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Early Middle Cambrian Representatives of the Superfamily Acrotretoidea (Brachiopoda) from Morocco

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Early Middle Cambrian Representatives of the Superfamily Acrotretoidea (Brachiopoda) from Morocco Z. dt. geol. Ges. 15011 p. 27-87, 26 fig., 9 tab., 6 pt. Stuttgart, Mai 1999 Early Middle Cambrian representatives of the superfamily Acrotretoidea (Brachiopoda) from Morocco MICHAEL STRENG* STRENG, M. (1999): Early Middle Cambrian repre­ ist bislang von Vorkommen außerhalb Marokkos sentatives of the superfamily Acrotretoidea (Bra­ bekannt. Zwei Arten, die nur durch jeweils ein chiopoda) from Morocco. [Frühe mittelkambri­ Exemplar repräsentiert sind, werden zusätzlich in­ sche Vertreter der Überfamilie Acrotretoidea formell beschrieben. Die verschiedenen Arten (Brachiopoda) aus Marokko.] - Z. dt. geol. Ges., gehören fünf Gattungen an, von denen die Cera­ 150: 27-87; Stuttgart. tretiden Acanthatreta, Almohadella und Mono­ phthalma und der Acrotretide Tingitanella neu Abstract: Acrotretoid brachiopods of the families sind. Die Gattung Vandalotreta wird emendiert. Im Acrotretidae and Ceratretidae from the early Gegensatz zu den untersuchten Acrotretidae wei­ Middle Cambrian of Morocco are described. The sen die ceratretiden Arten eine hohe Variabilität in fauna comprises seven species, five of which are den gemessenen Größenproportionen auf. Funkti­ new and two species are informally described. The onsmorphologische Untersuchungen an der Schale species are assigned to five genera of which the von Acanthatreta lassen eine primitive Artikulation ceratretid genera Monophthalma, Acanthatreta and der beiden Klappen vermuten. Die Schalenstruktur Almohadella, and the acrotretid Tingitanella are eines Teils der beschriebenen Taxa wurde unter­ newly introduced. The acrotretid genus Vandalotre­ sucht. Die acrotretide Gattung Vandalotreta besitzt ta is emended. In contrast to measurements on the eine zweilagige Schale. Im Gegensatz dazu haben Acrotretidae, those for most of the Ceratretidae die ceratretiden Gattungen Acanthatreta, Almoha­ show a wide variation of proportions. Functional della und Monophthalma eine typisch dreilagige morphologic observations on the shell of Acantha­ Schale. Bohrlöcher, die offenbar von räuberischen treta suggest a primitive hinge articulation, a Organismen erzeugt wurden, wurden auf zahlrei­ characteristic of many acrotretoid genera. The shell chen Klappen der untersuchten Stücke gefunden. structure of some of the Moroccan specimens is Insgesamt konnten fünf verschiedene Typen von studied and described. The acrotretid genus Vanda­ Bohrlöchern unterschieden werden. Die phyloge­ lotreta has a shell that consists of two layers in con­ netische Entwicklung der kambrischen Acrotre­ trast to the ceratretid genera Acanthatreta, Almoha­ toidea wurde unter besonderer Berücksichtigung della, and MOl1ophthalma, which are characterized der marokkanischen Gattungen und 12 weiterer by a three-Iayered shell. Borings have been ob­ kambrischer Gattungen untersucht. Die gewonne­ served in several specimens and are interpreted as nen Kladogramme lassen eine Unterteilung der Fa­ to be of predatory origin. Distinguishable are five milie Ceratretidae in zwei Gruppen vermuten, die different types of boreholes. The phylogeny of the als Unterfamilien interpretiert werden. Cambrian acrotretoid brachiopods is reevaluated based on special consideration of the Moroccan Keywords: Brachiopoda (Acrotretidae, Ceratreti­ species and 12 additional Cambrian genera. De­ dae), Middle Cambrian, classification, new taxa, rived cladograms suggest a subdivision of the Cera­ shell structure, biometry, borings, phylogeny, cladi­ tretidae into two subfamilies. stic analyses. Kurzfassung: Vertreter der Superfamilie Acro­ High Atlas, Antiatlas, Morocco. tretoidea (Brachiopoda) aus mittelkambrischen Schichten Marokkos werden beschrieben. Von den Contents insgesamt sieben Arten sind fünf neu und nur eine 1. Introduction ............................ 28 *Address of the author: Michael STRENG, Univer­ 2. Cambrian lithostratigraphy and biostrati­ sität Bremen, Fb-5 Geowissenschaften, Postfach graphy of the Souss Basin of southern 330440, D-28334 B'remen, Germany. Morocco ............................... 29 0012-0189/99/0150-0027 $ 15.25 © 1999 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart 28 M,Streng 1 AIlMersld 2 Akka Tzem 3 Aqdz 4 Assermo 5 Ber1<lk 6 EI Bolj 7 EI Hmam 8 Foum Tangarfa 9 Hassl Brahlm llrane d'Anti-Atlas l11ghels 121joukak 13 IIsiar I Tachguell 14 Jbel Amouri 15 Jbel Assadam 16 Jbel bou Iferslkl 17 Jbal Tirounbar 18lemdad 19 Qunein 20 Ourika Wawrmas 21 Tagragra 22 Tamouchl 23 TaleIl 24 Tin Ouagour 25 Tinifift lJ'l1l Areas presumed or known 10 Include h, Neoprolerozolc (?)-Cambrian rock G.::iJ Older Neoprolerozolc rock .nd +' crystalline basemenl o 100 km Fig, 1: Schematic geological map of the High Atlas and Anti-Atlas regions, Morocco, showing sampled localities and distribution of Late Proterozoic and Cambrian rocks, Modified after GEYER (1993, fig, 1), Abb, 1: Geologische Übersichtskarte des Hohen Atlas und des Anti-Atlas, Marokko, Dargestellt sind die Lokationen der beprobten Schichten und die Ausbisse spätproterozoischer und kambrischer Serien, Verän• dert nach GEYER (1993, fig, 1). 3. Methods ............................... 29 biozone ) and from probably basal Upper Cam­ 4. Systematic paleontology ................. 30 brian layers. MERGL (1983) reported billing­ 5. Quantitative analyses .................... 63 sellid and protorthid genera from the Middle 6. Shell structure .......................... 67 7. Evidence for predation from boreholes ..... 70 Cambrian of the Central Anti-Atlas and the 8. Phylogenetic analysis .................... 76 High Atlas, respectively. Obolellide brachiopods, 9. Acknowledgements ..................... 80 an abundant element of the Lower and Middle 10. References ............................. 81 Cambrian of Morocco have recently been 11. Appendices ............................ 85 described by GEYER (1994) and GEYER & MERGL (1995). 1. Introduction Lingulate brachiopods (sensu WILLIAMS et al. 1996) from the Cambrian of Morocco have Although brachiopods are a relatively common only been dealt with by MERGL (1988), who part of the Cambrian fauna of Morocco, they introduced the acrotretid Vandalotreta vavra have been neglected widely. The first record of and the botsfordiid Botsfordia epigona, and Moroccan Cambrian brachiopods was made by additionally mentioned an obolide brachiopod, TERMIER & TERMIER (1950) who described which was tentatively assigned to Lingulella sp. "Eoorthis Romingeri BARRANDE" from Cambrian The genus Vandalotreta MERGL 1988, described strata of the Moroccan Meseta. Further prot­ herein in detail, has been restricted to Morocco orthacean brachiopods have been described by until recently HOLMER et al. (1996), suggested HAVLICEK (1971) and GEYER & MERGL (1997) Lllhotreta MERGL & SLEHOFERovA 1990, from from the medial Middle Cambrian (Badulesia Bohernia to be a junior synonym (see Discus- Acrotretoidea (Brachiopoda) from Morocco 29 sion under Vandalotreta below). HOLMER & Akka Formation USHATINSKAYA (1994) also mentioned Vandalo­ Goulimine Formation Jbel Afraou Formation treta? sp. from glacial erratic boulders of pro­ bably late Early Cambrian age from Sweden. Tarhoucht Member HINZ-SCHALLREUTER (1997: pI. 1, fig. 17-19) Jbel Wawrmast Formation Breche a Mfcmacca figured an undetermined acrotretid from the ? Member Middle Cambrian of Bornholm which appears to be c10sely related to Vandalotreta. Further Cambrian lingulate brachiopods of western Africa are documented by POULSEN (1960) from the Zemmour region of northern Tislit Issafen Formation Formation Mauritania. The described fauna, which has been tentatively assigned to Early Cambrian Pig. 2: Lithostratigraphic units of the Lower-Middle age, yielded specimens distinguished as Neobo­ Cambrian transition of Morocco showing subdivi­ lus cf. N warthi WAAGEN 1885, Lingulella cf. L. sion of the upper Tata Group through the Feijas fuchsi REDLICH 1899, Botsfordia paucigranulata internes Group. After HELDMAIER & LANDING (in POULSEN 1960, Acrothele sougyi POULSEN 1960, press). Abb. 2: Lithostratigraphische Einheiten im Bereich and Acrothele spinulosa POULSEN 1960. des Übergangs vom Unter- zum Mittelkambrium. However, besides Vandalotreta no repre­ Nach HELDMAlER & LANDING (im Druck). sentative of the superfamily Acrotretoidea, has been recorded from Morocco so far, although they are an abundant element of the early Middle Cambrian fauna of Morocco. themselves are split into several formations (see GEYER & LANDING 1995; HELDMAIER & LANDING, in press). 2. Cambrian lithostratigraphy and The material described herein comes, for the biostratigraphy 01 the Souss Basin most part, from the Jbel Wawrmast and Jbel Afraou Formations (Feijas internes Group), 01 southern Morocco except two sampies from the Tatelt Formation The Cambrian lithostratigraphic framework of (Tata Group). the Anti-Atlas and High Atlas, Morocco, based For lithologic description, fossil re cord, and initially on investigations of Georges Choubert depositional environments of the various (e. g., CHOUBERT 1952; CHOUBERT 1953) and formations, as well as synonymy see GEYER & Henri Hollard (e. g., HOLLARD 1985) for the LANDING (1995), HELDMAIER & LANDING (in Lower Cambrian, and of DESTOMBES (1985) for press) and HELDMAIER (1997). the Middle to Upper Cambrian, has been sub­ sequently precised and c1arified by GEYER (1989, 1990a, 1990b), GEYER & LANDING 3. Methods (1995), and HELDMAIER & LANDING (in press). Biostratigraphic units based on trilobites were The material described herein was mainly first established by HupE (1953) and have been collected by G. Geyer (since 1981) and
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