, p. De scrip tion: Shell up to 31.7 mm in length, nearly as wide as long 100, figs 28, 29. (width-to-length ra tio 0.84 to 1.19, mean 1.02; N = 21), moder - v. 1998 Ivdelinia sp. – Kaufmann, p. 59, pl. 13, fig. 11. ately ventribiconvex, some times slightly asymmet ric; maxi mal * 2012 Ivdelinia pulchra sp. nov. – Franchi et al.: pp. 5–6, fig. width in the ante rior third. Dor sal valve broadly tri an gular, with a 3. flat-bot tomed sulcus in the an te rior half; interarea obscured by . 2013 Ivdelinia pulchra – Tessitore et al., p. 25, text-figs 2 over hang ing umbo. Ven tral valve par a bolic in an te rior pro file, [p.p.], 3, 5, 11. with a thick umbo and a strongly incurved beak; no fold; palin- Ma te rial: Over one hun dred ar tic ulated shells from Aferdou, trope high, concave, with well marked beak ridges, apsacline near mostly from the Ivdelinia bed, a large part of them com plete and the hinge mar gin. An te rior commissure unisulcate; tongue subtra- well pre served; col lec tion number ZPAL Bp 68/1/3. pezoidal, in a sin gle case subtriangular, its width 0.29 to 0.46 Ad di tional ma te rial: ar tic u lated shells SMF 54776, 54777 from (mean 0.35; N = 20) of that of the shell, its height 0.17 to 0.55 Jebel Issomour (northern MaÎder), fig ured by Struve (1995). (mean 0.35; N = 19) of the shell thick ness. Orna menta tion of ir reg - De scrip tion: Shell up to 47.5 mm in width, slightly trans verse ular costae and costellae, the lat ter aris ing by both bi fur ca tion and through aequidimensional to slightly elon gate, strongly ventribi- in ter ca la tion, 13–24 (mean 18; N = 18) on the dor sal valve, 17–30 con vex. Ven tral valve strongly con vex, umbo swollen, beak stron- (mean 21; N = 19) on the ventral valve. Costellae some times dis- gly incurved, nearly touch ing the dor sal valve; a low and flat fold ap pear be fore reach ing the an te rior mar gin. in the an te rior half. Ventral palintrope smooth, as wide as the

Fig. 9. Middle De vonian Pentamerida from southern MaÎder. A–E. Gypidula biplicata (Schnur, 1851), Aferdou. Artic ulated shell ZPAL Bp 68/1/2/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–R. Glyptogypa multiplicata (Roemer, 1856). F–J. Artic u lated shell ZPAL Bp 68/1/1/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. K–O. Ar tic u lated shell SMF 94867 from MadÀne el Mrakib in dor sal, ven tral, lat eral, pos te rior, and an te rior views. P. Ar tic u lated shell SMF 94871 from MadÀne el Mrakib in lat eral view. Q. Ven tral in te rior SMF 94855 from MadÀne el Mrakib. R. Dor sal in te rior SMF 94857 from MadÀne el Mrakib. S–X. Antirhynchonella sublinguifera (Mau rer, 1885). S–U. Frag mentary shell ZPAL Bp 68/2/18/1 from Maharch in dor sal, ven tral, and an te rior views. V–X. Frag mentary shell ZPAL Bp 68/1/18/1 from Aferdou in dor sal, lat eral, and poste rior views 262 A. T. HALAMSKI & A. BALIÑSKI

Fig. 10. Ivdelinia (Ivdelinia) pulchra Franchi, Schemm-Gregory and Klug, 2012. A–E. A middle-sized artic u lated shell ZPAL Bp 68/1/3/1 from marls over ly ing the lumachelle in Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–J, K–O. Two large ar- tic u lated shells ZPAL Bp 68/1/3/2,3 from the lumachelle in Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. P. Mag ni fi ca- tion of the growth lines ob served in the an te rior re gion of a ven tral valve of the spec i men ZPAL Bp 68/1/3/4 from the lumachelle

valve, me di ally lower, higher in mar ginal re gions, sep a rated from te ri orly in the sec tioned shell; outer hinge plates con verg ing on the the rest of the valve by an acute ridge. Dorsal valve mod erately valve floor and forming cruralium; an te rior tips of the in ner hinge con vex postero-me di ally, with el e vated flanks; sulcus flat and plates un supported give rise to the crescen tic in cross section crura shal low, vis i ble in the an te rior half. Dor sal palintrope ab sent. An - (Fig. 11). te rior commissure unisulcate, de flec tion rect an gu lar to trapezoi- Re marks: Ivdelinia pulchra is in cluded within the sub ge nus I. dal, rather low to mod er ately high, oc cu py ing a half to two-third of (Ivdelinia) on ac count of the presence of a well de vel oped ven tral the shell width. sep tum, de spite its ex ter nal sim i lar ity to I. (Ivdelinella) ellesme- Orna menta tion of thick, rounded costae and costellae, ob serv able rensis Brice, 1982 from the Emsian–Eifelian of Can ada (Brice, on both valves from about 1 cm from the umbones, sep a rated by 1982), the type spe cies of the sub ge nus Ivdelinella Brice, 1982 nar rower fur rows. Costae and costellae bi sected by shal low fur - [flat tened costae, as op posed to strongly raised costae or plicae in rows, in some cases a cost(ell)a passes in this way into a pair of I. (Ivdelinia); Blodgett et al., 2002]. This spe cies was iden ti fied in costellae; up to two bi fur ca tions of a pri mary costa have been ob - the Holy Cross Mountains, where it oc curs as a rare compo nent in served. Costae bor der ing the ven tral fold are stron ger than the the Eifelian–Givetian bound ary beds (Halamski, 2004a). The valid other ones. Costae and costellae 3–4 on the fold, 4–5 in the sulcus, name was in tro duced by Franchi et al. (2012), on the ba sis of the up to 5 on each lat eral flank. Con cen tric growth lines were ob - ex ter nal mor phol ogy of thir teen spec i mens. De tailed de scrip tion served at an te rior mar gin, 3–4 per mm (Fig. 10F). of ex ter nal and in ter nal mor phol ogy and syn on ymy lim ited to Ventral inte rior: shell substance thick; me dian septum well devel - North Afri can occur rences of the species are presented herein. The oped, sup port ing the spondylium pos te ri orly, then dis ap pear ing. palaeo ec ol ogy of the Ivdelinia brachi opod bed was dealt with by Dorsal in te rior: hinge plates very long, ex tend ing about 11 mm an- Tessitore et al. (2013). MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 263

Fig. 11. Trans verse se rial sec tions of Ivdelinia (Ivdelinia) pulchra Franchi, Schemm-Greg ory and Klug, 2012 through the shell ZPAL Bp 68/1/3/5. Dis tances mea sured in milli metres from the tip of the ven tral umbo

Dis tri bu tion: Middle De vo nian of Eifel, Holy Cross Mountains, Family Clorindidae Rzhonsnitskaya, 1956 and MaÎder (Halamski, 2004a). Ge nus Antirhynchonella 2hlert in Fischer, 1887 Type spe cies: Atrypa linguifera Sowerby, 1839; Stumps Ge nus Devonogypa Havlíèek, 1951 Wood, Malvern hills, western England; Wen lock Lime- Type spe cies: Gypidula (Devonogypa) spinulosa stone, Si lu rian Havlíèek, 1951; Èelechovice na Hané (Moravia); Middle Range: Eifelian to early Givetian. Up to now the last known oc - De vo nian cur rence of the ge nus (Boucot et al., 2002) was in the Mi³oszów Lime stone of the Holy Cross Mountains of un clear strati graphic Devonogypa spinulosa (Havlíèek, 1951) posi tion, con tain ing both Eifelian and Givetian spe cies (Halamski, Fig. 12A–M 2009) and there fore equiv ocal. The pres ence of Antirhynchonella * 1951 Gypidula (Devonogypa) spinulosa nov. nom. – Hav- in the lower Givetian is con firmed herein. líèek: pp. 5–7, pl. 2, figs 2, 3, 5. v. 1966 Gypidula (Devonogypa) spinulosa Havlíèek, 1951 – Antirhynchonella sublinguifera (Maurer, 1885) Biernat, pp. 28–29, pl. 1, figs 1–5, pl. 2, fig. 10. Fig. 9S–X . 2013 Devonogypa sp. – Tessitore et al., p. 25; text-figs 2 [p.p.], 4, 10b, 12. v* 1885 Pentamerus sublinguifer n. s. – Mau rer: pp. 218–221, Ma te rial: Twenty-four spec i mens from Aferdou, one complete, pl. 9, figs 9, 10. three subcomplete, the other frag mentary, ZPAL Bp 68/1/17/1– vp 1964 Dicamara prunulum Schnur – Sougy, p. 451, pl. 12, 24; two spec i mens from MadÀne el Mrakib (MB un numbered, V. fig. 13a–c; non pl. 42, fig. 1a–b. Ebbighausen’s col lec tion). v. 1966 Antirhynchonella linguiformis n. sp. – Biernat, pp. Ad di tional ma te rial: In com plete ar tic u lated shell MNHN.F.A 35–36, text-fig. 5, pl. 2, figs 2–5. 48135 (microornamentation not pre served) from the lo cal ity 2389, Ma te rial: Six in complete ar tic u lated shells, one from Aferdou, Zemmour Noir, Mau ri ta nia, coll. J. Sougy. ZPAL Bp 68/1/18/1, and three from Maharch, ZPAL Bp 68/2/18/ De scrip tion: Shell rounded in out line, most of ten lon ger than 1–2. wide, rarely wider than long, strongly ventribiconvex, up to ca. 67 Ad di tional ma te rial: Artic u lated shell MNHN.F.R10264 from mm in length. Dor sal valve weakly con vex, broadly tri an gular in the “Givetian”, 800 m SSE from Aguelt Nebka, Zemmour Noir, an te rior view. Ven tral valve strongly con vex with steep flanks and Mau ri ta nia, coll. J. Sougy; fig ured by Sougy (1964, pl. 12: 13). moder ately con vex to flat tened me dian re gion; umbo very large De scrip tion: Shell lon gi tu di nally el lip tic in out line; dor sal valve (one fifth to a quar ter of the to tal shell length), beak strongly trans verse (width-to-length ra tio 1.08–1.28), strongly ventribi- incurved. Ventral palintrope con cave, delthyrium oc cu pying ca. convex, up to ca. 40 mm in width. Dor sal valve in an te rior view re - one third of its width. An te rior commissure rectimarginate. Or na - sembling a Gauss curve. Ven tral palintrope weakly con cave. Ante - menta tion (sel dom preserved; Fig. 12L) of mi cro scopic spines, ca. rior commissure unisulcate in smaller indi vid u als, parasulcate in 0.1–0.2 mm in di ame ter; oth er wise shell smooth. Ven tral in te rior larger ones, tongue broad and low, rounded to subtrapezoidal. (Fig. 12A): spondylium; dor sal in te rior: weakly di ver gent in ner Shell en tirely smooth except for growth lines. hinge plates ex tend ing up to ca. one third of the valve length; oth - Re marks: The Maharch sample con tains both uniplicate and para- er wise not stud ied. plicate indi vid u als. This is an exam ple of intraspecific variabil ity Dis tri bu tion: Holy Cross Mountains, Moravia, MaÎder, and Mau- and an ar gument for tak ing the spe cies dis cussed broadly (see syn- ri ta nia; Mid dle De vo nian. onymy). Simi lar intraspecific variabil ity was described by Koz- 264 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 265

³owski (1929) and Baliñski (2012) in closely re lated Si lu rian to De vo nian (Eifelian and lower to mid dle Givetian). Mid dle De vo nian ge nus Clorinda. Dis tri bu tion: Rhenish Slate Moun tains, Holy Cross Mountains, Kransia subcordiformis (Schnur, 1853) MaÎder, and Mau ri ta nia; Mid dle De vo nian. Fig. 13JJ–NN v* 1853 Terebratula subcordiformis n. sp. – Schnur: p. 186, pl. Order Rhynchonellida Kuhn, 1949 25, fig. 6a–k. Family Hebetoechiidae Havlíèek, 1960 v. 1941a Uncinulus subcordiformis (Schnur) – Schmidt, pp. Ge nus Kransia Westbroek, 1967 19–20, pl. 2, fig. 20, pl. 4, fig. 71 [ubi syn.]. Type spe cies: Terebratula parallelepipeda Bronn, 1837 Ma te rial: A single slightly in complete and decorticated spec i men sensu Westbroek, 1967, Gerolstein Syncline, Eifel; from Aferdou, ZPAL Bp 68/1/4/1. De scrip tion: The sin gle avail able shell is 20.1 mm wide, 15.7 mm Eifelian long, and 11.5 mm thick, dorsibiconvex; the out line is roundly Kransia parallelepipeda (Bronn, 1834 in 1834–38 subrectangular. Ven tral valve with a rather shal low flat-bot tomed [“1835–37”]) sulcus, vis i ble in the an te rior half; umbo strong, beak incurved. Fig. 13K–II Dorsal valve with a flat fold de vel oped in the ante rior half, sepa - rated from lat eral flanks of the shell by steep slopes. An te rior v*p 1834 Terebratula Wilsoni [parallelepipeda] – Bronn: p. 73. commissure uniplicate, tongue semi cir cu lar, evenly con vex, ca. 10 1960 Uncinulus parallelepipedus Bronn – Gevin, p. 162. mm wide. Or na menta tion of flat costae and costellae, 7 on the top v. 1964 Uncinulus parallelepipedus (Bronn, 1837) – Sougy, of the fold, 2 on its slopes, ca. 17 on lat eral flanks. p. 445, pl. 42, fig. 7. Re marks: Kransia subcordiformis dif fers from K. parallelepi- v. 1966 Uncinulus parallelepipedus (Bronn, 1837) – Biernat, peda in larger size, usu ally greater width to length ratio, and a pp. 90–91, text-fig. 30, pl. 17, figs 8–11. slightly weaker fold. v. 2005 “Kransia parallelepipeda” (Bronn, 1837) – Schemm- Dis tri bu tion: Eifel, Holy Cross Mountains, Cantabrian Moun tains Greg ory and Jansen, p. 27. (Mohanti, 1972), MaÎder; Mid dle De vo nian (Givetian only?). Ma te rial: One hun dred ar tic ulated shells from Aferdou, sev eral of them complete and well pre served, ZPAL Bp 68/1/19/1–91. Kransia? coronata (Kayser, 1871) Ad di tional ma te rial: Six artic ulated shells (mostly complete; partly Fig. 13OO–SS mislabelled as “Uncinulus primipilaris”) MNHN.F.A 48128, 48129 from the lo cal ity 2073F, Zemmour Noir, Mau ri ta nia, coll. J. Sougy. v* 1871 Rhynchonella coronata n. sp. – Kayser: pp. 512–513, De scrip tion: Shell in shape of a rounded hep ta gon, usu ally sligh- pl. 9, fig. 5. tly trans verse (width-to-length ra tio 0.90 to 1.23, mean 1.08; N = v. 1941a Uncinulus coronatus (Kayser) – Schmidt, p. 24, pl. 2, 20), moder ately to mark edly dorsibiconvex, up to 15.8 mm in fig. 24, pl. 4, fig. 73, pl. 6, fig. 18. width but typ i cally about 11–12 mm wide. Dorsal valve with a flat ? 1971 “Uncinulus” coronatus (Kayser) – Drot, pp. 71–72, fold in the ante rior half. Ventral valve with a wide and shal low, text-pl. 1, pl. 3, fig. 1. nearly flat-bot tomed sulcus, most often bor dered by two slight el e - Ma te rial: Eighteen artic u lated shells (three com plete, three sub-c va tions of the dor sal valve per cep ti ble only in the im medi ate prox - omplete, and five in complete) from the Drotops beds at MadÀne el imity of the an te rior mar gin. Beak weakly incurved; fo ra men Mrakib, SMF 98178 and MB unnum bered (V. Ebbighausen’s col - permesothyrid, up to ca. 1 mm in diam e ter; umbo rel a tively fine. lec tion); a single artic ulated shell ZPAL Bp 68/1/5/1 from Afer- Ante rior commissure uniplicate to paraplicate; tongue high, roun- dou. ded, in some cases lateral slopes subvertical, subparallel to each De scrip tion: Shell up to 20.9 mm wide, trans verse (width-to- other. Or na menta tion of low, rounded costae and costellae, sep a - length ra tio 1.15 to 1.41, mean 1.24; N = 4), strongly dorsibicon- rated by sublinear fur rows, (3–)5–7 on the fold, 1–2 on the flanks vex. Dor sal valve with a low flat tened fold de vel oped in the an te - of the fold, 9–10 on each lat eral flank. rior third. Ven tral valve weakly con vex in pos te rior third, with a Re marks: This spe cies was treated in a broad sense, owing to the shallow flat-bottomed sulcus in the ante rior two-thirds; lat eral in suf fi ciently pre cise first de scrip tion (Bronn, 1835–37). The flanks con cave; umbo rather slen der but poorly pre served. An te - choice of a lectotype/neotype from the Eifelian of the Eifel (MCZ rior commissure uniplicate, tongue subtrapezoidal, its bot tom col lec tions; Halamski un pub lished) will al low to sub di vide the width 0.44–0.56, top width 0.33–0.48 of that of the shell, its height widely un der stood K. parallelepipeda s.l. and dis tin guish be tween 0.55–0.72 of the shell thick ness. Orna menta tion of strong, acute K. parallelepipeda s.s. (incl. Terebratula angulosa Schnur, 1851; costae and costellae, the lat ter ap pear ing by bi fur ca tion, 7–8 on the basal Eifelian to lower or middle Givetian) and its prob a ble evo lu - fold, 5–7 in the sulcus, 8–10 on lat eral flanks. In te rior not stud ied. tionary de scendant (as yet un named) in the late Givetian (e.g., Re marks: This spe cies is dis tin guished from re main ing rep re sen - Brice and Morzadec, 1983). ta tives of Kransia by the form of the dor sal valve and stronger, Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, and south ern acute costae, whence the quota tion mark afer the genus name. limb of the Tindouf Syncline (Schmidt, 1941a; Biernat, 1966; Dis tri bu tion: Eifel (Schmidt, 1941a), Holy Cross Mountains Halamski, 2004a; Schemm-Greg ory and Jansen, 2005); Mid dle (Biernat, 1966), MaÎder; Mid dle De vo nian (Eifelian only?).

Fig. 12. Devonogypa spinulosa (Havlíèek, 1951) from Aferdou. A–D. Ven tral valve in dor sal, ven tral, pos te rior, and lat eral views. E–G. In com plete ar tic u lated shell in dor sal, ven tral, and an te rior views. H–K. Subcomplete ar tic u lated shell in dor sal, ven tral, pos te rior, and lat - eral views. L–M. Ven tral valve in ven tral view; gen eral view (M) and microornamentation (L) 266 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 267

Ge nus Beckmannia Mohanti, 1972 Ma te rial: Three ar tic u lated shells from Aferdou (two complete), Type spe cies: Uncinulus minor beckmanni Schmidt, 1951; ZPAL Bp 68/1/21/1–3; seven in di vid u als from Maharch (one sub- Letmathe, Rhenish Slate Mountains; Middle De vonian complete ar tic u lated shell), ZPAL Bp 68/2/21/1–6, 68/2/43/1. (lower Givetian?) De scrip tion: Shell ap prox i mately iso met ric to trans verse, very strongly dorsibiconvex, up to 25.9 mm in width (incom plete speci - Beckmannia beckmanni (Schmidt, 1951) mens suggest a max i mal width up to 30 mm). Maxi mal width Fig. 13A–J about shell midlength, sel dom about the an te rior fifth of the shell length. Dor sal valve very strongly con vex, with a slightly flat tened * 1951 Uncinulus mi nor beckmanni n. ssp. – Schmidt: pp. fold in an te rior two-fifths. Ven tral valve moder ately to strongly 89–90, pl. 1, figs 1–3. con vex in umbonal re gion, flat on flanks, with a wide flat-bot - 1985 Beckmannia mi nor beckmanni (Schmidt 1951) – tomed to W-shaped sulcus an te ri orly; umbo low, beak incurved. Brice, pp. 135–136, text-fig. 3, pl. 1, figs 10–13 [ubi Ante rior commissure uniplicate, tongue very broad, oc cu pying syn.]. 0.6–0.7 of the shell width, bor dered by ver ti cal deflexions of the Ma te rial: ten ar tic ulated shells from Aferdou, ZPAL Bp 68/1/20/ commissure. Orna menta tion of low, flat tened, grooved costae, 1–10. sep a rated by very nar row furrows, 9–13(–19) on the tongue, 8–12 De scrip tion: Shell in out line of a rounded pen ta gon, as wide as on the fold, 12–16 on flanks. In te rior not stud ied. long, mod er ately ventribiconvex. Max i mal width about mid- Re marks: This is a very vari able species; the speci men ZPAL Bp length. Dor sal valve mod er ately con vex; a very low flat fold some - 68/2/43/1 from Maharch (Fig. 14F–J) is slightly out side of the times visi ble in the an te rior fourth. Ven tral valve con vex, with a usual extent of intraspecific variabil ity in its exceed ingly fine nearly imper cep ti ble sulcus in an te rior half; umbo fine, elon gate, costation (19 costae on the tongue com pared to 9–13 in the re - beak incurved. An te rior commissure uniplicate; tongue low, roun- maining mate rial) and in the po sition of the max i mal width of the ded, oc cu py ing ca. 0.6–0.8 of the shell width. Or na menta tion of shell (half vs. four-fifths of the shell length). It is quite simi lar to low, rounded costae sep a rated by very nar row interspaces, 3–4 on the in di vid ual SMF XVII 749a from the Eifel (Fig. 14P–T), mis- fold, 0–1 on its flanks, ca. 6 on each lateral flank. In te rior not stud - iden ti fied by Schmidt (1941a, pl. 5: 9) as Ladogifornix fornicatus ied. (Schnur, 1851). The holotype of the lat ter spe cies is il lustrated for Re marks: As far as the ex teri ors are concerned, the speci mens com par i son in Fig. 14A–E. stud ied are in dis tin guish able from the type col lec tion of Beck- Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder; Middle De- mannia beckmanni (Schmidt, 1951) from the Mid dle De vonian of vo nian. the Rhenish Slate Mountains. Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, Cantabrian Mountains (Mohanti, 1972), Montagne Noire (Massif Cen tral, Family Septalariidae Havlíèek, 1960 France; Brice, 1985); Mid dle De vonian (Givetian, mostly about Ge nus Septalaria Leidhold, 1928 Mid dle varcus Zone). Type spe cies: Terebratula ascendens Steininger, 1853; Eifel; Mid dle De vo nian Family Hypothyridinidae Rzhonsnitskaya, 1956 Septalaria gracilis (Gürich, 1896) Ge nus Glosshypothyridina Rzhonsnitskaya, 1978 Figs 15, 16K–O, Q–OO Type spe cies: Rhynchonella procuboides Kayser, 1871; v* 1896 Camarophoria gracilis n. sp. – Gürich: pp. 278–279, Eifel, Germany; Eifelian. pl. 7, fig. 3a–g. Range: Eifelian to early Givetian, emended herein (Eifelian only . 1909 Liorhynchus gracilis Gürich – Sobolew, p. 503, pl. 6, ac cord ing to Sav age in Savage et al., 2002) on ac count of pres ence figs 8–12, 16. of G. procuboides at Maharch. vp 1934 Septalaria ascendens (Steininger) – Torley, p. 78, text-fig. Gr. A, figs 11–12, pl. 2, figs 49–53, 58–57, non pl. 2, figs 54–57. Glosshypothyridina procuboides (Kayser, 1871) v. 1966 Septalaria cf. gracilis (Gürich, 1896) – Biernat, pp. Fig. 14F–Y 97–98, pl. 7, fig. 6. v* 1871 Rhynchonella procuboides n. sp. – Kayser: pp. 513– 1971 «Pugnax» pugnoides latus Herta Schmidt, 1941 – 514, pl. 9, fig. 3. Drot, pp. 89–90, pl. 3, fig. 4. vp 1941a Camarotoechia triloba fornicata (Schnur 1853) – v. 1975 Septalaria descendens n. sp. – Schmidt, pp. 93–96, Schmidt, pp. 12–13, pl. 5, fig. 9, non pl. 4, fig. 59. text-figs 2, 6, pl. 3, figs 15–16, pl. 4, figs 17–18 [ubi 1964 Hypothyridina procuboides (Kayser) – Sougy, p. 450, syn.]. pl. 42, fig. 8. 1985 Septalaria aff. descendens Schmidt 1975 – Brice, pp. v. 1966 Hypothyridina cf. procuboides (Kayser) – Biernat, 138–139, text-fig. 5, pl. 2, figs 1–3. pp. 105–106, pl. 21, fig. 2. Ma te rial: Twenty-two artic ulated shells (most of them complete)

Fig. 13. Middle De vonian Rhynchonellida from southern MaÎder. A–J. Beckmannia beckmanni (Schmidt, 1951). A–E. Speci men ZPAL Bp 68/1/20/1. F–J. Spec i men ZPAL Bp 68/1/20/2. K–II. Kransia parallelepipeda (Bronn, 1834 in 1834–38 [‘1835–37’]). K–O. Spec i men ZPAL Bp 68/1/19/1. P–T. Spec i men ZPAL Bp 68/1/19/2. U–Y. Spec i men ZPAL Bp 68/1/19/3. Z–DD. Spec i men ZPAL Bp 68/1/19/4. EE–II. Speci men ZPAL Bp 68/1/19/5. JJ–NN. Kransia subcordiformis (Schnur, 1853). Spec i men ZPAL Bp 68/1/4/1. OO–SS. Kransia? coronata (Kayser, 1871). Spec i men ZPAL Bp 68/1/5/1. All pho to graphs are of ar tic u lated shells from Aferdou in dorsal, ven tral, lat eral, pos te rior, and an te rior views 268 A. T. HALAMSKI & A. BALIÑSKI

Fig. 14. Middle De vo nian Rhynchonellida from southern MaÎder and Eu rope. A–E. Ladogifornix fornicatus (Schnur, 1851). Holotype from Schnur’s collec tion (Bonn Uni ver sity; un numbered) from the Mid dle De vonian of the Eifel. F–Y. Glosshypothyridina procuboides (Kayser, 1871). F–J. Spec i men ZPAL Bp 68/2/43/1 from Maharch. K–O. Spec i men ZPAL Bp 68/1/21/1 from Aferdou. P–T. Spec i men SMF XVII 749a from Uxheim (Eifel), Eifelian. U–Y. Spec i men ZPAL Bp 68/2/21/1 from Maharch. All pho to graphs are of ar tic ulated shells in dor sal, ven tral, lat eral, pos te rior, and an te rior views

from Aferdou, ZPAL Bp 68/1/22/1–22, and two (one com plete) gue broad, occu pying (0.5–)0.58–0.69 of the shell width, high, from Maharch, ZPAL Bp 68/2/22/1–2. trap e zoidal. Orna menta tion of rather low, rounded costae, ap pear - Other ma te rial ex am ined: one complete and three fragmen tary ing at ca. 5 mm from the umbo, sepa rated by nar rower fur rows, ar tic ulated shells MGUWr 1966s from Œniadka (£ysogóry Re gion, 6–11 on tongue, 8–18 on flanks. Holy Cross Mountains), syntypes of Camarophoria gracilis Gü- The inte rior of the single partly recrystallised sec tioned spec i - rich, 1896; two ar tic ulated shells SMF 29599/1–2 from Bilve- men (Fig. 15) does not re veal the pres ence of den tal plates. Dor sal ringsen (Sauerland), coll. K. Torley (sub Septalaria ascendens). in te rior with well de vel oped septalium; median sep tum high and De scrip tion: Shell subpentagonal in out line, in some cases asym- fairly long; pos te rior re gion of the septalium lined with thick ened met ric, most of ten trans verse, sel dom as wide as long, strongly car di nal pro cess; outer hinge plates subhorizontal with slightly dorsibiconvex, up to 27.4 mm in width. Dorsal valve with a flat - dor sally curved in ner mar gins; crura short, slightly lat er ally di ver - tened fold ap pear ing slightly an te ri orly to midlength, flanks steep. gent and with ven trally bent dis tal ends. Ven tral valve con vex in umbonal re gion, flat tened on flanks, with Re marks: Se lected met ric char ac ters of three pop u la tions of Sep- a wide and shal low flat-bot tomed sulcus an te ri orly; umbo fine, talaria gracilis are com pared in Table 1. beak mod er ately incurved. An te rior commissure uniplicate, ton- MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 269

Ta ble 1 Com par i son of se lected bio met ric char ac ter is tics of Septalaria gracilis from the Holy Cross Mountains, Sauerland, and MaÎder. Data for the Holy Cross Moun- tains, either after Gürich (1896) or calcu lated after photo - graphs in Sobolew (1909); those for Sauerland and MaÎder af ter pres ent au thors

Taxon Septalaria gracilis Character Holy Cross Mts. Sauerland MaÎder 14.4 19.5 24.3 Width [mm] (11–19) (17.4–20.7) (19.8–27.3) 1.15 1.25 1.34 Width to length ratio (1.04–1.36) (1.13–1.38) (1.21–1.43) 3.6 4.9 8.2 Costae on tongue [(2–)4] (2–7) (7–11) 0.27 0.25 0.34 Costae/shell width (0.18–0.31) (0.11–0.34) (0.26–0.40) 0.48 0.52 0.65 Costae/tongue width Fig. 15. Trans verse se rial sec tions of Septalaria gracilis (Gürich, (0.33–0.51) (0.42–0.65) (0.55–0.82) 1896) through the shell ZPAL Bp 68/1/22/6 from Aferdou. Dis - Number of specimens 5/8 8 8 tances mea sured in millimetres from the tip of the ven tral umbo

Each sample has its own biomet ric characteristicts, but all three Langenaubach has two costae on each flank, is larger, and has also are con sidered conspecific, on the ba sis of over all sim i lar ity of a reg u larly con vex dor sal valve. shape and intergrading val ues of bio metric char ac ters. In view of such marked vari abil ity, Septalaria aff. descendens sensu Brice Paulinaerhynchia new genus (1985) from La Serre in the Montagne Noire (Mas sif Cen tral, Type spe cies: Paulinaerhynchia paulinae gen. et sp. nov., France) should prob a bly be con sid ered as rep re sent ing the spe cies dis cussed, as well. Un for tu nately, the Mo roc can ma te rial of Septa- as be low laria, de scribed by Drot (1964), could not be traced at the MNHN. Derivatio nominis: As for the type species. Dis tri bu tion: Eifel, Holy Cross Mountains (North), and MaÎder, Di ag no sis: Large-sized, strongly costate Pugnacidae; dorsal sep- prob a bly also Montagne Noire; Middle De vo nian. tum and septalium absent; dental plates close to lat eral walls and not reach ing the ven tral valve floor. Family Pugnacidae Rzhonsnitskaya, 1956 Re marks: This ge nus is quite close to Pugnax in ex ter nal form and anat omy; it dif fers in hav ing nu merous costae (few in Pugnax) Ge nus Parapugnax Schmidt, 1964 and in its raduliform crura (later ally flattened and subparallel in Type spe cies: Pugnax pugnus brecciae Schmidt, 1941 Pugnax; Savage et al., 2002, p. 1164). Pugnax is in need of re vi - [1941b]; Langenaubach near Haiger, Lahn Syncline, Ger- sion, as tes ti fied by its improb a ble alleged stratigraphic range many; Iberg Limestone, Frasnian (Mid dle De vo nian to Up per Perm ian, Sav age et al., 2002; see also Sartenaer, 1968). Parapugnax? cf. skalensis (Biernat, 1966) Fig. 16A–J, P Paulinaerhynchia paulinae new genus and spe cies cf. 1966 ?Nemesa skalensis n. sp. – Biernat: pp. 100–101, pl. Figs 17, 18 22, figs 14–17. Type ma te rial: Holotype (a complete artic u lated shell) ZPAL Bp Ma te rial: Four spec i mens (two subcomplete) from Aferdou, 68/2/24/1 and twenty paratypes, incl. four complete ar tic u lated ZPAL Bp 68/1/23/1–4. shells; four paratypes seri ally sec tioned) from Maharch, ZPAL Bp De scrip tion: Shell ir reg u larly pen tag o nal (nearly tri an gu lar) in 68/2/24/2–21; three du bi ous frag men tary spec i mens from Afer- out line, trans verse (width-to-length ra tio ca. 1.3–1.5), strongly dou, ZPAL Bp 68/1/24/1–3. dorsibiconvex, up to ca. 20 mm in width. Max i mal width at 1/2 to Et y mol ogy: In hon our of Paulina Halamska, senior au thor’s spouse. 1 of the shell length. Ven tral valve moder ately con vex in umbonal Di ag no sis: As for the ge nus. re gion, flattened to slightly con cave on flanks, with a wide U-sha- De scrip tion: Shell subpentagonal in out line, trans verse with ped sulcus an te ri orly. Dor sal valve very deep (the thick ness of the width to length ratio 1.3–1.7 (N = 5), strongly dorsibiconvex, up to shell is roughly equal to its length), approx i mately semicir cu lar in 41.2 mm in width (di mensions given in Ta ble 2). Max i mal width poste rior view. Ven tral beak poorly pre served. An te rior commis- about mid-length or slightly an te ri orly. Dor sal valve strongly con - sure uniplicate, tongue broad, high, subtrapezoidal. Or na menta - vex (shell thick ness equal ling 0.8–1.3 its length), with a rounded, tion of high, strong, acute costae de vel oped in the ante rior half of rel a tively high fold and steep flanks. Ven tral valve weakly con vex the valves, sep a rated by V-shaped fur rows of approx i mately the in umbonal re gion, flat tened on flanks, with a U-shaped deeply ex - same width as the costae, 4 on fold, 3 on flanks. ca vated sulcus an te ri orly. Both umbones thick, ventral beak in cur- Re marks: Parapugnax skalensis (Biernat, 1966) from the up per ved. An te rior commissure uniplicate, tongue very broad, oc cu py - Eifelian of Ska³y in the Holy Cross Mountains has a reg ularly con - ing 0.67–0.86 of the shell width, and high, with subvertical lat eral vex dor sal valve (Biernat, 1966). P. brecciae from the Frasnian of slopes and semi cir cu lar top. Or na men ta tion of dis tinct, rounded 270 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 271

Ta ble 2 Atrypa triloba (Sowerby, 1840) “a hand some shell” (Sowerby, 1840, [717], pl. 56: 14) from the (Mid dle?) De vo nian of Plymouth Bio met ric char ac ter is tics of Paulinaerhynchia paulinae (possi bly pres ent also in the Boulonnais; Devos, 1962) is sim i lar in gen. et sp. nov.; measure ments in millimetres out line and about the same size as Paulinaerhynchia paulinae gen. et sp. nov., but its fold is higher and ap pears closer to the umbonal Spe- re gion; the costation is stron ger. It is not cer tain, whether Camaro- W L T w W/L T/L w/W Nc Nl Nc/W cimen toechia triloba sensu Schmidt (1941a) be conspecific with the 24/1 49.0 29.0 30.7 32.7 1.69 1.06 0.67 11 7 0.34 above-men tioned Eng lish taxon. All Mid dle De vonian Pugnacidae from Germany (Schmidt, 24/4 38.5 28.5 27.3 33.0 1.35 0.96 0.86 14 ? 0.42 1941a, b) are smaller and have fewer costae. Pugnax proboscidea 24/2 34.0 25.7 24.2 26.9 1.32 0.94 0.79 8 4? 0.30 Nalivkin, 1930 from the Up per De vonian of East Ferghana is simi - 24/3 28.0 28.7 32.0 28.0 1.32 0.80 0.74 8 6? 0.29 larly large-sized, but the costae are fewer in num ber (Nalivkin, 24/5 40.0 23.9 30.3 30.9 1.67 1.27 0.77 9 ? 0.29 1930, p. 89–90, 186–187, pl. 6: 14, 31); the in te rior is un known. Dumestre and Illing (1967) reported a few brachi opod taxa from W – width; L – length; T – thickness; w – width of the tongue; Nc – number a Givetian reef in the Uein Terguet re gion (near the town of of costae and costellae on the tongue; Nl – number of costae and costellae per a dor sal lat eral flank Semara, Western Sahara, Morocco; western part of the Tindouf Syncline) and among them a “large rhynchonellid super fi cially re - sem bling Hypothyridina or Uncinulus but which proved on sec - tion ing to have to tally dif fer ent in ter nal struc ture and to be near est costae, sep a rated by nar rower fur rows, 7–14 on tongue, (4–)6–8 to Ladogioides” (writ ten com muni ca tion of H.M. Muir-Wood in less strong ones on flanks, but or na menta tion only seen in the an te - Dumestre and Illing, 1967). One may wonder, if this does not re fer rior half of large shells. to the same genus as here. Ladogioides has simi larly ori ented den- Ven tral in te rior with very short, ru di men tary, ven trally di ver- tal plates, but pos sesses a dorsal sep tum and a septalium. Unfor tu - gent dental plates situ ated close to the lateral valve walls (result ing nately, this ma te rial could not be traced in the Natu ral History in nar row den tal cavi ties) and not reach ing the floor of the valve; Mu seum, in Lon don (S. Long, per sonal com mu ni ca tion, 6th Nov., teeth slen der and short. In the dor sal valve septalium and me dian 2012). sep tum absent but a high myophragm re vealed in one of the sec - Halamski (2004a) re ported a sin gle frag mentary shell of a sim i - tioned spec i mens; hinge plates divided; outer hinge plates flat, lar rhynchonellide from the Eifelian to Givetian bound ary beds at nearly hor i zon tal, with well marked crural bases forming sharp Sitka in the Holy Cross Mountains and re ferred to it as “Hypo- ridges along in ner mar gins of outer hinge plates; crura curv ing thyridinidae? gen. et sp. nov.”. ven trally, tri an gu lar in sec tion, raduliform. A low, tri an gu lar in Type lo cal ity: Guelb el Maharch. cross-sec tion me dian ridge de vel oped at some dis tance from the Type level: Guelb el Maharch mud mound, Taboumakhloõf For- hinge mar gin in a sin gle spec i men (Fig. 18D) can be in ter preted as mation; lower Givetian. a myophragm di vid ing mus cle at tach ment area. Strati graphic dis tri bu tion: lower Givetian. Re marks: The spec i mens shown in Fig. 17 il lustrate the vari abil - De po si tion of types: ZPAL. ity of the ex ter nal mor phol ogy, ex pressed mainly in the shell Dis tri bu tion: The new taxon remains uncon firmed, out side the thickness (compare the shell illus trated in Fig. 17A–E with the re- type local ity and stratum. main ing three), as well as the width and the shape of the bot tom of the ventral sulcus; the latter is usually slightly concave to (seldom) very weakly con vex. Rhynchonellida fam., gen. et sp. indet. Com par i sons of Paulinaerhynchia paulinae gen. et sp. nov. Fig. 30A–E with simi lar rhynchonellides are hindered, because in several ca- ses, the in te rior fea tures are un known. Homoeomorphic taxa do Ma te rial: Two ar tic u lated shells (one subcomplete, one frag men- ex ist, as ex em pli fied by rep re sen ta tives of Amissopecten Havlíèek, tary) from Aferdou, ZPAL Bp 68/1/50/1. 1960 (Camarotoechiidae; Havlíèek, 1961, Havlíèek and Kukal, De scrip tion: The sin gle mea surable shell is subpentagonal in out - 1990) or Ladogioides Mc Laren, 1961 (Yunnanellidae; Mc Laren, line, with greatest width at about two-thirds of the shell length, 1962). dorsibiconvex, ca. 10 mm wide, 9.4 mm long, and 6.5 mm thick. “Rhynchonella acuminata var. platiloba Sowerby” sensu Mau - Ante rior commissure uniplicate, ca. 8 mm broad, high, rounded. rer (1885, p. 207, pl. 8: 35) from the ap prox i mately co eval strata of Beak incurved. Shell mac ro scop i cally smooth. In te rior un known. Grube Hainau at Waldgirmes (mate rial exam ined: HLMD-Mr. Re marks: This brachi o pod is iden ti fied as a mem ber of the Rhyn- 10770, 10771) and Brilon (ma te rial ex amined: BGR, Koch’sche chonellida, on account of its astrophic shell, hidden interareas, and Sammlung, ZGI, no number) is very simi lar in shape to Paulina- absence of pedicle fora men. It is certainly differ ent from all the erhynchia paulinae gen. et sp. nov., but is smaller and lacks costae forms iden ti fied above, but the pos si bil ity of any iden ti fi ca tion is on lat eral flanks. The tongue is striate (4–5 per mm), but this fea - precluded by the scarcity of mate rial. It is illus trated here to call at- ture might have been lost in the ma te rial of the pres ent study. The ten tion to the presence of smooth rhynchonellides in the fauna in te rior is un known. stud ied.

Fig. 16. Middle De vo nian Rhynchonellida from the south ern MaÎder and Eu rope. A–J, P. Parapugnax? cf. skalensis (Biernat, 1966). All spec i mens from Aferdou. A–E. Artic ulated shell ZPAL Bp 68/1/23/1 in dor sal, ven tral, lat eral, pos te rior, and ante rior views. F–J. Ar - tic u lated shell ZPAL Bp 68/1/23/2 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. P. Frag mentary spec i men ZPAL Bp 68/1/23/3 in an te rior view. K–O, Q–OO. Septalaria gracilis (Gürich, 1896). Ar tic u lated shells in dor sal, ven tral, lat eral, pos te rior, and an te rior views. K–O. Spec i men ZPAL Bp 68/1/22/5 from Aferdou. Q–U. Spec i men ZPAL Bp 68/1/22/3 from Aferdou. V–Z. Spec i men SMF 29599.1 from Bilveringsen (Sauerland). AA–EE. Spec i men ZPAL Bp 68/1/22/2 from Aferdou. FF–JJ. Spec i men ZPAL Bp 68/1/22/1 from Aferdou. KK–OO. Speci men ZPAL Bp 68/1/22/4 from Aferdou 272 A. T. HALAMSKI & A. BALIÑSKI

Fig. 17. Paulinaerhynchia paulinae gen. et sp. nov. Ar tic ulated shells in dor sal, ven tral, lat eral, pos te rior, and an te rior views. A–E. Paratype ZPAL Bp 68/2/24/10 from Maharch. F–J. Paratype ZPAL Bp 68/2/24/3 from Maharch. K–O. Paratype ZPAL Bp 68/2/24/2 from Maharch. P–T. Holotype ZPAL Bp 68/2/24/1 from Maharch

Order Atrypida Rzhonsnitskaya, 1960 kib, SMF 94845–94847, 98179 and MB un numbered (V. Ebbig- Family Atrypidae Gill, 1871 hausen’s col lec tion). Ad di tional ma te rial: three ar tic ulated shells MNHN.F.A48101, Ge nus Atrypa Dalman, 1828 50414 from local ity 363o, Zemmour Noir, Mau ri ta nia, coll. J. Type spe cies: Anomia reticularis Linnaeus, 1758; Sougy. Gotland; lower Hemse Beds, Ludlow, Silu rian Other ma te rial ex am ined: Paratypes SMF 94878, col lected near Atrypa (Planatrypa?) confusa (Struve, 1992) Ahütte railway sta tion (St. 190; both this and the fol lowing lo cal - ity number re fer to an un pub lished reg ister of Struve’s col lect ing Figs 19G–JJ, 20 lo cal i ties kept at the SMF), and SMF 97879 from the quarry Mül- vp 1964 Atrypa reticularis – Sougy, p. 383. lertchen at Ahütte (Fp. 2162); both Hillesheim Syncline, Eifel; v* 1992 Planatrypa confusa n. sp. – Struve: pp. 552–554. Ahbach For ma tion, mid dle Eifelian. . 1995 Atrypa (Planatrypa) confusa (Struve, 1992) – Gode- De scrip tion: Shell heptagonal, rounded, or shield-shaped in out- froid, pp. 87–89, text-fig. 8,; pl. 2, figs 6–8. line, strongly dorsibiconvex to planoconvex, usually lon ger than v. 2001 Planatrypa confusa Struve 1992 – Thormann and wide, typi cally 21 to 26 mm wide, more seldom up to 36 mm wide. Weddige, pl. 2, fig. 15. Shoulder an gle 130°–150°(–160°). Dor sal valve reg u larly arched. Ma te rial: Over ninety spec i mens (mainly complete or subcom - Ven tral valve weakly to dis tinctly con vex in umbonal re gion, plete, ar tic ulated shells) from Aferdou, ZPAL Bp 68/1/48; about flanks weakly convex to concave, in the latter case redressed at sixty ar tic ulated shells from the Drotops beds at MadÀne el Mra- margins; beak suberect to incurved; fo ra men obscured by pedicle MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 273

Fig. 18. Trans verse se rial sec tions of Paulinaerhynchia paulinae gen. et sp. nov. through shells ZPAL Bp 68/2/24/6 (A), 68/2/24/7 (B), 68/2/24/8 (C), and 68/2/24/9 (D) from Maharch. Dis tances mea sured in milli metres from the tip of the ven tral umbo

callist. Ante rior commissure nearly straight to uniplicate; tongue orig i nal ma te rial of Struve (1992) (five spec i mens only) did not al - usually rather nar row and low. Or na menta tion of fine ribs, (5–) low the au thor to ac count for the vari abil ity of this spe cies, which 6–9 per 5 mm, inter rupted by fre quent growth lines. Frills un com- is illus trated more fully herein. A speci men from the Zemmour monly pre served, up to 5 mm long. Noir (Mau ri ta nia) is il lustrated (Fig. 19I–M) to doc ument the pres - Ventral inte rior with pedicle callist and pedicle collar (Fig. 20B: ence of A. confusa in that re gion. 1.4 and 1.65); teeth large, long, and strong, nearly ver ti cal, with Dis tri bu tion: Eifel, Ardennes, MaÎder, and Zemmour Noir; Mid- poorly de vel oped lateral lobes; den tal nu clei distinct (Fig. 20B: dle De vo nian. 1.2, 1.4, and 1.65). Dor sal valve with massive hinge plates; car di - nal pit deep with dis tinct at tach ments of diductor muscles (car di - Ge nus Atryparia Copper, 1966 nal pro cess) lin ing also in ner socket ridges poste ri orly; the me dian Type spe cies: Atryparia instita Copper, 1966; Mühlenberg ridge di vid ing adductor mus cle dis tinct and fairly long. near Niederehe, Hillesheim Syncline, Eifel; Ahbach Beds, Re marks: The dis cussed brachi opod was included within the ge - nus Planatrypa by Struve (1992) [sub ge nus Atrypa (Planatrypa) upper Eifelian (Copper, 1966b) Struve, 1966 herein]. However, it dif fers from Atrypa (Plana- Re marks: A poten tially mislead ing re port of “a spec i men, possi - trypa) in pos sess ing frills and dental nuclei; the lat ter charac ter bly re lated to Atryparia, from the De vo nian near the Al ge ria-Mo - distin guishes it also from Kyrtatrypa Struve, 1966, which has solid rocco bor der” (Cop per, 1966b, p. 983), therefore in close proxi- teeth (Cop per, 2002). It re sembles respresentatives of Atrypa mity to the area studied in the present paper, is corrected here: as a (Atrypa) Dalman, 1828, a sub ge nus to this time known up to the mat ter of fact, the brachi o pod orig i nated in the Mouydir Ba sin, Emsian (Copper, 2002), in hav ing convexoplane to dorsibiconvex some 800 km further south-east (Flamand, 1911). shell, uniplicate commissure, obscured fo ra men, and mul ti ple Atryparia dispersa (Struve, 1966) frills (di ag nostic char ac ters af ter Cop per, 2002). The frills be ing an un cer tain di ag nos tic char ac ter, be cause of preservational bias Figs 21F–O, 22, 28L (P. Cop per, per sonal com mu ni ca tion April 2013), the sub ge ner ic * 1966 Atrypa (Hyponeatrypa) dispersa n. sp. – Struve: pp. ap pur te nance of the brachi o pod dis cussed is con sid ered here as 142–143, text-fig. 11, pl. 16, fig. 10. un re solved; it is re ported un der the con ser va tive head ing “Atrypa non 1970 Atryparia dispersa (Struve, 1966) – Godefroid, pp. (Planatrypa?) confusa“. It is worth not ing that Cop per (1967c) re - 97–98, text-fig. 5, pl. 1, fig. 2, pl. 2, figs 1–2 [A. ported the presence of de flected growth lamellae and minute den - instita; fide Godefroid, 1995]. tal nuclei in some species of Atrypa (Planatrypa) from the Eifelian 1987 Atryparia (Atryparia) dispersa (Struve, 1966) – Ko- of Germany. marov, pp. 106–109, text-fig. 33, pl. 2, figs 4, 5. Re in ves ti ga tion of the type ma te rial of “Planatrypa confusa . 1995 Atryparia dispersa (Struve 1966) – Struve, p. 100, Struve, 1992” from the Eifel per mitted the discov ery of frills (Fig. figs 33–34. 19Z) and a plate cov er ing the delthyrium, char ac ters ex clud ing as - 1995 Atryparia sp., aff. dispersa (Struve 1966) – Struve, p. sign ment to Atrypa (Planatrypa) and con cor dant with the pres ent 100, figs 35–36. ma te rial. These in ter nal char ac ters are also ev i dent in the ma te rial non 2007 Atryparia dispersa (Struve 1966) – Hubert et al., p. from the Ardennes (Godefroid, 1995, fig. 8, espe cially 0.4). The 260 [A. instita]. 274 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 275

Fig. 19. Middle De vo nian Atrypida from south ern MaÎder and Eu rope. A–F. Desquamatia (D.) microzonata Struve, 1966. All spec i - mens from MadÀne el Mrakib. A–E. Ar tic u lated shell SMF 94842 in dorsal, ven tral, lat eral, pos te rior, and ante rior views. F. Ven tral in te - rior SMF 94840. G–JJ. Atrypa (Planatrypa?) confusa Struve, 1992. G, H. Artic ulated shell ZPAL Bp 68/1/48/1 from Aferdou in dor sal and ven tral views. I–M. Artic ulated shell MNHN.F.A50414 from local ity 346o sensu Sougy (1964) in the Zemmour Noir (Mauri ta nia). N–R. Artic ulated shell ZPAL Bp 68/1/48/2 from Aferdou. S–W. Ar tic ulated shell ZPAL Bp 68/1/48/3 from Aferdou. X. Ar tic ulated shell ZPAL Bp 68/1/48/4 from Aferdou in ven tral view, showing frills. Y–DD. Ar tic u lated shell SMF 94879 from the Ahbach beds (upper Eifelian) of the quarry “Müllertchen”, 470 m S from the Ahütte rail way sta tion, Eifel (Fp. 2162 sensu Struve un pub lished) in dor sal, ven - tral, lat eral, pos te rior, and an te rior views. EE, FF, II. Ven tral valve SMF 94847 shown in exter nal (FF) and in ter nal (II) views; en large - ment of the umbo (EE). GG. Ar tic ulated shell ZPAL Bp 68/1/48/7 from Aferdou, show ing traces of the dor sal in te rior. HH, JJ. Two ven tral in te ri ors SMF 94846, 94845 from MadÀne el Mrakib

Ma te rial: Over one hun dred and twenty ar tic ulated shells (sev eral Ventral inte rior with thick, well devel oped pedicle callist, on the of them complete and well pre served) from Aferdou, ZPAL Bp sec tioned spec i men funnelled into col lar; teeth strong, with lat eral 68/1/28/1. lobes and small dental nuclei. Dorsal valve with a large cardi nal De scrip tion: Shell usu ally over 30 mm in width, up to ca. 38 mm, pit, mod er ately strong hinge plates and widely tri an gular in trans- lon ger than wide, sel dom wider than long. Ven tral valve subcir- verse sec tion myophragm; sock ets with large in ner socket ridges cular to shield-shaped in out line with great est width, re spec tively, and a dis tinct me dian ridge. about midlength or pos te ri orly (up to pos te rior fifth of the valve), Re marks: In the ma te rial from the north ern MaÎder (7 speci mens), moder ately con vex in umbonal re gion, some what flat tened on Struve (1995) sep a rated A. dispersa and A. aff. dispersa, on the ba - flanks; beak poorly pre served, pos sibly incurved. Dor sal valve sis of the greatest width of the shell, ei ther at mid-length or more con vex to strongly con vex. An te rior commissure nearly straight to poste ri orly. In the much more abun dant ma te rial, stud ied by the strongly uniplicate, usu ally with a dis tinct but poorly delimitated pres ent au thors, these morphotypes intergrade, where fore they are plica. con sid ered conspecific. Ornamentation of strong, flat tened ribs, some of them bifur cat - Dis tri bu tion: In the type area, this spe cies is known from the ing up to three times on the ven tral valve, more sel dom on the dor- Junkerberg to Ahbach for mations (middle to up per Eifelian; sal one, (2.5–)3–4 per 5 mm at an te rior mar gin, in ter rupted by Struve, 1966). This spe cies was also re ported from Transcaucasia strong growth lines. Frills (Fig. 28L) pre served up to 9 mm in (Komarov, 1997) and er ro ne ously from the Ardennes (Godefroid, length, with ribs ei ther simi lar to those on the shell or much finer 1970; the er ror was cor rected by Godefroid, 1995, but was re - (ca. 10 per 5 mm). peated by Hubert et al., 2007).

Fig. 20. Trans verse se rial sec tions of Atrypa (Planatrypa?) confusa through shells ZPAL Bp ZPAL Bp 68/1/48/5 (A) and 68/1/48/6 (B) from Aferdou. Distances mea sured in milli metres from the tip of the ven tral umbo 276 A. T. HALAMSKI & A. BALIÑSKI

Fig. 21. Middle De vo nian Atrypida from south ern MaÎder. A–E. Gruenewaldtia latilinguis (Schnur, 1851). Spec i men ZPAL Bp 68/1/30/1. F–O. Atryparia dispersa (Struve, 1966). F–J. Spec i men 68/1/28/1. K–O. Spec i men 68/1/28/2. All speci mens are ar tic ulated shells from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views

Fig. 22. Trans verse se rial sec tions of Atryparia dispersa (Struve, 1966) through shell ZPAL Bp 68/1/28/3 from Aferdou. Dis tances mea - sured in millimetres from the tip of the ven tral umbo MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 277

Ge nus Invertina Copper and Chen, 1995 inter preted to reflect infraspecific and/or ontogenetic variabil ity. Type spe cies: Atrypa aspera var. sinensis Kayser, 1883; Kerpina atrypoides Struve, 1961 is a poorly known taxon from the Yunnan; Givetian? same strata of the Eifel (Struve, 1961), of simi larly smaller size than the ma te rial described here. This is the first report of Kerpina Invertina cf. struvei Godefroid, 2000 from Africa. Fig. 23A–E Dis tri bu tion: Kerpina vineta was de scribed orig i nally from the late Eifelian of the Eifel area in Germany, where it is confined to cf. 2000 Invertina struvei n. sp. – Godefroid: pp. 269–271, the “Rasenriff” or thicket reef biotope (Cop per, 1967b). text-figs 2–6, pl. 1, figs 1–8. Ma te rial: A single spec i men from Aferdou, ZPAL Bp 68/1/45/1. De scrip tion: The single available shell is ellip tic in outline, elon- Ge nus Spinatrypa Stainbrook, 1951 gate, venribiconvex, 10.3 mm wide, 11.8 mm long, and 4.9 mm Type spe cies: “Atrypa hystrix var. occidentalis Hall” thick. Dor sal valve weakly con vex in umbonal re gion, with a me - (errore pro Atrypa aspera var. occidentalis Hall, 1858) dian sulcus in the poste rior fourth, nearly flat in lat eral and ante rior sensu Stainbrook, 1945; Iowa, USA; Cedar Valley Forma - re gions. Ven tral valve tri an gu lar in an te rior view; umbo very fine, tion, Frasnian strongly incurved. An te rior commissure nearly straight. Or na men- ta tion of rounded ribs, 6–7 per 5 mm. Spinatrypa (Spinatrypa) cf. trigonella (Davidson, 1882) Re marks: The speci men described is of small size and nearly Fig. 25K–HH planoconvex; it is prob a bly a ju ve nile, like that of Invertina sinen- cf. 1882 Atrypa? trigonella, Dav. – Davidson, p. 40, pl. 1, figs sis il lus trated by Cop per and Chen (1995, fig. 2: 11–15) from the 19, 19a, 19b. middle Givetian of Sichuan. The com par i son with the slightly vp 1964 Atrypa aspera (Schlotheim, 1813) – Sougy, pp. 445– youn ger (Up per varcus Zone) Invertina struvei from the north ern 446, pl. 50, fig. 7. MaÎder (Godefroid, 2000) is there fore dif fi cult; open no mencla - cf. 1965 Spinatrypa trigonella (Davidson) – Cop per, p. 360. ture has been used for this rea son. As for other Mid dle De vo nian ventribiconvex atrypides, Kerpina is usu ally asym metric (Struve, Ma te rial: Forty-eight artic ulated shells, several of them complete 1961; Cop per, 1967b) and Invertina fasciplicata is nearly aequibi- and well pre served, from Aferdou, ZPAL Bp 68/1/29/1–48. con vex (Cop per, 1967a). Ad di tional ma te rial: Artic u lated shell MNHN.F.R10288 from OudeÎ Askaf (Zemmour, Mauri ta nia) fig ured by Sougy (1964, pl. 50: 7); fourteen mostly complete artic ulated shells MNHN.F.A Ge nus Kerpina Struve, 1961 48102, 50415 from the lo cal ity 2108A, det. P. Cop per; ar tic ulated Type spe cies: Kerpina vineta Struve, 1961; Eifel, Eifelian shell MNHN.F.A48104 from the lo cal ity 2428; three ar tic ulated shells MNHN.F.A48103 from the local ity 346c; all the above from Kerpina vineta Struve, 1961 the Zemmour Noir, Mau ri ta nia, coll. J. Sougy. Figs 23F–T, 24 De scrip tion: Shell ap proxi mately as wide as long, more rarely v* 1961 Kerpina vineta n. g., n. sp. – Struve: p. 333, pl. 1, figs trans verse, ventribiconvex, usually about 17 to 23 mm in width, 3, 4. sel dom up to ca. 28 mm. Orna menta tion of un du lat ing ribs, 14–16 (–22) in to tal, 3(–4.5) per 5 mm; tubu lar spines pres ent. Ma te rial: Seventy-seven spec i mens from Aferdou El-Mrakib, Re marks: This spe cies is included into Spinatrypa, on ac count of some of them well pre served, a large part slightly de formed, some wave-like ribs and the pres ence of spines; it is re ferred to the in ad - in complete, ZPAL Bp 68/1/27/1–77. e quately known species Spinatrypa trigonella (Davidson, 1882), De scrip tion: Shell circu lar to ellip tic in outline, slightly longer de scribed from the Mid dle De vonian (Givetian; Elliott, 1961 as than wide to slightly wider than long, moder ately ventribiconvex “Upper Givetian”, but prob a bly lower if the Bilveringsen fauna is to moder ately dorsibiconvex, up to 28.0 mm in width. Dorsal an equiv a lent; Cop per, 1965) of Lummaton near Torquay (Devon- valve uni formly con vex. Ven tral valve moder ately in flated or with shire; Davidson, 1882) on ac count of simi lar shape and or na men- flat tened attach ment area in umbonal region, some times slightly tation. The type mate rial has not been restud ied recently, where- geniculated in the lateral profile, with a steep marginal region; fore open no mencla ture is used as a pre cau tion. Spinatrypa trigo- beak straight to moder ately incurved, with an api cal fo ra men; ven - nella prob a bly oc curs also in the Rhenish Massif (Büchel in the tral interarea very high, trans versely striate, weakly curved to Bergisches Land; Copper, 1965, p. 360). Spinatrypa cf. trigonella nearly flat, usually apsacline, seldom weakly anacline. Ante rior from Aferdou resem bles early Eifelian Spinatrypa variaspina commissure uniplicate, tongue oc cu py ing about two-thirds of the Copper, 1967 [1967a] from the Eifel, which has, how ever, finer shell width. or na menta tion. Two other spe cies from the Eifel show some more Orna menta tion of strong costae and costellae, 6–8 per 5 mm, distant resem blance to the ma te rial studied: the early Givetian crossing with strong growth lines. Spinatrypa curvirostra Copper, 1967 [1967a] has coarser or na - In ter nally, shell wall thick, es pe cially pos te ri orly; thick pedicle menta tion, while the middle Givetian Spinatypa orthoclina Cop- callist and massively thick ened deltidial plates form wide, irreg u - per, 1967 [1967a] is flat ter (Cop per, 1967a). lar pedicle collar (Fig. 24); teeth solid, without dental nuclei, thick. Dis tri bu tion: This spe cies is known from the Mid dle De vonian of Dorsal in te rior with low and thick hinge plates; me dian myo- MaÎder and of Zemmour Noir (Mau ri ta nia); most prob a bly also of phragm wide, well de vel oped. Devonshire and the Rhenish Mas sif. Re marks: This spe cies is eas ily distin guished from co-oc cur ring atrypides by its excep tion ally high ventral interarea, asymme try, and a steep mar ginal re gion of the ven tral valve. The type col lec - Spinatrypa (Spinatrypa) globulina Copper, 1967 [1967a] tion of Kerpina vineta Struve, 1961 from the late Eifelian Freilin- Fig. 25A–J gen Forma tion of the Eifel con tains in di vid uals that are smaller (usually ca. 15 mm wide, max i mum width 20.2 mm; Copper, * 1967a Spinatrypa globulina n. sp. – Cop per: pp. 506–507, 1967b, and the present au thors), have approx i mately the same or - pl. 78, figs 11–24. na menta tion den sity and flat to weakly con vex dor sal valve. The Ma te rial: Two ar tic u lated shells (one subcomplete, one in com- dif fer ences of size and con vex ity, al though clear and con stant, are plete) from Aferdou, ZPAL Bp 68/1/47/1–2. 278 A. T. HALAMSKI & A. BALIÑSKI

Fig. 23. Mid dle De vo nian Atrypida from south ern MaÎder. A–E. Invertina cf. struvei Godefroid, 2000. Artic u lated shell ZPAL Bp 68/1/45/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–T. Kerpina vineta Struve, 1961. Ar tic u lated shells from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–J. Spec i men ZPAL Bp 68/1/27/1. K–O. Spec i men ZPAL Bp 68/1/27/2. P–T. Spec i men ZPAL Bp 68/1/27/3

De scrip tion: Shell approx i mately cir cular in outline, weakly to Ge nus Desquamatia Alekseeva, 1960 moder ately dorsibiconvex, up to ca. 20 mm in width. An te rior Sub ge nus Desquamatia (Desquamatia) Alekseeva, 1960 commissure uniplicate, tongue broad and low. Orna menta tion of Type spe cies: Atrypa (Desquamatia) khavae Alekseeva, rounded, undulose, broad ribs, 1–2 per 5 mm at an te rior mar gin. 1960; Urals, Rus sia; lower Eifelian The ven tral midrib pair is stron ger than the other ribs. Re marks: This spe cies is dis tin guished from the co-oc cur ring Desquamatia (Desquamatia) microzonata Struve, 1966 Spinatrypa cf. trigonella by its coarser ribs and the presence of a Fig. 19A–F stron ger ven tral midrib pair. Dis tri bu tion: In the type area, S. globulina is found in the Loogh vp 1964 Atrypa reticularis – Sougy, p. 448. For ma tion (low er most Givetian, Eifel, Ger many; Cop per, 1967a). v* 1966 Desquamatia (Synatrypa) microzonata n. sp. – Struve: pp. 151–153, pl. 16, fig. 12. 1970 Desquamatia (Synatrypa) microzonata Struve, 1966 MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 279

– Godefroid, p. 117–119, fig. 21, pl. 1, fig. 3. 1995 Desquamatia (Synatrypa) microzonata Struve, 1966 – Godefroid, p. 94, pl. 3, fig. 5. Ma te rial: Five ar tic ulated shells SMF 94839–94842 and MB un- numbered (V. Ebbighausen’s col lec tion) from the Drotops beds at MadÀne el Mrakib. Ad di tional ma te rial: Three ar tic ulated shells MNHN.F.A48116 from the “basal Givetian” of Aguelt Oudiate el Khyam (local ity 401), Zemmour Noir, Mau ri ta nia, coll. J. Sougy, det. P. Cop per [as “Desquamatia (Synatrypa) cf. microzonata”]. De scrip tion: Shell subcircular in out line, up to 16.9 mm wide, mod er ately to mark edly dorsibiconvex. Or na men ta tion of ribs, 8–9 per 5 mm at an te rior mar gin. In te rior not stud ied. Re marks: As far as the exte ri ors are concerned, the mate rial stud- ied is indis tin guish able from the type col lec tion from the Eifel. Dis tri bu tion: Desquamatia microzonata is known from the mid- dle Eifelian of the Eifel (Struve, 1966) and of the Ardennes (Gode- froid, 1970, 1995), as well as from the Mid dle De vo nian of MaÎder and of the Zemmour Noir. Desquamatia cf. microzonata was re- ported from the Mid dle De vo nian of Padaupkin in Burma (Ander - Fig. 24. Trans verse se rial sec tions of Kerpina vineta Struve, son et al., 1969). The subge nus Desquamatia (Synatrypa) is a 1961 through shell ZPAL Bp 68/1/27/4 (A) from Aferdou. Dis- syn onym of D. (Desquamatia) (Cop per, 2002). tances measured in millimetres from the tip of the ven tral umbo

Desquamatia (Desquamatia) deserti new species 26A–E) is ten ta tively re ferred to the discussed spe cies. Figs 26A–O, 27 Type lo cal ity: Guelb el Maharch. Type ma te rial: Holotype, complete artic u lated shell ZPAL Bp Type level: Guelb el Maharch mud mound, Taboumakhloõf For- 68/2/26/1 and thirty nine paratypes, ZPAL Bp 68/2/26/2–40, mation; lower Givetian. (among them some complete artic u lated shells), all from Maharch. Strati graphic dis tri bu tion: lower Givetian. Et y mol ogy: Desertum, Latin for desert. De po si tion of types: ZPAL. Di men sions: See Table 3. Dis tri bu tion: Ex cept for the single du bi ous spec i men from Afer- Di ag no sis: Desquamatia (Desquamatia) with nearly flat ventral dou, the new taxon is unknown outside the type lo cal ity and stra tum. valve and fine or na menta tion, 7–9(–13) ribs per 5 mm at an te rior mar gin. Sub ge nus Desquamatia (Independatrypa) Copper, 1973 De scrip tion: Shell more fre quently lon ger than wide, usually over Type spe cies: Atrypa independensis Web ster, 1921, Iowa, 35 mm in width, up to ca. 40 mm. Shoul der an gle ca. 140–150°. USA; middle Givetian Dorsal valve high. Ven tral valve subtriangular in an te rior view, con vex in umbonal re gion, oth er wise flat tened. An te rior commi- Desquamatia (Independatrypa) circulareformis Biernat, ssure broadly uniplicate. Or na menta tion of fine ribs, 7–9(–13) per 1964 5 mm at ante rior mar gin. Fig. 26U–DD Ventral valve with pedicle callist with tendency to de velop into a short col lar; teeth long, strong with a lat eral lobe and small den - v* 1966 Desquamatia circulareformis n. sp. – Biernat: pp. tal cavi ties. Dorsal valve with a rather small cardi nal pit and a tri- 326–327, pl. 8, fig. 8, pl. 12, figs 2–3. an gu lar myophragm; hinge plates with comblike car di nal pro cess Ma te rial: Four speci mens (three subcomplete artciulated shells) de vel oped pos te ri orly; crural bases well vis i ble, ex pand ing into from Aferdou, ZPAL Bp 68/1/25/1–4. sharply de flected later ally and feathered crura (Fig. 27). De scrip tion: Shell weakly trans verse (width to length ra tio 1.13– Re marks: Desquamatia (D.) deserti sp. nov. may be com pared to 1.17), strongly dorsibiconvex, up to 43 mm in width. Dorsal valve some fine-ribbed and large-sized atrypides, al though none of them moder ately con vex (thick ness to width ra tio 0.53–0.58). Ven tral is closely simi lar. Desquamatia (D.) ovata Copper, 1966 from the valve rounded, mod er ately con vex, tri an gu lar in an te rior view; beak Eifelian of the Eifel (Nohn Forma tion) is quite sim i lar in outline elon gate, incurved. An te rior commissure uniplicate, tongue subtra- (Cop per, 1966a); it dif fers in its more con vex ven tral valve. The pezoidal to rounded, wide, moder ately high. Or na menta tion of ribs, finely ribbed Atrypa desquamata var. alticola Frech, 1891 from the Givetian of the Carnic Alps (Frech, 1891) prob a bly does not rep re sent the ge nus Desquamatia (see Baliñski, 1979, 1998). The Ta ble 3 new spe cies shows some exter nal simi lar ity to the finely ribbed Desquamatia (D.) alticoliformis Rzhonsnitskaya, 1975, re ported Bio met ric char ac ter is tics of Desquamatia (D.) deserti sp. from the late Frasnian of Russia (Rzhonsnitskaya, 1975; Rzhon- nov.; mea sure ments in millimetres snitskaya et al., 1998), Eifel (Godefroid and Hauser, 2003), Arde- nnes (Godefroid and Helsen, 1998; Mottequin, 2008a), and Po land Specimen W L T R-10 Ra (Racki and Baliñski, 1998). Besides hav ing a dif fer ent stratigra- 2/26/1 35.1 36.6 21.6 8-10 8–9 phic range, the spe cies from Aferdou dif fers also in hav ing a not as 2/26/2 37.5 36.3 24.2 15 11–13 wide, rather shield-shaped out line of the shell, with a nar rowly elon gated an te rior re gion. The late Givetian Desquamatia (D.?) 2/26/5 36.1 42.1 25.5 11-12 9 globosa (Gürich, 1896) has a more con vex ven tral valve (Racki W – width; L – length; T – thickness; R-10 – num ber of ribs per 5 mm and Baliñski, 1981). counted at 10 mm from the beak; Ra – numer of ribs per 5 mm at ante rior The sin gle spec i men ZPAL Bp 68/1/26/1 from Aferdou (Fig. mar gin 280 A. T. HALAMSKI & A. BALIÑSKI

Fig. 25. Mid dle De vo nian Atrypida from north west ern Af rica. A–J. Spinatrypa globulina Cop per, 1967. In com plete ar tic u lated shells from Aferdou in dorsal, ven tral, lat eral, poste rior, and an te rior views. A–E. Spec i men ZPAL Bp 68/1/47/1. F–J. Spec i men ZPAL Bp 68/1/47/2. K–HH. Spinatrypa cf. trigonella (Davidson, 1882). K, L. Ar tic u lated shell ZPAL Bp 68/1/29/1 from Aferdou in dor sal and ven tral views. M–Q. Artic ulated shell MNHN.F.A50415 from the local ity 2108A sensu Sougy (1964) (Zemmour Noir, Mau ri ta nia) in dorsal, ven tral, lat eral, pos te rior, and ante rior views. W, X. Ar tic ulated shell ZPAL Bp 68/1/29/2 from Aferdou in dor sal and ven tral views. Y–CC. Ar tic u lated shell ZPAL Bp 68/1/29/3 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. DD–HH. Ar tic - u lated shell ZPAL Bp 68/1/29/4 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 281

4–6 per 5 mm at an te rior mar gin; growth lines sparse, lamellose. Family Carinatinidae Rzhonsnitskaya, 1960 Re marks: The above-men tioned ma te rial is con sidered conspecific with D. circulareformis, a poorly known, ap prox i mately co eval spe - Ge nus Carinatina Nalivkin, 1930 cies from Mi³oszów, in the Holy Cross Mountains (Biernat, 1964). Type spe cies: Orthis arimaspus Eichwald in von Buch, Three poorly pre served very large shells from Aferdou (ZPAL 1840; Bogoslowsk, Russia; Middle Devo nian Bp 68/1/25B/1–3) are simi lar to D. (I.) circulareformis, as above, but dif fer ing in their larger size (up to 51 mm), nearly flat dor sal Carinatina arimaspus (Eichwald in von Buch, 1840) valves and very high tongues, with subparallel lat eral slopes; they Fig. 28F–K are tenta tively re ferred to as D. (I.) cf. circulareformis. * 1840 Orthis Arimaspus Eichw. – von Buch: pp. 108, 112. Dis tri bu tion: Holy Cross Mountains and MaÎder, upper Eifelian 1978 C[arinatina] arimaspus (Eichwald, in Buch 1840) – to lower–middle Givetian. Copper, p. 310, pl. 7, figs 1–6. Ma te rial: Fourteen spec i mens (three subcomplete) from Aferdou, Desquamatia? sp. 1 ZPAL Bp 68/1/31/1–14. Fig. 26P–T De scrip tion: Shell trans verse (width-to-length ra tio 1.3–1.4), up Ma te rial: Four speci mens (two complete artic u lated shells) from to ca. 28 mm in width, very low (up to 7 mm thick). Poste rior mar - Aferdou, ZPAL Bp 68/1/25A/1. gin straight. Dorsal valve with a deep U-shaped me dian sulcus. De scrip tion: Shell ap proxi mately as wide as long (width-to- Ventral valve with a median carina, lateral flanks poste ri orly flat, length ra tio 0.99–1.02), markedly dorsibiconvex, up to 41 mm in an te ri orly con vex; interarea apsacline. An te rior commissure recti- width. Shoul der line composed of two con cave seg ments. Dorsal marginate. Or na menta tion (Fig. 28K) of high, acute, fre quently bi - valve mark edly con vex (thick ness to width ra tio 0.65–0.69). Ven - fur cat ing ribs, 3–7 per 5 mm at an te rior mar gin (higher or lower tral valve shield-shaped, con vex in umbonal re gion, lateral flanks numbers de pend on pres ence or ab sence of bi fur ca tion in close flattened to concave; beak small. Ante rior commissure uniplicate, prox imity to the an te rior mar gin, re spec tively). Growth lines reg u - tongue subtriangular, wide, high. Or na menta tion of ribs, 4–5(–6) lar, dense. per 5 mm at an te rior mar gin; growth lines dense, lamellose. Dis tri bu tion: Bo he mia, Lower De vo nian (Suchomasty Lime - Re marks: The large size of this rare form would support ap purte - stone, Emsian; Havlíèek and Kukal, 1990); Holy Cross Mountains nance to Desquamatia (Independatrypa), yet the dense orna men ta - and MaÎder, Mid dle De vo nian. tion in di cates D. (Desquamatia). It is diffi cult to find any compa- rable spe cies. Family Paraferellidae Spriesterbach, 1942 Ge nus Gruenewaldtia Tschernyschew, 1855 Family Davidsoniidae King, 1850 Type spe cies: Terebratula latilinguis Schnur, 1851; Ge nus Prodavidsonia Havlíèek, 1956 Ger many; Eifelian Type spe cies: Prodavidsonia dalejensis Havlíèek, 1956; Gruenewaldtia latilinguis (Schnur, 1851) Bo he mia; Choteè For ma tion, Eifelian Figs 21A–E, 29 Prodavidsonia sp. * 1851 Terebratula latilinguis, n. sp. – Schnur: p. 7. Fig. 28A–E 1955 Grünewaldtia latilinguis latilinguis (Schnur 1851) – Ma te rial: A single subcomplete spec i men from Aferdou El- Struve, pp. 228–229, text-fig. 8a. Mrakib, ZPAL Bp 68/1/32/1. v. 1964 Gruenewaldtia latilinguis latilinguis (Schnur, 1851) De scrip tion: Shell 12.6 mm wide, >8.3 mm long, and 1.7 mm – Biernat, pp. 332–334, text-fig. 20, pl. 14, figs 1–6, thick, its out line be ing best de scribed as an ellipse cut by the 8–9. straight line of the pos te rior margin. Dorsal valve flat, with a very 1965 Gruenewaldtia latilinguis (Schnur) – Cop per, pp. nar row (ca. 1 mm) median sulcus be gin ning <1.5 mm from the 369–371, text-fig. 8, pl. 47, figs 1–3. umbo and con tin uing to the ante rior mar gin. Ven tral valve very 1970 Gruenewaldtia latilinguis (Schnur, 1851) – Gode- low, but acutely tri angu lar in ante rior view, with barely per cepti - froid, pp. 123–125, text-fig. 23. ble low median carina; interarea very broad, nearly flat, catacline. Ma te rial: Five ar tic ulated shells from Aferdou, ZPAL Bp 68/1/ Ante rior commissure rectimarginate. Shell smooth ex cept for fine 30/1–5. growth lines. In te rior un known. De scrip tion: Shell rounded, slightly wider than long or longer Re marks: This speci men can not be matched with any repre sen ta - than wide, weakly ventribiconvex to mod er ately dorsibiconvex, tive of Prodavidsonia (incl. Quasidavidsonia Havlíèek, 1987, fol - ca. 30 to 40 mm in width. Dor sal valve slightly flat tened me di ally. low ing Cop per, 2002), de scribed up to now. P. tenuissima (Bar- Ven tral valve reg u larly con vex; tongue present but sulcus im per - rande, 1879) and P. mediocarinata Havlíèek, 1967, both from the cep ti ble; umbo very massive, beak incurved; deltidial plates fused. Eifelian of Bohe mia, have simi lar width-to-length ratios, but their Ante rior commissure unisulcate, tongue very broad, rounded. Or- ventral valves are concave ante ri orly (Barrande, 1879; Havlíèek, na menta tion of un in ter rupted sub acute ribs, 5–8 per 5 mm at an te - 1967). P. dalejensis Havlíèek, 1956 from the Eifelian of Bo he mia rior mar gin. has a simi lar weakly con vex ven tral valve and flat dor sal one, but Ven tral in te rior with mas sive mus cle plat form sup ported an te ri - its shape is more trans verse (Havlíèek, 1956). In P. vicina Hav- orly by two thick and coarse crystal line septa and ex tend ing an te ri - líèek, 1967 from the Eifelian of Moravia the ca rina and the sulcus orly for about one fourth of the valve length; pedicle collar not are less dis tinct (Havlíèek, 1967). The same can be said about the ob served in the sin gle sec tioned spec i men, but dis tinct in ter nal Givetian P. havliceki García-Al calde, 2010 from the Asturias, pro jec tions of deltidial plates are pres ent (Fig. 29, distances 3.0 which is, moreover, smaller. The late Eifelian to early Givetian P. and 3.2 mm). In the dor sal valve the in ner socket ridges thick; dis - scalensis (Sobolew, 1904) from the Holy Cross Mountains has a tinc tive stri ated car di nal pro cess lin ing pos te ri orly in ner socket con cave dor sal valve (Halamski, 2004a). ?Prodavidsonia sp., re- ridges and car di nal pit; mus cle plat form thick, well devel oped and ported by Drot (1961a) from the up per Givetian of Drâa val ley in rised somewhat an te ri orly from the umbo, supported by two coarse Morocco, is also concavo-con vex. crys tal line lat eral septa and a poorly dif fer en ti ated me dian sep tum 282 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 283

Fig. 26. Middle De vonian Atrypida from south ern MaÎder. A–O. Desquamatia (D.) deserti sp. nov. A–E. Spec i men ZPAL Bp 68/1/26/1 from Aferdou ten ta tively re ferred to the spe cies. F–J. Holotype ZPAL Bp 68/2/26/1 from Maharch. K–O. Paratype ZPAL Bp 68/2/26/2 from Maharch. P–T. Desquamatia? sp. 1 from Aferdou. Ar tic u lated shell ZPAL Bp 68/1/25A/1 in dor sal, ven tral, lat eral, an te rior, and pos te - rior views. U–DD. Desquamatia (Independatrypa) circulareformis Biernat, 1964 from Aferdou. U–Y. Spec i men ZPAL Bp 68/1/25/1. Z–DD. Spec i men ZPAL Bp 68/1/25/2. All pho to graphs are of ar tic u lated shells in dor sal, ven tral, lat eral, pos te rior, and an te rior views

(Fig. 29, dis tances 12.5 and 13.1 mm); thick me dian ridge with mens (usually complete artic u lated shells) from Maharch, ZPAL rounded top, an te ri orly medi ally grooved, ap pears at about 8 mm Bp 68/2/40. an te ri orly from the umbo, running on the up per sur face of the mus- Ad di tional ma te rial: Five partly decorticated artic u lated shells cle platform. MNHN.F.A48141 from a small es carp ment 2 km ENE from Re marks: Differ ences in the inter nal structures of the Gruene- Aguelt Oudiate el Khyam (sec tion D-13, lev els 5–6); “up per Cou- waldtia taxa, occur ring in the Eifel area, are best inter preted as vinian” (bed 80), Zemmour Noir, Mau ri ta nia, coll. J. Sougy. rep re sent ing spe cific dis tinc tions, whereas ex ter nal mor phol ogy is De scrip tion: Shell cir cu lar to tear drop-shaped in out line, aequibi- a poor tax onomic char ac ter (Cop per, 1965 and P. Cop per, pers. con vex to moder ately ventribiconvex, up to ca. 15 mm in width. comm., April 2013). The mate rial described here is sim i lar to G. Ven tral umbo elon gate, fine, beak strongly incurved. An te rior latilinguis (Struve, 1955, fig. 8a) in hav ing the ven tral plat form sup- commissure straight to very weakly uniplicate. Growth lines usu- ported by two lat eral septa with out a me dian sep tum, in con trast to ally well vis i ble, but rare, be coming conspicouous and crowded in G. matutina Struve, 1955 with sev eral ven tral septa (Struve, 1955, mar ginal re gion. fig. 8b–i). In ter nally, thick shelled, es pe cially pos te ri orly; den tal plates Dis tri bu tion: Eifel (Eifelian), Ardennes (Couvin, Co2c), Holy thin, subparallel to slightly diver gent ventrally; den tal cav i ties Cross Mountains (Ska³y beds), Eng land (Chercombe Bridge sha- large and wide; small deltidial plates de vel oped. In the dor sal les), Nakhichevan Re public (Djahannamdarasi val ley, up per Eife- valve car di nal pit wide and shal low; hinge plates low, in ner socket lian), MaÎder; Mid dle De vonian (Struve, 1955; Biernat, 1964; ridges nearly hor i zon tal, wide. Copper, 1965; Godefroid, 1970; Mamedov, 1974). Dis tri bu tion: This species was described from upper Eifelian Freilingen For mation of the Eifel (Cop per, 1986). This is the first Family Lissatrypidae Twenhofel, 1914 re port of this ge nus in Africa (MaÎder and Zemmour Noir). Ge nus Peratos Copper, 1986 Order Athyridida Boucot, Johnson and Staton, 1964 Type spe cies: Peratos arrectus Copper, 1986; Germany; Family Athyrididae Davidson, 1881 up per Eifelian Ge nus Athyris M’Coy, 1844 Peratos arrectus Copper, 1986 Type spe cies: Terebratula concentrica von Buch, 1834; Figs 31A–J, 32 Eifel, Germany; Eifelian * 1986 Peratos arrectus n. sp. – Cop per: pp. 859–860, text- figs 16–17, pl. 74, figs 32–36, pl. 75, fig. 1. Athyris ex gr. concentrica (von Buch, 1834) Fig. 30F–O Ma te rial: Over sev enty spec i mens (mostly com plete ar tic u lated shells) from Aferdou, ZPAL Bp 68/1/40 and over fif teen spec i - ex gr. 1996 Athyris (Athyris) concentrica (von Buch 1834) –

Fig. 27. Trans verse se rial sec tions of Desquamatia (D.) deserti sp. nov. through shells ZPAL Bp 68/2/26/3 (A) and 68/2/26/4 (B) from Maharch. Distances mea sured in millimetres from the tip of the ventral umbo 284 A. T. HALAMSKI & A. BALIÑSKI

Fig. 28. Middle De vo nian Atrypida from south ern MaÎder. A–E. Prodavidsonia sp. Ar tic u lated shell ZPAL Bp 68/1/32/1 from Aferdou in dorsal, ventral, lateral, poste rior, and ante rior views. F–K. Carinatina arimaspus (Eichwald in von Buch, 1840). Ar tic u lated shell ZPAL Bp 68/1/31/1 from Aferdou in dor sal, ven tral (G, K), lat eral, pos te rior, and an te rior views. L. Atryparia dispersa (Struve, 1966). Artic u lated shell ZPAL Bp 68/1/28/4 from Aferdou in dor sal view show ing frills

Alvarez et al., pp. 78–82, text-figs 12, 14, pl. 1, figs this dis cus sion (Alvarez and Brunton, 2005); a cau tious ap proach 1–6. and open nomen cla ture are pre ferred. Ma te rial: Four ar tic ulated shells (two subcomplete) from Afer- dou, ZPAL Bp 68/1/34/1–4. Athyris? aff. curvata (von Schlotheim, 1820) De scrip tion: Shell rounded to roundedly pen tag onal in out line, Fig. 30P–T weakly trans verse (width to length ra tio 1.16–1.18), aequibicon- vex to weakly dorsibiconvex, up to 14.9 mm in width. Dorsal aff. 1996 Athyris (Triathyris?) curvata (Schlotheim 1820) – valve subtriangular to rounded in an te rior view; an indis tinct roun- Alvarez et al., pl. 8, figs 46–47. ded fold ap pear ing in an te rior sixth. Ven tral valve with a shal low Ma te rial: One poorly pre served ar tic u lated shell from Aferdou, U-shaped sulcus ap pear ing slightly poste ri orly to midlength, nar - ZPAL Bp 68/1/46/1. row pos te ri orly, some what broad en ing an te ri orly. Ven tral beak De scrip tion: The sin gle avail able shell is ir reg u larly pen tag o nal short, fine, strongly incurved; fo ra men medium-sized, cir cu lar, (nearly tri an gular) in out line, strongly dorsibiconvex, 18.5 mm palintrope small. An te rior commissure uniplicate; tongue mod er - wide, >13.9 mm long, and 11.8 mm thick. Max i mal width at about ately nar row to moder ately broad (0.3–0.5 of the shell width), three-fifths of the shell length. Shoul der an gle ca. 120°. Dor sal rounded to tri an gu lar, very low (Fig. 30J) to moder ately high (Fig. valve strongly con vex, subtrapezoidal in an te rior view be cause of 30O). Growth lines prom i nent. In te rior not stud ied. steep flanks and narrow flat tened fold in the an te rior third; umbo Re marks: Athyris concentrica (von Buch, 1834) has been in ter - massive, beak incurved. Ven tral valve moder ately con vex in um- preted both nar rowly (Alvarez et al., 1996) and widely (Grunt and bonal re gion, with a wide, shal low, U-shaped (nearly flat-bot - Weyer, 2002). The stud ied ma te rial does not al low par tic i pa tion in tomed) sulcus, be gin ning at valve mid-length. An te rior commis- MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 285

Fig. 29. Trans verse se rial sec tions of Gruenewaldtia latilinguis through shell ZPAL Bp 68/1/30/2 from Aferdou. Distances mea sured in millimetres from the tip of the ven tral umbo sure uniplicate, tongue 9.5 mm wide, rounded, high. Traces of Camarium sp. strong growth lines, oth er wise shell smooth. In te rior un known. Figs 31K–Z, 33 Re marks: This sin gle spec i men is ten ta tively compared, on ac - count of simi lar ity in shape, to the poorly known early to middle v. 1934 Dicamara plebeja (Sowerby) – Torley, p. 116–117, Eifelian Athyris curvata (von Schlotheim, 1820) illus trated, but text-fig. D, figs 51–54. not de scribed, by Alvarez et al. (1996). Cleiothyridina blacourti Ma te rial: Seven (mostly subcomplete) ar tic ulated shells from Brice, Mottequin and Loones, 2008 from the Givetian (Mid dle to Maharch, ZPAL Bp 68/2/50. Up per varcus zones) of the Boulonnais is simi lar in shape, but the Other ma te rial ex am ined: SMF XVII 278h from Bilveringsen microornamentation is dif fer ent (Brice et al., 2008). (Sauerland), coll. K. Torley (Fig. 33B). De scrip tion: Shell subpentagonal in out line, weakly to mark edly Family Meristidae Hall and Clarke, 1893 ventribiconvex, up to 17.4 mm in length. Dor sal umbo thick, beak Ge nus Camarium Hall, 1859 very short, strongly incurved. Ven tral umbo thick, rather long, Type spe cies: Camarium typum Hall, 1859; Cumber land, beak incurved. An te rior commissure uniplicate with broad, low Maryland, USA; Lower Helderberg Group, Lochkovian trap e zoidal tongue. Shell smooth. Ventral inte rior with long, thin, subparallell dental plates, sup- (after Alvarez and Brime, 2000; Alvarez and Rong, 2002) ported by a long shoe-lifter pro cess; mystrochial plates not re - Re marks: Camarium Hall, 1859 is re ported to dif fer from Dica- vealed. In the dor sal valve septalium short, nar row, supported by a mara Hall and Clarke, 1893 in the lack of mystrochial plates high and long me dian sep tum; lat er ally di rected spiralia of at least (Alvarez and Rong, 2002, p. 1572). This tiny struc ture can be ob- seven whorls (Fig. 31K). served solely through se rial sec tions, but would not ap pear on in - Re marks: At Maharch, this species co-occurs with the exter nally ter nal moulds. Hall and Clarke (1894, p. 772) se lected Atrypa sim i lar Peratos arrectus. The lat ter has straight ante rior comis- plebeia Sowerby, 1840 as the type spe cies of Dicamara, but their sure, is more rounded, and has more distinct growth lines. in ter pre ta tion is based on in ter nal moulds and iso lated valves of a Dis tri bu tion: Sauerland, MaÎder; Givetian. pre sumed Dicamara scalprum Roemer, 1844 (Hall and Clarke, 1894, pl. 33: 16–19; figs 18, 19 are those of a Late De vo nian Family Meristellidae Waagen, 1893 brachi o pod from the Harz, prob a bly not the same spe cies as D. scalprum from the Eifel rep re sented in figs 16, 17). Siehl (1962, Ge nus Meristella Hall, 1859 pl. 32: 1) fig ured se rial sec tions of a “Dicamara plebeja” from Type spe cies: Atrypa laevis Vanuxem, 1842; Albany Gerolstein in the Eifel, but without repre sent ing the exte rior. Alva- County, New York, USA; Lower Heidel berg Froup, Lower rez and Rong (2002, p. 1572, fig. 1069.4) ad mitted the synon ymy Devo nian (see Alvarez and Brime, 2000 for a detailed be tween D. plebeia from Mount Wise near Plymouth, Eng land and dis cus sion) D. scalprum from Germany. Struve (1964, p. 516) ar gued for sep a - rat ing both spe cies. It fol lows that no spec i men of the type spe cies Meristella cf. iconensis Struve, 1964 of Dicamara from the type area in England has ever been se rially Fig. 31AA–EE sec tioned. This is espe cially unfor tu nate in a group full of homo- eomorphic species (e.g., “Dicamara plebeia” sensu Torley, 1934 v cf. 1964 Meristella iconensis n. sp. – Struve: p. 510–513, figs 2–5. is a Camarium; see be low, Fig. 33B). Charac ters of Dicamara Ma te rial: Two ar tic ulated shells (one decorticated) from Aferdou, should be con sidered as un cer tain. ZPAL Bp 68/1/51/1–2. 286 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 287

De scrip tion: Shell pen tag onal, elongate (width to length ra tio 0.80– 1973 Bifida lepida (Archiac and Verneuil 1842) – Cop per, 0.82), mod er ately to mark edly ventribiconvex, up to 22.4 mm long. pp. 123–128, text-figs 1, 2[a], 3, pl. 4, figs 5–8, pl. 5, Ven tral umbo elon gate, beak incurved. Ante rior commissure uni- figs 3–4, pl. 6, figs 1–4, pl. 7, figs 23–24. plicate, tongue subtrapezoidal, moder ately high. Ven tral in te rior: Ma te rial: Four ar tic ulated shells SMF 94896–94898 from Ma- very long traces of the shoe-lifter can be ob served nearly up to valve dÀne el Mrakib, in clud ing one at tached to a bryozoan cov er ing a midlength. part of a shell of Glyptogypa multiplicata, SMF 94871 (Figs 9P, Re marks: These two speci mens differ from Camarium sp., oc cur - 29U, V). ring in Maharch, because of its elon gate shape. They are identi cal De scrip tion: Shell rounded in out line, up to 2.9 mm in width, lon - in ex ter nal char ac ters with Meristella iconensis Struve, 1964 (Me- ger than wide, planoconvex to strongly ventribiconvex, with in - ristellidae; see Struve, 1964). Ap prox i mately co eval Dicamara curved ven tral umbo and ap par ently ab sent fo ra men. The ven tral prunulum (Schnur, 1851) (Meristidae) usu ally has a nar rower or na menta tion con sists of a median pair of thick, subparallel co- tongue. In the absence of serial sections of the Maharch mate rial, stae, sep a rated by a narrow median fur row, and an exter nal pair of open no mencla ture has been used as a pre cau tion. lower and thin ner costae; the dor sal or na menta tion of a me dian costa, a strong pair of costae situ ated later ally from the former, and Family Neoretziidae Dagys, 1972 an exter nal pair of faint costae; the growth lines are sublamellose. Re marks: The mate rial described is included in the Eifelian spe- Ge nus Plectospira Cooper, 1942 cies Bifida lepida. The Givetian B. nitida (Schmidt, 1951) is larger Type spe cies: Terebratula ferita von Buch, 1834; Eifel; and has a more con vex dor sal valve (Schmidt, 1951; Cop per, Eifelian 1973; Halamski, un pub lished). The shell SMF 94871b may be in - terpreted as preserved in the in vivo po si tion (Figs 9P, 29U, V), be - Plectospira ferita (von Buch, 1834) ing at tached dur ing life by a short pedicle, al though a preserva- Fig. 30W–AA tional artefact (two shells cemented post mortem ) can not be ex - cluded. v* 1834 Terebratula ferita n. – von Buch: p. 76, pl. 17, fig. 4. Dis tri bu tion: This spe cies is known from the Eifelian of the Eifel, 1982 Plectospira ferita (v. Buch) – Biernat and Baliñski, Ardennes (Maillieux, 1938 and B. Mottequin, pers. comm. June pp. 857–866, text-fig. 1, pl. 92, figs 1–6, pl. 93, figs 2013), of the Holy Cross Mountains, and MaÎder (Cop per, 1973, 1–7, pl. 94, figs 1–7, pl. 95, figs 1–6. sup ple mented). Ma te rial: A single subcomplete (ventral umbo lack ing) ar tic u- lated shell from Aferdou El-Mrakib, ZPAL Bp 68/1/33/1. Family Kayseriidae Boucot, Johnson and Staton, 1964 De scrip tion: The sin gle avail able shell is subpentagonal in out - line, 13.0 mm wide, >12.2 mm long, and 6.7 mm thick, weakly Ge nus Kayseria Davidson, 1882 dorsibiconvex. Dor sal beak fine, strongly incurved. An te rior com- Type spe cies: Orthis lens Phillips, 1841; England; missure zig zag ging. Or na menta tion of stout plicae, nine dor sal and Mid dle De vo nian eight ven tral ones, sep a rated by wide U-shaped fur rows, in which strong growth lines are visi ble. In te rior not stud ied. Kayseria alvea Copper, 1973 Dis tri bu tion: Plectospira ferita is known from the Middle De vo - Fig. 30HH–KK nian (Eifelian, possi bly also Givetian) of the Eifel, the Holy Cross Mountains, Moravia (Ficner and Havlíèek, 1978), Burma (Ander - * 1973 Kayseria alvea sp. nov. – Cop per: pp. 134–136, text- son et al., 1969), and Tur key (Gourvennec and Hoºgör, 2012). fig. 6, pl. 7, figs 12–22. Ma te rial: Two ar tic u lated shells (one com plete), SMF 98180, 98181, from MadÀne el Mrakib. Family Anoplothecidae Schuchert, 1894 De scrip tion: Shell subcircular in out line, up to 3.9 mm wide, slig- Ge nus Bifida Davidson, 1882 htly lon ger than wide, weakly dorsibiconvex. An te rior mar gin me- di ally in dented. Both valves with me dian sulci. Orna menta tion of Type spe cies: Terebratula lepida d’Archiac ra dial ribs, begin ning in umbonal re gion. On the dor sal valve (Fig. and de Verneuil, 1842; Eifel; Eifelian 30 HH, II), a strong pair of ribs bor der ing the sulcus with fine and Bifida lepida (d’Archiac and de Verneuil, 1842) dense ribs in side and me dium-sized ribs on lat eral flanks; on the Fig. 30U–V, BB–GG ven tral valve (Fig. 30 KK), ribs of ap prox i mately the same size. Re marks: Kayseria alvea is distin guished from the slightly older * 1842 Terebratula lepida Goldfuss, Bonn Mus. – d’Archiac Kayseria lens (Phillips, 1841) and K. dividua (Schnur, 1851) (ex - and de Verneuil, p. 368, pl. 35, figs 2, 2a–c. ter nally in distin guishable and of the same age, there fore prob a bly

Fig. 30. Middle De vo nian Rhynchonellida and Athyridida from south ern MaÎder. A–E. Rhynchonellida fam., gen. et sp. indet. Ar tic u - lated shell ZPAL Bp 68/1/50/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. F–O. Athyris ex gr. concentrica (von Buch, 1834), Aferdou. F–J, K–O. Artic u lated shells ZPAL Bp 68/1/34/1, 2 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. P–T. Athyris? aff. curvata (von Schlotheim, 1820). Artic u lated shell ZPAL Bp 68/1/46/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. U–V, BB–GG. Bifida lepida (d’Archiac and de Verneuil, 1842). Spec i mens from MadÀne el Mrakib. BB–DD. Artic ulated shell SMF 94896 in ven tral, lat eral, and dor sal views. EE–GG. Ar tic u lated shell SMF 94897 in dorsal, lat eral, and ven tral views. U, V. Ar - tic ulated shell SMF 94871b in proba ble in vivo posi tion [see dis cussion in the text] on the ven tral valve of Glyptogypa multiplicata (cf. Fig. 9P) in ven tral and lat eral views. W–AA. Plectospira ferita (von Buch, 1834). Ar tic ulated shell ZPAL Bp 68/1/33/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. HH–KK. Kayseria alvea Copper, 1973. Ar tic ulated shells from MadÀne el Mrakib. HH. Spec i men SMF 98180 in dorsal view. II–KK. Spec i men SMF 98181 in dorsal, lat eral, and ven tral (with an at tached palaeocopide ostracod, pos sibly a rep re sen ta tive of Primitiopsidae; det. E. Olempska-Roniewicz) views 288 A. T. HALAMSKI & A. BALIÑSKI

Fig. 31. Middle De vonian Atrypida and Athyridida from south ern MaÎder. A–J. Peratos arrectus Copper, 1986. Ar tic ulated shells in dor sal, cen tral, lat eral, poste rior, and an te rior views. A–E. Spec i men ZPAL Bp 68/2/40/3 from Maharch. F–J. Spec i men ZPAL Bp 68/1/40/2 from Aferdou. K–Z. Camarium sp. from Maharch. K. Partly bro ken ar tic u lated ZPAL Bp 68/2/50/4 shell show ing spiralium; dorsal view. L–P. Ar tic ulated shell ZPAL Bp 68/2/50/1 dorsal, ventral, lateral, poste rior, and an te rior views. Q–U. Artic ulated shell ZPAL Bp 68/2/50/2 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. V–Z. Ar tic u lated shell ZPAL Bp 68/2/50/3 in dor sal, ven tral, lat eral, pos te rior, and an te rior views. AA–EE. Meristella cf. iconensis Struve, 1964. Ar tic ulated shell ZPAL Bp 68/1/51/1 from Aferdou in dor - sal, cen tral, lat eral, pos te rior, and an te rior views MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 289

Fig. 32. Trans verse se rial sec tions of Peratos arrectus through shells ZPAL Bp 68/2/40/1 from Maharch (A), 68/1/40/1 from Aferdou (B), and 68/2/40/2 (C) from Maharch. Dis tances mea sured in milli metres from the tip of the ven tral umbo

syn on y mous; Davidson, 1882, p. 24) in pos sessing a stron ger rib Microornamentation (Fig. 34L) of gran ules ar ranged on microfila, pair, bor der ing the sulcus (ab sent in the lat ter). the lat ter 2–3 per mm. Dis tri bu tion: Eifel, upper most Eifelian in the tradi tional sense Re marks: This spe cies is included into Spinocyrtia, on ac count of (Ahbach Beds, mostly Müllert ho ri zon; Cop per, 1973), there fore macro- and microornamentation and rel a tively low ven tral inter- mostly low er most Givetian ac cord ing to the mod ern def i ni tion area (John son, 2006a). It is quite simi lar to S. elburzensis from the (Walliser et al., 1995; Bultynck and Hollevoet, 1999); MaÎder, Middle De vonian of Elburz (Iran) which is, how ever, al ways Mid dle De vo nian (Eifelian?). ventribiconvex (Brice, 1973). Prob a bly conspecific rep re sen ta - tives of this ge nus were de scribed by Struve (1995) from co eval Order Spiriferida Waagen, 1883 de posits of the north ern MaÎder. Family Spinocyrtiidae Ivanova, 1959 Ge nus Spinocyrtia Fredericks, 1919 Type spe cies: Delthyris granulosa Conrad, 1839; New York, USA; Givetian Spinocyrtia cf. elburzensis Brice, 1973 Fig. 34A–H, L cf. 1973 Spinocyrtia elburzensis nov. sp. – Brice in Brice et al.: pp. 203–206, text-fig. 14, pl. 22, figs 9–10, pl. 23, figs 1, 5, pl. 24, figs 2–3. 1995 Spinocyrtia (Spinocyrtia) elburzensis elburzensis Brice – Struve, p. 100, figs 38–41. Ma te rial: Forty-six spec i mens SMF 94880–94892, MB.B.9384c, and MB un numbered (V. Ebbighausen’s col lec tion) from the Dro- tops level at MadÀne el Mrakib. De scrip tion: Shell up up to 56.4 mm in width, transverse (width- to-length ratio 1.32–1.84; mean 1.50; N = 7), more frequently dorsibiconvex (among mea surable spec i mens fourteen were dorsi- biconvex and five were ventribiconvex, the lat ter be ing usu ally smaller); hinge line nearly straight. Ventral valve convex, with a deep V-shaped sulcus orig i nat ing at beak, the lat ter incurved; interarea concave, subrectangular, apsacline, delthyrial cover not pre served. Dor sal valve con vex with a rounded fold (ca. 12 mm wide and 5 mm high in an in di vid ual ca. 45 mm wide) be gin ning at Fig. 33. Trans verse se rial sec tions of Camarium sp. through beak; the fold with a me dian groove, the width of which is from shells ZPAL Bp 68/2/50/5 from Maharch (A) and SMF XVII 278h one-third to a half of that of the fold. Or na menta tion of straight pli - from Bilveringsen (B). Dis tances mea sured in milli metres from ca tions, 14–20 per flank (Nc/w 0.28 to 0.44; mean 0.36; N = 6). the tip of the ven tral umbo 290 A. T. HALAMSKI & A. BALIÑSKI

Fig. 34. Middle De vo nian Spiriferida from southern MaÎder. A–H, L. Spinocyrtia cf. elburzensis Brice, 1973. Both spec i mens from MadÀne el Mrakib. A–C, L. In complete ar tic ulated shell SMF 94880 in dorsal, ven tral, and lat eral views; en large ment of microorna- mentation. D–H. Ar tic ulated shell MB.B. 9384c in dor sal, ven tral, lat eral, ante rior, and pos te rior views. I–K. Reticularioidea fam., gen. et sp. indet. Ven tral valve ZPAL Bp 68/1/53/1 from Aferdou in ven tral, pos te rior, and lat eral views

Family Delthyrididae Phillips, 1841 aequibiconvex, 20.8 mm wide, 14.8 mm long, and 12.0 mm thick. Dorsal valve wid est near the hinge line, with three pairs of thick Ge nus Ivanothyris Havlíèek, 1957 lat eral plica tions and a high me dian fold; lateral plica tions low and Type spe cies: Spirifer gibbosus Barrande, 1879; Bohe mia; acute, be gin ning in umbonal re gion, the most lat eral pair nearly Lochkovian im per cep ti ble; me dian fold be gin ning at umbo. Dor sal umbo thick, beak short, strongly incurved. Ven tral valve wid est about its mid- Ivanothyris aculeata (Schnur, 1851) length, with a me dian V-shaped sulcus be gin ning at umbo and Fig. 35A–E three pair of thick plica tions, dimin ish ing in size to wards lat eral * 1851 Sp[irifer] aculeatus nob. – Schnur: pp. 10–11. flanks. Ven tral interarea occu pying less than half of the shell . 1853 Sp[irifer] aculeatus m. – Schnur, p. 203, pl. 34, fig. width, apsacline; umbo elon gate, rather thick, beak incurved. 2a–f. Growth lines strong, 3–4 per mm. 1963 Delthyris aculeata – Vandercammen, pp. 112–116, Dis tri bu tion: This spe cies is known from the Eifelian of the Eifel pl. 11, figs 1–8. (Schnur, 1853), of the Ardennes (Jemelle For mation; Vander- 1997b “Spirifer aculeatus” Schnur 1853 – Gourvennec in cammen, 1963), of the Holy Cross Mountains (Halamski, un pub- Boumendjel et al., p. 109–110. lished), and of the MaÎder; it is equally rare in all four regions. It has been re ported from the lower part of the Chefar el Amhar For- Ma te rial: Single subcomplete ar tic u lated shells, one from Afer- mation (up per most Emsian or Eifelian) of the Saoura val ley (Go- dou, ZPAL Bp 68/1/35/1, and one from MadÀne el Mrakib, MB urvennec in Boumendjel et al., 1997b). un num bered (V. Ebbighausen’s col lec tion). De scrip tion: The single available shell is rhombic in outline, MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 291

Ge nus Quiringites Struve, 1992 Ma te rial: A single frag mentary ven tral valve from Aferdou, Type spe cies: Spirifera elegans Steininger, 1853; ZPAL Bp 68/1/48/1. Gerolstein, Eifel; Nohn Forma tion, lower Eifelian De scrip tion: The available speci men is an incom plete half of a ventral valve, length ca. 17 mm, total width esti mated at ca. 28 Quiringites arensentiae Schemm-Gregory, 2009 mm. Outline subtrapezoidal in an te rior view with steep flanks and Fig. 35F–P, R a ca. 5 mm wide (esti mated) deep median sulcus. Beak elongated, incurved. Interarea trian gu lar, apsacline, ca. 3 mm high. Orna men - v* 2009 Quiringites arensentiae sp. nov. – Schemm-Gregory: ta tion con sist ing of both ra dial and con cen tric el e ments: seven pp. 11–13, figs 6B, 7–9. rounded pli ca tions, di minish ing in size to wards the lat eral flank, Ma te rial: Twenty-five complete or subcomplete artic u lated shells sep a rated by U-shaped interspaces; strong, un du lat ing, sublamel- and sev eral more frag mentary ones, SMF 94872, 94873, 94893– lose growth lines, 7–8 per 5 mm, covered by ra dial striae (Fig. 94895, and MB un numbered (V. Ebbighausen’s col lec tion); four 35S; capillae sensu Krans, 1971). Me dian sep tum high. dor sal and three ven tral valves SMF 94843, 94844 and MB unnum - Re marks: This frag mentary spec i men may be in cluded con fi - bered (V. Ebbighausen’s col lec tion); all from MadÀne el Mrakib. dently in Cyrtinopsis, on ac count of the char ac ter is tic or na men ta- De scrip tion: Shell trans verse, mucronate, up to 24.9 mm in width, tion and the ven tral me dian sep tum. Spe cific iden ti fi ca tion is more ventribiconvex. Dor sal valve with a rounded fold orig i nat ing at prob lematic; a rounded and mod er ately trans verse out line in dica- beak. Ven tral valve with wide, shallow, U-shaped sulcus orig i nat - tes af fin i ties with C. brachyptera, the only spe cies pres ent in Nort- ing at beak; the latter strongly incurved; interarea low, apsacline, hern Af rica, ac cord ing to Struve (1965b). Com par i son with the subrectangular, and strongly con cave. Orna menta tion of strong, sin gle chronosubspecies, in par tic u lar with C. brachyptera maide- rounded pli ca tions, (4–)5–6(–10) per flank of the dor sal valve. rensis Drot, 1964 is not pos sible, given the scar city of the ma te rial. Microornamentation (Fig. 35R) of ir reg ularly placed swol len Dis tri bu tion: The type mate rial of C. brachyptera co mes from the spine bases. Eifelian Jemelle For mation (B. Mottequin, pers. comm., 2013). Ven tral in te rior (Fig. 35O): teeth not pre served; den tal plates The stratigraphic dis tri bution of this spe cies is re ported to be con- embed ded in a strongly de vel oped cal lus; muscle scars not visi ble; fined to the middle part of the Eifelian (Struve, 1965b), whereas muscle area pro longed an te ri orly by a short and low myophragm; the spec i men discussed be longs proba bly to the late Eifelian, al - gonoglyphs present abmedially from dental plates. Dor sal in te rior though its col lec tion from the rub ble pre cludes any de fin i tive con - (Fig. 35P): ctenophoridium with ca. 11 lamellae; dental sock ets clu sions. deep; a pair of strong plates is formed by inner socket ridges coales - cent with crural bases (these are “brachiophores” sensu Schemm- Greg ory, 2009; the ter minol ogy of Brunton et al., 1996 and of Ba- Family Reticulariidae Waagen, 1883 liñski and Sun, 2010: fig. 5 is fol lowed) poorly pre served, di ver - gent; a thin, low, rounded median myophragm ex tends up to ca. Ge nus Rhenothyris Struve, 1970 two third of the valve length. Type spe cies: Rhenothyris rhenana Struve, 1970; Eifel; Re marks: Char ac ters sep a rat ing Q. arensentiae and Q. elegans mid dle Eifelian given by Schemm-Gregory (2009, p. 11) are largely of low tax o- nomic value. In par tic ular, de vel opment of the callus does not seem Rhenothyris sinuata (Gürich, 1896) to be a good discriminator at spe cies level. More over, as shown by Fig. 36M–Q the pres ent study, Q. arensentiae has (al though sel dom) up to ten * 1896 Reticularia sinuata nov. nom. – Gürich: pp. 259–260, costae per flank (Fig. 35K); this charac ter cannot therefore be used pl. 9, fig. 10. for dis tin guishing the taxa dis cussed [Schemm-Gregory (2009) sug- v. 1966 Eoreticularia eifeliensis (Frech in Scupin) – Biernat, gested that Q. arensentiae has up to 7 pli ca tions, whereas Q. elegans pp. 126–128, text-figs 43–44, pl. 30, figs 13–19 [non has up to 10 pli ca tions]. However, Q. arensentiae and Q. elegans Reticulariopsis eifliensis (Scupin, 1900)]. are judged to be sep a rate spe cies, on ac count of the incurvation of Ma te rial: Two subcomplete ar tic u lated shells from Aferdou, the ven tral umbo (Fig. 35I, L, M), con stantly much greater in the ZPAL Bp 68/1/36/1–2. for mer taxon. De scrip tion: Shell subrectangular to transversely el liptic in out- Dis tri bu tion: MaÎder (north and south), middle Eifelian to mid dle line (hinge line and me dian part of the an te rior commissure subpa- Givetian (Schemm-Gregory, 2009, sup ple mented). Q. cf. arensen- rallel) with broadly rounded postero-lateral mar gins, trans verse tiae was re ported from the Drâa val ley (Schemm-Gregory, 2009). (width-to-length ra tio 1.53–1.56), strongly ventribiconvex, up to 24 mm in width. Maxi mal width at midlength of the dor sal valve. Dorsal valve with a low, rounded fold ap pear ing in pos te rior fourth; Family Cyrtinopsidae Wedekind, 1926 interarea low, ap proxi mately orthocline. Ven tral valve with a shal - Ge nus Cyrtinopsis Scupin, 1896 low U-shaped sulcus bor dered by a pair of thick ribs; the sulcus and the ribs ap pear ing in umbonal re gion; interarea high, apsacline, Type spe cies: Spirifer undosus Schnur, 1853; Gerolstein gently curved; umbo thick, beak incurved. An te rior commissure Syncline, Eifel; Middle De vonian uniplicate; tongue as broad as ca. one-third of the shell, high, Cyrtinopsis cf. brachyptera (Maillieux, 1914) rounded. Shell mac roscop i cally smooth; microornamentation of Fig. 35Q, S concentric growth lamellae bear ing bands of mar ginal spine bases, 3–4 per mm. Inte rior un known. cf. 1914 Cyrtina undosa Schnur sp. var. brachyptera nov. var. Re marks: The mate rial described is very simi lar to the late Eife- – Maillieux: pp. 4–6, figs 3–4. lian to early Givetian taxon, de scribed by Gürich (1896) from the p cf. 1963 Cyrtinopsis undosa (J. Schnur, 1851) Vandercam- Holy Cross Mountains and re ported by Biernat (1966) un der er ro - men, pp. 101–104, pl. 10, figs 3–8. ne ous ge neric and spe cific names (Halamski, 2004a). This is the cf. 1964 Cyrtinopsis undosa maiderensis ssp. n. – Drot, pp. youngest known member of Rhenothyris (Struve, 1970; Halamski, 69–70, pl. 6, figs 1–8. 2004a). cf. 1965b Cyrtinopsis brachyptera (Maillieux 1914) – Struve, Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder; Middle De- pp. 13–22, pl. 1, figs 2–4, pl. 2, figs 1–2, 4, pl. 3, fig. 4. vo nian. 292 A. T. HALAMSKI & A. BALIÑSKI

Fig. 35. Middle De vonian Spiriferida from southern MaÎder. A–E. Ivanothyris aculeata (Schnur, 1851). Artic ulated shell ZPAL Bp 68/1/35/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and ante rior views. F–P, R. Quiringites arensentiae Schemm-Gregory, 2009 from MadÀne el Mrakib. F–J. Artic ulated shell SMF 94872 in dorsal, ven tral, lat eral, pos te rior, and ante rior views. K, L. Artic ulated shell SMF 94893 in dorsal and pos te rior views. M, N, R. Artic u lated shell SMF 94894 in dorsal and pos te rior views, and microornamentation (R). O. Ven tral in te rior SMF 94843. P. Dorsal in te rior SMF 94844. Q, S. Cyrtinopsis cf. brachyptera (Maillieux, 1914). Frag mentary ven - tral valve ZPAL Bp 68/1/48/1 from Aferdou (Q) and microornamentation (S) MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 293

Ge nus Deltospirifer Wang and Rong, 1986 U. rzhonsnitskajae Mamedov, 1961 from the Mid dle De vonian of Type spe cies: Elytha transversa Wang, 1956; Guangxi, Danzik in the Nakhitchevan Re public (Cau ca sus), which has, southern China; Emsian however, only one or two costae per flank (Mamedov, 1961). This taxon was in cluded in Nakazatothyris Minato and Kato, 1977, pro - Deltospirifer? new species posed as a sub ge nus of Undispirifer to accom mo date three Lower Fig. 36R–V to Mid dle De vo nian spe cies with ob so les cent costation (Minato and Kato, 1977). However, Gracianova et al. (1990) re clas si fied Ma te rial: A single subcomplete ar tic u lated shell from Aferdou, U. rzhonsnitzkajae within Deltospirifer Wang and Rong, 1986; ZPAL Bp 68/1/51/1. Chen and Tazawa (2003) reclas si fied U. (N.) vandercammeni De scrip tion: Shell widely, ir reg u larly pen tag o nal in out line, with Minato and Kato, 1977, the type spe cies of Nakazatothyris, within gently rounded antero-lat eral mar gins and car di nal ex trem i ties, Kymatothyris Struve, 1970. Carter and Gourvennec (2006) con sid- trans verse (width to length ra tio 1.64), mod er ately ventribiconvex, ered Nakazatothyris as a ju nior subjec tive synonym of Undispi- 26 mm in width. Maxi mal width poste ri orly to midlength of the rif er. The mate rial described does not allow partic i pation in this dor sal valve. Dorsal valve with a low, flat fold ap pear ing in um- dis cus sion. bonal re gion, bor dered by a pair of dis tinct fur rows; interarea low, Rep re sen ta tives of Undispirifer from Zemmour (Sougy, 1964), apsacline. Ven tral valve with a shal low sulcus, pos te ri orly V- Boulonnais (U. rigauxi Brice and Loones, 2002), and the Lahn shaped, an te ri orly ap prox i mately flat-bot tomed, bor dered by a pair Syncline (Halamski unpub lished; SMF col lec tions) are strongly of thick ribs; the sulcus and the ribs ap pear ing in umbonal re gion; costate. Geograph i cally and stratigraphically re stricted samples of interarea high, apsacline, incurved; umbo not pre served. An te rior Undispirifer usually show rather re stricted vari a tion (e.g., Brice commissure paraplicate; tongue as broad as ca. one-third of the and Loones, 2002); this is against the wide in ter pre ta tion of the shell, low, subtrapezoidal. A sin gle pair of low, rounded, ca. 1.8 taxa (e.g., Schnur, 1853; Davidson, 1864, p. 36–37; Cottreau, mm wide costae ad ja cent to the fur rows border ing the dor sal fold, 1940, p. 198), as observed al ready by Sandberger and Sandberger oth er wise shell smooth; microornamentation of spine bases ar - (1850–56, p. 314) and more re cently by Struve (1970, 1981a). ranged in dense con cen tric lines, 8–9 per mm. Inte rior un known. Re marks: This shell differs from Rhenothyris sinuata co-occur - ring in the same out crop, in microornamentation den sity, shell out - Family Thomasariidae Cooper and Dutro, 1982 line (more trans verse, po sition of the great est width), in cli na tion of the dorsal interarea, smaller ventribiconvexity, incip i ent costation, Ge nus Thomasaria Stainbrook, 1945 and form of the tongue. It is ten ta tively in cluded in Deltospirifer, Type spe cies: Thomasaria altumbona Stainbrook, 1945; on account of its overall simi lar ity to the Emsian D. trans ver sus Iowa; In de pend ence Shale, Frasnian (Wang, 1955) (Wang and Rong, 1986, pl. 71: 1–18). Dif fer ences Re marks: Pyramidalia Nalivkin, 1947 (type spe cies Spirifera in clude higher interarea and less de vel oped costation in the Afer- sim plex Phillips, 1841) has been con sidered (John son, 2006b: dou spec i men; it rep re sents a new spe cies, which is not named, be - 1882) as a ju nior subjec tive synonym of Squamulariina Frede- cause the ma terial is too scarce. ricks, 1916 (type spe cies Cyrtina parva Gürich, 1896; Cyrtinidae, Spiriferidina). This view is pro vi sionally re jected here, on ac count Ge nus Undispirifer Havlíèek, 1957 of the impunctate shell of the Moroc can mate rial of Thomasaria? sim plex (Phillips, 1841) in ves ti gated; this however, should be con - Type spe cies: Spirifer undiferus Roemer, 1844; Rhenish firmed on the type ma te rial of that spe cies from south Devon Slate Mountains and Eifel [Roemer (1844, p. 73) indi cates (Phillips, 1841). Fol low ing Baliñski (1979), Pyramidalia is pro vi - three local i ties for his new species; this problem has not sionally con sidered to be a ju nior subjec tive synonym of Thoma- been resolved in the revi sion made by Vandercammen saria Stainbrook, 1945 although spines di ag nostic for the lat ter ge - (1967, p. 6)]; Middle Devo nian nus could not be found in the ma te rial stud ied, ow ing to ae olian corrasion and not with stand ing the fact that Frasnian T. sim plex Undispirifer aff. rzhonsnitzkajae Mamedov, 1961 sensu Baliñski (1979) is prob a bly not conspecific with the Mid dle Fig. 36W–AA De vo nian Spirifera simplex Phillips, 1841 (Halamski, 2004a). aff. 1961 Undispirifer rzhonsnitzkajae Mamedov, sp. nov. – Mamedov: pp. 53–55, pl. 5, fig. 2. Thomasaria? sim plex (Phillips, 1841) aff. 1974 Undispirifer rzhonsnickajae Mamedov, 1961 – Abra- Fig. 36A–L mân, pp. 62–63, pl. 16, fig. 1. Ma te rial: Seven mostly fragmen tary spec imens from Maharch, * 1841 Spirifera simplex – Phillips: p. 71, pl. 29, fig. 124a; pl. ZPAL Bp 68/2/37/1–7. 60, fig. 124a. De scrip tion: Shell subrectangular to rounded in out line (width-to- v. 1966 Pyramidalia cf. sim plex (Phillips, 1841) – Biernat, p. length ra tio 1.02–1.25), weakly to strongly ventribiconvex, up to 136, pl. 27, fig. 1. at least 28 mm in width. Dorsal valve with a flat fold ap pear ing at ? 1985 Thomasaria? serrensis n. sp. – Brice, pp. 144–146, poste rior fourth; the fold has a nar row median fur row; maxi mal text-fig. 12, pl. 3, figs 3–5. width of the dor sal valve about its midlength; interarea very low, Ma te rial: Six spec i mens (in clud ing four subcomplete ar tic u lated anacline. Ven tral valve with a narrow and rather low, apsacline shells) from Aferdou, ZPAL Bp 68/1/39/1, and two subcomplete interarea and a shallow but rela tively wide sulcus appear ing at ar tic u lated shells from Maharch, ZPAL Bp 68/2/39/1. poste rior fourth. An te rior commissure uniplicate; tongue subtrape- De scrip tion: Shell py rami dal (thick ness to width ra tio 0.55–0.6), zoidal, rather low, its width 0.40–0.55 of that of the shell. Or na - trans verse (width-to-length ra tio ca. 1.4–1.5), strongly ventribi- menta tion of rounded costae, weak or very weak (to ev a nes cent?) con vex, up to ca. 40 mm in width. Hinge line straight. Max i mal ex cept the pair bor der ing the ven tral sulcus, which are some what width be tween the hinge line and midlength of the dor sal valve. higher and thicker, 3–5 (?) per flank; a single case of bifur ca tion Dorsal valve weakly to strongly con vex; a flat fold in the ante rior ob served. Microornamentation not pre served. In te rior: long den tal third, rela tively low, but distinct to imper cep ti ble; interarea linear, plates in the ven tral valve; oth er wise un known. more or less orthocline. Ven tral valve pyram i dal with a shallow, Re marks: The scarce mate rial, described here, is most simi lar to U-shaped sulcus, be gin ning in umbonal re gion and tri an gularly 294 A. T. HALAMSKI & A. BALIÑSKI MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 295

wid en ing an te ri orly; interarea very high, slightly procline, more sel - Order Spiriferinida Ivanova, 1972 dom catacline, flat, more rarely weakly concave, as wide as the Family Cyrtinidae Fredericks, 1911 hinge line length; delthyrium nar row (0.2–0.3 of the interarea width), with api cal an gle at tain ing about 20°, cov ered in the up per Ge nus Cyrtina Davidson, 1858 third or fourth by an api cal plate. An te rior commissure uniplicate; Type spe cies: Cyrtina heteroclita Defrance, 1828; Néhou, tongue rounded to subtrapezoidal, moder ately high to high, oc cu py - Normandy; Pragian (or lower Emsian?) ing 0.35–0.5 of the shell width. Shell smooth. In te rior not stud ied. Re marks: The spe cies-level tax onomy of Cyrtina is diffi cult to Re marks: The ma te rial stud ied al lows an ap pre ci a tion of the elab o rate. Al though usu ally there is rel a tively small vari a tion ontogenetic and intraspecific vari a tion of the spe cies dis cussed. within a sample, differ ent samples show a lack of simi lar ity. It is Smaller in di vid u als have the maxi mal shell width near the hinge un clear whether mor pho log i cal char ac ters, such as the num ber of line (like shells fig ured by Phillips, 1841), while in larger ones it is costae, valve form, or interarea po sition, are un der ge netic or en vi - sit uated near the mid-length of the dor sal valve. The ven tral inter- ronmen tal control. This may result in either all-lumping or all- area can be flat or concave; the latter variant was distin guished split ting tax on o mies. (Sobolew, 1909) as Spirifer simplex var. subsimplex Sobolew, Late Eifelian to early Givetian popu la tions of Cyrtina from the 1909, but this char ac ter does not seem to be tax o nomi cally signif i - Holy Cross Mountains and the Eifel have been shown to rep re sent cant. Thomasaria? serrensis Brice, 1985 from the Givetian (Mid - C. sauvagei Rigaux, 1908 (Halamski unpub lished) and not its evo - dle varcus Zone) of the Montagne Noire is ex ter nally indistin- lution ary Pragian (to early Emsian?) prede ces sor C. intermedia guishable from the small spec i mens of T.? sim plex from Aferdou 2hlert, 1887 (for this spe cies, see Gourvennec, 1989), the main (Fig. 36C–G): synon ymy of the two species therefore is ten ta tively dif fer ence be ing in the shape of the dor sal valve (mod er ately trans - proposed; however, it should be re membered that the type mate rial verse in the former and strongly transverse in the latter; see Table of this species is in need of re vi sion (see above, re marks on the ge- 4). The mea gre ma terial from MaÎder is diffi cult to inter pret. Sam- nus). Spirifer pyramidalis Schnur, 1853 and S. nudus Schnur, 1853 ples from Aferdou and Maharch show bio met ric char ac ter istics of (both from the Eifel) are also quite close to T. sim plex (Mottequin, C. sauvagei and C. intermedia, respec tively (Ta ble 4); the speci - 2008b, p. 516). mens from the latter local ity are sim i lar to those from the late Dis tri bu tion: Eifel, Holy Cross Mountains, MaÎder, Montagne Emsian to early Eifelian strata of Saoura (Le Maître, 1952a; see Noire?; Mid dle De vo nian. also Boumendjel et al., 1997b). Given the eco log i cal sep a ra tion between the two local i ties, the two samples are described sepa - Fam ily un known rately; one of them in open no mencla ture. Co-oc cur rence of sev - eral species of Cyrtina in strata of the same age has already been Reticularioidea fam., gen., et sp. indet. re ported (Keyes and Pitrat, 1978). For ex ample, two spe cies of Fig. 34I–K Cyrtina are present in the Givetian of La Serre (Montagne Noire; Brice, 1985); one (Cyrtina sp. 1 sensu Brice, 1985) is quite simi lar Ma te rial: One in complete ventral valve ZPAL Bp 68/1/53/1 from to C. sauvagei, whereas the other (Cyrtina sp. 2 sensu Brice, 1985) Aferdou. might be compared to C. ex gr. intermedia. Cyrtina hamiltonensis De scrip tion: The single available ventral valve is subelliptic in (Hall, 1857) is an approx i mately coeval species from the eastern out line, 13.1 mm wide (in complete; es ti mated wisth ca. 14 mm) United States: it is very simi lar to C. sauvagei; dif fer ences in clude and 10.5 mm long. A faint sulcus is pres ent an te ri orly. Beak thick; slightly more trans verse shape and slightly larger num ber of co- interarea apsacline, weakly con cave; delthyrium open, its apical stae. It is not ex cluded these taxa are conspecific but this can not be an gle ca. 80°. Shell mac ro scop i cally smooth except rare promi - de cided at pres ent. Re ports of C. hamiltonensis from Africa (Ge- nent, sublamellose growth limes; microornamentation not pre - vin, 1964; Schemm-Greg ory and Jansen, 2005) may possi bly re fer served. Poorly pre served den tal plates pres ent; oth er wise, in te rior to C. sauvagei. un known. Re marks: This poorly pre served spec i men dif fers from Rheno- thyris sinuata co-oc cur ring in the same out crop, in or na menta tion; Cyrtina sauvagei Rigaux, 1908 it is in cluded in the superfamily Reticularioidea Waagen, 1883, on Fig. 36BB–FF ac count of an open delthyrium, a mac ro scop i cally smooth shell, and sublamellose growth lines. The overall simi lar ity to Yeothyris * 1908 Cyrtina Sauvagei – Rigaux: p. 6, 20; pl. 1: 8. Struve, 1992 is noted, yet the lack of microornamentation and in - ? 1940 Cyrtina heteroclita Defr. – Cottreau, p. 198, pl. 7, figs ter nal struc tures pre cludes the pos si bil ity of iden ti fi ca tion. 13, 13a.

Fig. 36. Middle De vonian Spiriferida and Spiriferinida from south ern MaÎder. A–L. Thomasaria? sim plex (Philips, 1841). A, B. In com- plete artic ulated shell ZPAL Bp 68/2/39/1 from Maharch in dorsal and an te rior views. C–G. Ar tic ulated shell ZPAL Bp 68/1/39/2 from Aferdou in dor sal, ven tral, lat eral, ante rior, and pos te rior views. H–L. Ar tic u lated shell ZPAL Bp 68/1/39/1 from Aferdou in dor sal, ven - tral, lat eral, an te rior, and pos te rior views. M–Q. Rhenothyris sinuata (Gürich, 1896). Ar tic u lated shell ZPAL Bp 68/1/36/1 from Aferdou in dor sal, ven tral, lat eral, pos te rior, and an te rior views. R–V. Deltospirifer? sp. nov. Ar tic u lated shell ZPAL Bp 68/1/51/1 in dor sal, ven - tral, lat eral, pos te rior, and an te rior views. W–AA. Undispirifer aff. rzhonsnitzkajae Mamedov, 1961. Artic u lated shell ZPAL Bp 68/2/37/1 from Maharch in dor sal, ven tral, lat eral, pos te rior, and ante rior views. BB–FF. Cyrtina sauvagei Rigaux, 1908. Ar tic ulated shell ZPAL Bp 68/1/38/1 from Aferdou in dorsal, ven tral, lat eral, pos te rior, and ante rior views. GG–KK. Cyrtina ex gr. intermedia 2hlert, 1887. GG, HH. Ar tic u lated shell ZPAL Bp 68/2/38/2 from Maharch in dor sal and lat eral views. II–KK. Ar tic u lated shell ZPAL Bp 68/2/38/2 from Maharch in dor sal, lat eral, and pos te rior views 296 A. T. HALAMSKI & A. BALIÑSKI

Ta ble 4 Com par i son of se lected bio met ric char ac ter is tics of se lected Lower and Mid dle De vo nian rep re sen ta tives of Cyrtina

Character Number of Width to length ratio Age Source Taxon and sample costae minimal value average maximal value N C. intermedia topotypic (Brittany) Pragian 3–6 1.44 1.94 2.72 22 Gourvennec, 1989 late Emsian to this work (original C. intermedia Saoura (2–)3 1.56 1.84 2.29 12 early Eifelian material of Le Maître) C. hamiltonensis Traverse Group, Mi. Middle Devonian 5–8 1.40 1.65 1.98 24 Keyes and Pitrat, 1978 C. hamiltonensis Onondaga Fm., N.Y. Eifelian 5–8 1.19 1.56 1.75 9 Feldman, 1994 late Eifelian to C. sauvagei Holy Cross Mts. (3–)4–6(–7) 1.34 1.50 1.74 22 Halamski, unpublished early Givetian C. sauvagei topotypic (Boulonnais) early Givetian 5-7 1.30 1.46 1.62 9 Brice, 1988 late Eifelian to C. sauvagei Aferdou 4–5 1.49 1.62 1.76 3 early Givetian this work C. ex gr. intermedia Maharch early Givetian 3–4 1.79 1.89 2.00 2

? 1964 Cyrtina Hamiltonensis Hall – Sougy, p. 161. Cyrtina with five costae per flank was de scribed from the pre - v. 1966 Cyrtina heteroclita intermedia – Biernat, p. 133–135, sumed upper Givetian of the Zap river val ley in south-east ern Tur - text-fig. 47, pl. 23, figs 9–27, pl. 32, fig. 14. key under the name of C. intermedia (Gourvennec and Hoºgör, ? 1985 Cyrtina sp. 1 – Brice, p. 147, pl. 3, figs 15–16. 2012); it is differ ent from the mate rial dealt with here. 1988 Cyrtina sauvagei Rigaux 1908 – Brice, pp. 359–360, pl. 43, figs 11–12. ? 2005 Cyrtina cf. hamiltonensis (Hall, 1857) – Schemm- Greg ory and Jansen, p. 27. PALAEOBIOGEOGRAPHY Ma te rial: Three speci mens (two artic u lated shells) from Aferdou, ZPAL Bp 68/1/38/1–3. Mid dle De vo nian brachi o pod fau nas De scrip tion: Shell up to 18.0 mm in width, wider than long (av er - from north-west ern Af rica age width-to-length ra tio 1.62, N = 3), very strongly ventribicon- vex. Max i mal width and thickness at hinge line. Ven tral valve The brachio pod fauna of the Tindouf Syncline (Fig. 1B) high, py rami dal; interarea mod er ately apsacline, high, transver- is es pe cially im por tant for com par i sons with the brachi o pods, sely striate, flat or somewhat con cave. Delthyrium narrow, oc cu- dealt with in the present pa per. Middle Devo nian brachi o- py ing less than one-sev enth of the shell width, closed by a strongly pods, first briefly reported by Menchikoff (1935), Jacquet con vex deltidium. Pedicle fo ra men in the mid dle of the deltidium (1936), and Menchikoff and Monod (1936), were first illus - or closer to the apex. Dor sal valve slightly con vex to nearly flat. trated by Cottreau (1940) on the basis of collec tions, made by Orna menta tion of rounded costae, 4–5 on each flank. Strong Odette du Puigaudeau et Marion Sénones. The best doc u- median dor sal fold with flat tened top, ex cep tion ally with a faint mented fauna was described by Sougy (1964; see also Drot, median groove on it. Ven tral sulcus flat-bot tomed. In te rior not 1961b) from the Zemmour Noir (area at the NW edge of the stud ied. Reguibat Shield, proba bly a northern exten sion of the Mau- Re marks: For a compar i son with Cyrtina ex gr. intermedia, see Re marks on the ge nus. ritanides fold belt, mostly in Mauri ta nia; Lécorché et al., Dis tri bu tion: Boulonnais, Holy Cross Mountains, MaÎder, Eifel?; 1991). Proxim ity of brachio pod faunas from the southern Mid dle De vo nian. MaÎder and the Zemmour Noir may be exem pli fied by co- oc cur rence of Atrypa confusa and Spinatrypa cf. trigonella, two oth er wise rare spe cies, at Aferdou and at the lo cality Cyrtina ex gr. intermedia 2hlert, 1887 346 of Sougy (1964, p. 382–383) in the Zemmour. Species Fig. 36GG–KK lists without either descrip tions or figures were given from ex gr. 1887 Cyrtina heteroclita, var. intermedia 2hl. – 2hlert: the southern flank of the Tindouf Syncline (the northern p. 42, pl. 3, figs 29–35. border of the Reguibat Shield, mostly in Alge ria, partly also ex gr. 1952a Cyrtina heteroclyta var. intermedia Oehlert – Le in Mauri ta nia) by Gevin (1960). The western part of the Maître, p. 132. Tindouf Syncline is more poorly known (Königshof et al., ex gr. 1989 Cyrtina intermedia (Oehlert) – Gourvennec, pp. 2003); a few taxa from this area (Western Sa hara, south ern 56–59, text-figs 28, 30–31, pl. 2, figs 20–36 [ubi Morocco) were reported by Dumestre and Illing (1967; syn.] identi fi ca tions H. Muir-Wood) and by Schemm-Gregory Ma te rial: Two subcomplete shells from Maharch, ZPAL Bp 68/2/ and Jansen (2005, 2008). 38/1–2. De scrip tion: Like the pre vi ous, ex cept: shell strongly trans verse The De vo nian of the Ougarta chain, cropping out in the (av er age width-to-length ra tio 1.89, N = 2); interarea weakly apsa- Oued Saoura valley (western Alge ria; Saoura in Fig. 1B; for cline; costae 3–4 per flank; ven tral sulcus V-shaped. re gional de tails see Alimen et al., 1952; Boumendjel et al., Re marks: For a compar i son with Cyrtina sauvagei, see Remarks 1997a), yielded a rich brachio pod fauna, described by Le on the ge nus. A moder ately trans verse (width to length ra tio 1.5) Maître (1952a) and restud ied by Boumendjel et al. (1997b) MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 297

and Brice et al. (2011). Most of this fauna is Early Devo nian row oce anic zone (McKerrow et al., 2000), against the op- in age; a part belongs to the Middle Devo nian. po site conclu sion (a wide Rheic Ocean: Ziegler, 1990; Plus- There follows an enumer a tion of other Middle Devo - quellec et al., 1997; Tait et al., 2000; Torsvik and Cocks, nian brachio pod faunas from northern and north-western 2011; see also the summary of this conten tion in Be³ka and Af rica. Termier and Termier (1950) de scribed sev eral De - Narkiewicz, 2008, p. 383–384). Secondly, not a single Amer- vo nian taxa from a large number of lo cali ties, scattered ican-type species is present in the southern MaÎder (unless from northern Mo rocco to Mauri ta nia (but appar ently not a Cyrtina sauvagei be conspecific with C. hamiltoni and thus a sin gle Mid dle De vo nian brachi o pod from MaÎder among widely distrib uted species), not even an Appallachian-type them). A few Eifelian to Givetian taxa from the Drâa (Dra) ge nus (sensu Boucot et al., 1969). val ley were de scribed by Becker et al. (2004) and by Ebbig- The same con clu sion was made by Brice (in Boumen- hausen et al. (2007). Givetian brachio pods have also been djel et al., 1997b) for the Lower to Midle Devo nian brachi o- described from the Adrar (Racheboeuf et al., 2001) and pod faunas from the Saoura. It may be noted too that the from the Taoudeni Basin (Racheboeuf et al., 2004). Eifelian anal ysis of latest Emsian brachi o pods from the Cantabrian and/or Givetian brachio pods were described from the Zone showed also great simi lar ity of faunas from northern Tamesna Basin (South Ahaggar Massif) by Mergl and Gondwana and from Euramerica, and therefore minor sig- Massa (2004). Givetian brachio pods from Libya and Niger nif icance of the Variscan Sea as a bar rier to faunistic ex - were described by Boucot et al. (1983). changes (García-Alcalde, 1995, 2001). This is not aston ish - ing, inso far as palaeocurrents are re con structed as hav ing circu lated along the margins of the ocean (Oczlon, 1990; Anal y sis of the fauna stud ied Dopieralska, 2009), thus forming a system that allowed the The palaeobiogeographic anal ysis was con ducted on easy disper sion of benthic faunas. the ba sis of the distri bu tion of 56 out of the 62 spe cies re- Mid dle De vo nian brachi o pod fau nas from the Tindouf ported in the present paper, includ ing 13 taxa described in Syncline have not been un der detailed research. However, open no men cla ture, the re main ing 6 be ing iden ti fied only at gener ally speaking, they are composed partly of the same the ge nus or fam ily level, and therefore not suit able for this species as in the southern MaÎder (14 or 19 common spe- purpose (see Halamski, 2008 for details of method ol ogy; for cies, Ta ble 3). Ameri can af fin ities of these fau nas have been the Otsuka co ef fi cient, see Cheetham and Ha zel, 1969). This invoked, on account of the presence of Tropidoleptus (Ge- mid dle Eifelian to early Givetian fauna was ana lysed as a sin - vin, 1960; Schemm-Gregory and Jansen, 2005) and Amphi- gle unit [i.e., not sepa rat ing the Eifelian and Givetian parts; genia (Gevin, 1960). As a matter of fact, the former genus is see Halamski (2008) for argu ments for this approach]. cosmo pol i tan (Harper, 2007) and the latter widely distrib - First of all, it must be observed that no common species uted (Lee et al., 2006, p. 1996). Three taxa “simi lar to forms were found with the follow ing coeval faunas from north- from eastern North Amer ica” have been re ported from Wes- west ern Af rica: from the Tamesna Basin (Mergl and Massa, tern Sahara (Schemm-Gregory and Jansen, 2005, p. 27): 2004) and from Libya (Boucot et al., 1983; Mergl and Cyrtina cf. hamiltonensis (Hall, 1857), Devonochonetes cf. Massa, 1992). scitulites (Cooper, 1945), and Rhipidomella cf. vanuxemi The dis tri bu tion of the spe cies in ves ti gated in se lected (Hall, 1857). All three are identi fied un der open nomen cla - other regions is summa rised in the Table 5. “Tindouf South” ture, where fore the ac tual ex tent of pre sumed Ameri can in - includes faunas from the Zemmour (Sougy, 1964), those flu ences can not be es ti mated be fore the sys tem atic re vi sion from the southern flank of the Tindouf Syncline proper (Ge- of this ma te rial. vin, 1960), as well as those from the Western Sahara (Du- Racheboeuf (1991) and Racheboeuf et al. (2001, p. mestre and Illing, 1967; Schemm-Gregory and Jansen, 2005, 153) divided the Devo nian of West Africa into two major 2008). The Rhenish Massif includes the Eifel (left or western regions: the Taoudeni Basin (south of the emerged area of side of the Rhine, accord ing to the tradi tional German termi - the Reguibat Shield; Golonka et al., 2006, fig. 20) with fau- nology) and the Sauerland, Bergisches Land, Siegerland, and nas showing “close faunal rela tion ships with the Northeast - Westerwald (right or eastern side of the Rhine) (Schnur, ern Amer ican Realm” (an ob ser va tion al ready made by Le 1853; Kayser, 1871; Torley, 1934; Schmidt, 1941a, b, and Maître, 1952b) and the Zemmour and Tindouf Basins with espe cially numer ous papers by Struve, listed in Weddige and Eu ro pean-type fau nas. This broad sub di vi sion was es tab- Ziegler, 2000). The “Holy Cross Mountains North” refers to lished mainly on the basis of chonetoid brachio pod distri bu - the Middle Devo nian faunas of the Bodzentyn Syncline in tions. It remains to see to what ex tent these conclu sions per- the £ysogóry region of the Holy Cross Mountains (Poland), tain to the entire brachio pod faunas. described by Gürich (1896), Sobolew (1904, 1909), Siemi- The available incom plete data on Middle Devo nian radzki (1909), Biernat (1959, 1964, 1966), and revised by non-chonetoid brachio pods from the Taoudeni Basin do not Halamski (2004a, 2009; see also Halamski, 2005; Halamski support the strong sepa ra tion with more northern faunas, ad- and Racki, 2005). vo cated by Racheboeuf et al. (2001). Affin i ties with the Ap- The main conclu sion of this enumer a tion is the remark - pallachian region were postu lated on the basis of taxa, repor- able sim i lar ity be tween brachi o pod fau nas of the north ern ted under open nomen cla ture. On the contrary, taxa identi fied (Euramerican) and southern (Gondwanan) shores of the at the spe cies level are mainly Euro pean forms: Xystostro- Variscan Sea. Out of the 62 spe cies pres ent in the MaÎder, at phia umbraculum, Schizophoria schnuri (see above), Atrypa least forty are pres ent also ei ther in the Eifel or in the Holy Planatrypa) collega, A. (P.) “squamifera”, and Kyrtatrypa Cross Mountains. This supports the idea of a rela tively nar- culminigera (Copper in Racheboeuf et al., 2001). Simi larly, 298 A. T. HALAMSKI & A. BALIÑSKI

Ta ble 5 Syn op sis of occur rences of taxa pres ent in the studied mate rial in se lected co eval fau nas.

Region Gondwana Euramerica (Laurussia) MaÎder Rhenish Holy Cross Tindouf (S) Other Taxon MadÀne Aferdou Maharch Massif Mts. (N) Deliella deliae + + Craniida Deliella aff. rhenana + + Leptagonia analogaeformis + + + + + Protodouvillina interstrialis + + + + + + Strophomenida Radiomena irregularis + + + + Parastrophonella anaglypha + + + + + Protodouvillininae indet. + Devonaria sp. + Productida Poloniproductus varians + + + + Xystostrophia umbraculum + + + + + + Orthotetida Iridistrophia cf. undifera + + + + Aulacella prisca + + + + + + Tyersella canalicula + + ? + + Orthida Phragmophora schnuri + + + + Schizophoria schnuri + + + + + Schizophoria? sp. + Gypidula biplicata + + + ? Glyptogypa multiplicata + + + + + Pentamerida Ivdelinia pulchra + + + Devonogypa spinulosa + + ? + + + Antirhynchonella sublinguifera + + + + + Kransia parallelepipeda + + + + Kransia subcordiformis + + + + Kransia? coronata + + + + Beckmannia beckmanni + + ? + Rhynchonellida Glosshypothyridina procuboides + + + + Parapugnax? cf. skalensis + cf. Septalaria gracilis + + + ? Paulinaerhynchia paulinae + + ? Rhynchonellida indet. + Atrypa (Planatr.?) confusa + + + + + Atryparia dispersa + + + Invertina cf. struvei + Spinatrypa cf. trigonella + + cf. cf. Spinatrypa globulina + + Desquamatia deserti + Desquamatia microzonata + + + Atrypida Desquamatia circulareformis + + Desquamatia sp. 1 + Kerpina vineta + + Prodavidsonia sp. + Carinatina arimaspus + + + + Gruenewaldtia latilinguis + + + + Peratos arrectus + + + + Plectospira ferita + + + + Athyridida Athyris ex gr. concentrica + ? ? Athyris? aff. curvata + MID DLE DE VO NIAN BRACHI O PODS FROM THE SOUTH ERN MADER (MAROCCO) 299

Table 5 continued

Region Gondwana Euramerica (Laurussia) MaÎder Rhenish Holy Cross Tindouf (S) Other Taxon MadÀne Aferdou Maharch Massif Mts. (N) Camarium sp. + + Meristella cf. iconensis + cf. Athyridida Bifida lepida + + + Kayseria alvea + + Spinocyrtia cf. elburzensis + cf. Ivanothyris aculeata + + + + Quiringites arensentiae + Cyrtinopsis cf. brachyptera + cf. Spiriferida Rhenothyris sinuata + + + Deltospirifer? sp. nov. + Undispirifer aff. rzhonsnitzkajae + aff. Thomasaria? simplex + + + + ? Reticularioidea indet. + Cyrtina sauvagei + ? ? + ? Spiriferinida Cyrtina ex gr. intermedia + ? TOTAL 20 49 11 38 (42) 31 (33) (absolute number of taxa in the 14 (19) – MaÎder and of species in 62 40 (46) common with other regions) Total, for palaeobiogeography 66 (75) % 55 (59) % and percentages of species in 56 26 (35) % – common between MaÎder and a 71 (82) % given region Total, species (in parentheses: 300 120 (75) fit for palaeobiogeographic 62 »70 – analysis) 348 (329) BRACHIOPODA Total, genera (in parentheses: 106 84 (69) fit for palaeobiogeographic 52 (49) – – analysis) 133 (118) Total, genera in common 42 37 (39) between the MaÎder and a given 49 – – region 44 (45) 29 (32) 47 (51) Otsuka coefficient, species (%) 100 – – 29 (34) 58 66 (70) Otsuka coefficient, genera (%) 100 – – 58 (59) Biogeographic data, unless other refer ence is given, after Halamski (2008). Greyed cells denote taxa unfit for palaeobiogeographic analy sis. See text for fur ther ex pla na tion

“close af fin i ties with the NEA Realm” were pos tu lated for af ter sys tem atic re vi sion of these fau nas will have been com- brachi o pods com ing from the Hodh (south ern Mau ri ta nia) on pleted. account of pres ence of “?Arcuaminetes, Longispina, Eleu- A pre lim i nary ob ser va tion, con cern ing links of north- therokomma, and Cupularostrum” (Racheboeuf et al., 2004, west ern Af ri can Mid dle De vo nian brachi o pod fau nas with p. 100); as a matter of fact, the oldest Eleutherokomma is South China, as exem pli fied by the genera Deltospirifer known from the Holy Cross Mountains (Halamski, 2004a) (present only in those two re gions; Carter and Gourvennec, and Cupularostrum is cos mo pol i tan (Sav age et al., 2002, p. 2006 and data presented herein; however, it should be borne 1056). It seems proba ble that faunal ex changes with ter rito - in mind that the iden tifi ca tion of the single spec imen at the ries, sit u ated in the pres ent North America, were more in - disposal of the authors is tenta tive), Invertina (only those two tense south of the Reguibat Shield, owing to the latter and the also; Godefroid, 2000; Copper, 2002) and Paracrothyris Acadian Highlands act ing as barri ers (see fig. 1 in Stigall (present in Western Sahara, South China, and Nevada; Rode and Lieberman, 2005 and refer ences therein). How- Schemm-Gregory and Jansen, 2008), may be made, as well. ever, the extent of Old and New World faunal influ ences A detailed palaeobiogeographic anal ysis of early Give- north and south of the Reguibat Shield may only be esti mated tian rugosan corals from Jebel Driss in the southwest ern 300 A. T. HALAMSKI & A. BALIÑSKI

MaÎder was conducted by Pedder (1999), who concluded thus very simi lar in a bi o log ical sense (same spe cies, not that there were strong simi lar i ties be tween the north and only same genera). In contrast, no Appallachian (Eastern south shores of the Variscan Sea and very weak links with North Ameri can realm) brachi o pod spe cies or gen era were the Appallachian Re gion (Eastern Amer icas Realm). Simi - noted in the study area. larly, May (2008) deduced that there were close biogeogra- 7. Palaeobiogeographic proxim ity of the north ern Gon- phic rela tion ships between the Givetian tabu lates of Mo- dwana and southern Laurussia shores favours palaeogeo- rocco and of central Europe, without any special influ ence graphic recon struc tions with a narrow Variscan Ocean, like of the Eastern Amer icas Realm. A de tailed anal ysis of bra- that of McKerrow et al. (2000). chiopods, crinoids, trilobites, corals, and chitinozoans from the Lower to Middle Devo nian of Saoura (Boumendjel et al., 1997b) showed that the presumed North Ameri can af- Ac knowl edg ments fin ities of this fauna (Le MaÐtre, 1952a) as a matter of fact, The au thors were in tro duced to the ge ol ogy of Morocco by were weak at best. These con clu sions are fully con cor dant Zdzis³aw Be³ka (Adam Mickiewicz Univer sity, Poznañ) dur ing a with those, made on the basis of brachio pods from the fieldtrip he led in Feb ru ary and March 2010. Other par tic i pants south ern MaÎder, studied by the present authors. were B³a¿ej Berkowski (same Univer sity), Janusz Grebowicz, and Ken neth John son (both Univer sity of Houston Downtown). Ulrich Jansen (SMF) kindly al lowed the au thors to study the Mid dle De - CON CLU SIONS vonian part of his col lec tion from MadÀne el Mrakib and granted ac cess to the com par a tive col lec tions. Ma te rial from the MNHN was ex amined through the cour tesy of Didier Merle and Jean- 1. A di verse Middle Devo nian (middle? Eifelian to Michel Pacaud and that from the HLMD through the cour tesy of middle Givetian; mainly Taboumakhloõf Forma tion, Gabriele Gruber and Marisa Blume. The au thors are in debted to subordinately proba bly also the El Otfal Forma tion) brachi - Paul Cop per’s (for merly Lau ren tian Uni ver sity, Sudbury, On tario) opod fauna is present at MadÀne el Mrakib, Aferdou el er u di tion, of which they took ad van tage through the me dium of the Mrakib, and Guelb el Maharch in the southern MaÎder (east - iden ti fi ca tion la bels that he had put in sev eral mu seum col lec tions ern Anti-At las, Mo rocco). Sixty-two taxa (2 rep re sen ta tives all over the world. A number of hard-to-find publi ca tions were of Craniida, 5 of Strophomenida, 2 of Productida, 2 of Or- provided by Denise Brice (Faculté libre des Sciences, Lille), thotetida, 5 of Orthida, 5 of Pentamerida, 9 of Rhyn- François Atrops (Université Claude Ber nard – Lyon 1), and Miko³aj Zapalski (Univer sity of Warsaw). Fieldwork of ATH in chonellida, 14 of Atrypida, 7 of Athyridida, 9 of Spiriferida, Morocco was fi nanced by the Pol ish Minis try of Sci ence and Hig- and 2 of Spiriferinida) are described, on the basis of >1,300 her Ed uca tion (Grant N307 016237 “Bi otic and functional struc- spec i mens. The fauna is dom i nated by rep re sen ta tives of the ture of sub marine vent eco sys tems in the De vonian of Mo rocco” to order Atrypida (14 taxa, about half of the mate rial). B³a¿ej Berkowski). The au thors would like to thank A. Mjid and 2. Strophomena porrecta Maurer, 1885 and Zopho- E. Rjimati (MinistÀre de l’Énergie et des Mines, Morocco) for strophia? aaatos Halamski, 2009 are junior subjec tive syn- work per mits and lo gis tic ad vice. The pa per was re viewed by Ber - onyms of Radiomena irregularis (Roemer, 1844). Septa- nard Mottequin (Université de LiÀge) and Zdzis³aw Be³ka. The laria descendens Schmidt, 1975 is a junior subjec tive syn- Eng lish text was thor oughly re viewed by Frank Simpson (Univer - onym of S. gracilis (Gürich, 1896). sity of Wind sor). All the above-mentioned per sons and in sti tu tions are grate fully ac knowl edged. 3. Paulinaerhynchia is proposed as a new genus of Pugnacidae (Rhynchonellida) with type species P. paulinae gen. et sp. nov. from Maharch. It is closest to Pugnax, from which it differs in its strong costation, lack of sep tum and REF ER ENCES septalium, and ru di men tary den tal plates. Abramân, M. S., 1974. Devonskaâ sistema, brahiopody. In: Ako- 4. Glosshypothyridina Rzhonsnitskaya, 1978, known pân, V. T. (ed.), At las iskopaemoj fauny Armânskoj SSR. up to now from the Eifelian, is reported in the lower Give- Edi tions of the Ar menian SRR Acad emy of Sci ences, Erevan, tian. A ques tion able oc cur rence of Antirhynchonella in the 48–64. [In Rus sian]. lower Givetian is confirmed. Alekseeva, R. E., 1960. O novom podrode Atrypa (Desquamatia) 5. Desquamatia (D.) deserti sp. nov. from Maharch is a subgen. nov. iz sem. Atrypidae Gill (Brahiopody). Doklady new large-sized and finely costate rep re sen tative of the ge - Akademii Nauk SSSR, 131: 421–424. nus. It is clos est to Desquamatia (D.) ovata Copper, 1966 Alimen, H., Le Maître, D., Menchikoff, N., Petter, G. & Poueyto, [1966a] from the Eifelian of the Eifel, from which it differs A., 1952. Les chaînes d’Ougarta et la Saoura. XIXÀme Con- Àre in a more convex ventral valve. grÀs géologique in ter na tional, Monographies régionales,1 6. The Middle Devo nian brachio pod fauna from the Série : Algérie – n° 15. Alger, 114 pp. southern MaÎder is surpris ingly simi lar to co eval fau nas Alvarez, F. & Brime, C., 2000. Type spec i mens of athyridid brachi - from central Europe: out of 56 taxa fit for palaeobiogeogra- o pods from the James Hall col lec tion. The Uni ver sity of Kan - sas, Paleontological Con tri bu tions, New Se ries, 12: 1– 15. phic analy sis (those described in open no mencla ture are Alvarez, F. & Brunton, C. H. C., 2005. On the name-bear ing type mostly excluded), 66% (possi bly up to 75%) of species are of Athyris concentrica (von Buch, 1834). Lethaia, 38: 86–87. com mon with the Eifel and 55% (59%?) of spe cies with the Alvarez, F., Brunton. C. H. C. & Struve, W., 1996. On Athyris northern region of the Holy Cross Moun tains; if both the (Brachiopoda) and its type spe cies ‘Terebratula‘ concentrica latter fau nas are treated as a sin gle unit, the ra tio of species von Buch. Senckenbergiana lethaea, 76: 65–105. in common is as high as 71 (82)%. 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