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Dignomia munsterii (Brachiopoda, Lingulata) from the Ordovician of Bolivia, with redescription of the genus Christian C. EMIG Centre d’Océanologie de Marseille, CNRS-UMR 6540, Station marine d’Endoume, rue de la Batterie-des-Lions, F-13007 Marseille (France) [email protected] Zarela HERRERA Departamento de Ciencias de la Tierra (Paleontología), Facultad de Ciencias, Universidad de Zaragoza, E-50009 Zaragoza (Spain) [email protected] Emig C. C. & Herrera Z. 2006. — Dignomia munsterii (Brachiopoda, Lingulata) from the Or- dovician of Bolivia, with redescription of the genus. Geodiversitas 28 (2) : 227-237. ABSTRACT Specimens of Dignomia have been recorded in the Anzaldo Formation (Up- per Ordovician) in three localities in the Cochabamba area of the Cordillera Oriental, Bolivia. Th ey had been attributed to Lingula munsterii d’Orbigny, 1842 found in company with L. submarginata d’Orbigny, 1850, both are in- dubitable Dignomia. Th ese species were collected in the Cordillera Oriental in the Cochabamba region and in the Chuquisaca area, near Zudáñez the type locality and belong in fact to Dignomia. Dignomia munsterii is described and a new generic diagnosis is proposed. Th e emended diagnosis is based on char- acters of the shell and anatomic traits that permit its recognition as a discrete genus of the Lingulidae, the family to which it belongs. Th e type species is Lingula alveata Hall, 1863. Dignomia is found in strata ranging in age from KEY WORDS Brachiopoda, Lower Silurian to Middle Devonian in North and South America. At least six Lingulidae, previously described species can be referred to Dignomia but all are inadequately Dignomia, Ordovician, described without reference to the scars indicating the sites of attachment of Bolivia. the visceral organs. GEODIVERSITAS • 2006 • 28 (2) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 227 Emig C. C. & Herrera Z. RÉSUMÉ Dignomia munsterii (Brachiopoda, Lingulata) de l’Ordovicien de Bolivie, et redescription du genre. Des exemplaires de Dignomia ont été récoltés dans la Formation Anzaldo (Ordovicien supérieur) dans trois localités de la région de Cochabamba (Bolivie) dans la Cordillère Orientale des Andes. Ils ont été attribués à Lingula munsterii d’Orbigny, 1842 qui appartient sans conteste au genre Dignomia, de même que L. submarginata d’Orbigny, 1850 : ces deux espèces ont été décrites de la région de Cochabamba (Cordillère Orientale) et de celle de Chuquisaca (à Zudáñez qui est la localité type). Une redescription de Dignomia munsterii et une nouvelle diagnose générique sont proposées. Grâce à l’étude des caractères de la coquille et des caractéristiques anatomiques, le genre Dignomia, dont la diagnose a été amendée, a été reclassé dans la famille des Lingulidae et se distingue bien des autres genres de cette famille. L’espèce type est Lingula alveata Hall, 1863. MOTS CLÉS L’extension géologique du genre va du Silurien inférieur au Dévonien moyen Brachiopoda, et la distribution géographique se situe en Amérique du Nord et du Sud. Au Lingulidae, moins six espèces peuvent être rapportées au genre Dignomia, mais toutes ont Dignomia, Ordovicien, été décrites de façon inadéquate car sans référence aux traces donnant la dispo- Bolivie. sition des caractères anatomiques du corps. INTRODUCTION In the last century several French and German expeditions worked in Bolivia, including the impor- Lingulids are the most common inarticulated bra- tant investigations of Dereims (1906) and of Stein- chiopods encountered in the Upper Ordovician mann & Hoek (1912). Th eir collections included of Bolivia. Alcide d’Orbigny (1842) was the fi rst L. munsterii and three new forms: Lingula lineata, to describe them in his exceptional Voyages en L. ellipsiformis and L. boliviana. More recently, Gag- Amérique Méridionale. Of the three new species nier et al. (1986, 1996) recorded Dignomia from of Lingula described by this author (1842: 25-27, three localities in the vicinity of Cochabamba. Emig pl. 2), Lingula submarginata d’Orbigny, 1850 (pro in Gagnier et al. (1996) described them briefl y and Lingula marginata d’Orbigny, 1842, not Phillips, referred them to a new species of Dignomia. Recently, 1836) and L. munsterii, collected from Zudáñez Aceñolaza et al. (2003) reviewed the Ordovician (= Tacopaya) in the Chuquisaca region and from lingulid shell beds in South America. Palta-cueva in the Cochabamba region belong with certainty to the genus Dignomia (by the way, the genus Lingula arose only in the Tertiary, or MATERIAL perhaps in the Late Cretaceous [Biernat & Emig 1993; Emig 2003]).Th e syntypes of L. munsterii Th e fossil specimens were collected in Bolivia in are deposited in the d’Orbigny collection in the the Cochabamba region from the Anzaldo Forma- Muséum national d’Histoire naturelle in Paris. tion (sensu Rodrigo de Walker & Toro 1987) at the D’Orbigny never fi xed a holotype. No specimen following localities (Figs 1; 2): Cerro San Pedro referred to L. submarginata (or to L. marginata) in eastern suburb of Cochabamba Ciudad; Cerro could be found in d’Orbigny’s collection. Chakeri in the north of the pueblo of Sacabambilla, 228 GEODIVERSITAS • 2006 • 28 (2) Ordovician Dignomia (Brachiopoda) from Bolivia FIG. 1. — Maps of Bolivia showing the distribution of the Ordovician (from Pareja & Ballon 1978). about 50 km east of Cochabamba; Sacabamba ties where Dignomia has been recorded are in the in the Rio Challaque, about 60 km south of the Anzaldo Formation (also called the Cuchupunata town of Cochabamba. Th irty more or less com- Formation by many Bolivian geologists) which plete valves of lingulides from these localities were may attain a thickness of 2000 m (Fig. 2). In the studied (Figs 1; 3). Cochabamba area, Steinmann & Hoek (1912) Th e specimens described in this paper are housed in diff erentiated three units based mainly on fauna, the “Typothèque des Invertébrés fossiles du Muséum from bottom to top (Fig. 2): “Bilobites Sandstein”, national d’Histoire naturelle”: MNHN R06654 “Lingula Sandstein”, and “oberer Quarzit”. Th e latter – the syntypes of Lingula munsterii (a dorsal and has been named the San Benito Formation, while a ventral valve) from Zudáñez (= Tacopaya), Chu- the two lower levels have been combined as the quisaca region; MNHN A24313 – Cerro San Pedro Anzaldo Formation. Ordovician rocks in this region in Cochabamba (17°23’45”S, 66°07’40”W); MNHN form the Cochabamba Group (Fig. 2) comprised A24314 – Cerro Chakeri, Puntata Province, north of the Independencia Formation, the Capinota of Sacabambilla (17°30’00”S, 66°47’58”W). A large Formation, the Anzaldo Formation and the San collection from Cerro San Pedro in Cochabamba Benito Formation. Th e Independencia Formation Ciudad is also housed in the Museo de la Univer- is dated Arenigian, the Capinota and the lower part sidad de Zaragoza. of the Anzaldo Formation are Llanvirnian-Llandeil- ian, and upper portion of the Anzaldo Formation and the San Benito Formation are of Caradocian GEOLOGICAL SETTING AND AGE age. Th e precise age of the upper portion of the Anzaldo Formation remains under debate. Súarez- In Bolivia, the most nearly complete sequence of Soruco (1976) considers the Dignomia localities to the Ordovician is in the Cordillera Oriental (Mar- be Caradocian, but Rodrigo de Walker & M. Toro tínez et al. 1971; Gagnier et al. 1996). Outcrops (pers. comm.) argue for a Llanvirnian-Llandeilian extend from the Peruvian border in the northeast age based on trilobites. Gagnier et al. (1996) report to Argentina in the South (Fig. 1). Th e three locali- that vertebrates from these localities are similar to GEODIVERSITAS • 2006 • 28 (2) 229 Emig C. C. & Herrera Z. of argillaceous sandstone separates each fossiliferous sandstone level. In Cerro Chakeri, Dignomia oc- curs in thin beds (0.5-1 cm thick) of fi ne to coarse sandstones, consisting exclusively of fl at- lying valve fragments. In addition, some complete, rather well preserved, large, solitary valves were collected in a layer of coarse sandstone. Below and above the lingulid beds, there occur many of the vertical cylin- drical burrows which may be interpreted as traces of lingulid burrows. Th e three localities studied include several phosphorite levels, composed principally of crushed Dignomia valves, but containing some small vertebrate fragments. Th ese large concentrations of lingulid valves have been referred (see Súarez- Soruco 1976) to Lingula ellipsiformis Hoek, 1912, L. munsterii, Bistramia elegans Hoek, 1912; other invertebrates have been identifi ed, among them the trilobite Huemacaspis bistrami (Hoek, 1912) and the vertebrate Sacabambaspis janvieri Gagnier, 1987 FIG. 2. — Stratigraphical sequence of the Ordovician of the central (see Gagnier et al. 1986; Toro et al. 1994). Cordillera Oriental with the position of the studied Dignomia beds indicated in deep grey. Th e invertebrate fauna represents a marine, in- tertidal or subtidal benthic assemblage. In light of recent ecological investigations of living Lingula early Llanvirnian forms from Australia (Ritchie & Bruguiére, 1797 (cf. Emig 1986, 1997), fossiliza- Gilbert-Tomlinson 1977), and therefore the two tion of fl at-lying valves may occur preferentially localities must be contemporaneous. Th e latest pal- when the environment changes drastically, e.g., a aeontological data (Gagnier et al. 1996) regarding large decrease in salinity, the eff ects of storms and the Anzaldo Formation suggest a Caradocian age subaqueous slumps both of which produce thick (or perhaps older) because the underlying Capinota deposits of coarse or fi ne sediment. Th e absence of Formation is of Llanvirnian-Llandeilian age and the fossil lingulid shells in other sandstone beds does overlying San Benito Formation is also Caradocian not preclude the presence in them of an original in age (Fig. 2). population of Dignomia because in normal environ- Lingulid specimens from Rio Challaque (Sacabam- ments the shell of the lingulides is not preserved ba) were collected in the basal part of the stratigraphic (Emig 1990).