Lower Ovetian (Lower Cambrian) trilobites and biostratigraphy of the Pedroche Formation (Sierra de...

Original article

Lower Ovetian (Lower Cambrian) trilobites and biostratigraphyof the Pedroche Formation (Sierra de Córdoba, southern Spain)

Trilobites et biostratigraphie de l’Ovétien inférieur (Cambrien inférieur)de la Formation Pedroche (Sierra de Córdoba, sud de l’Espagne)

Eladio Liñán a,*, M Eugenia Dies a, José Antonio Gámez Vintaned a, Rodolfo Gozalo b,Eduardo Mayoral c, Fernando Muñiz c

a Departamento de Ciencias de la Tierra, Museo de Paleontología, Facultad de Ciencias, Universidad de Zaragoza, 50009 Zaragoza, Spainb Departamento de Geología, Facultad de Ciencias Biológicas, Universitat de València, 46100 Burjassot, Spain

c Departamento de Geología, Facultad de Ciencias Experimentales, Campus de El Carmen, Universidad de Huelva, Avda. de las Fuerzas Armadas, s/n.,21819 Huelva, Spain

Received 17 June 2003; accepted 28 November 2003

Abstract

The low Lower Cambrian rocks from the Sierra de Córdoba, which consist of well exposed mixed facies and abundant fossil assemblagesshowing long stratigraphic ranges throughout the Pedroche Formation, represent one of the best successions of this age in Europe. The fossilassemblages include diverse Ovetian archaeocyaths, trilobites, small shelly fossils, calcimicrobia, trace fossils and stromatolites. Trilobitesare still poorly known, and thus they are the main objective of this work. The trilobites studied originate from three sections. At the Arroyo dePedroche 1 section, cf. Bigotinella and Bigotina bivallata are replaced towards the top by Lemdadella linaresae, Lemdadella perejoni sp. nov.and, finally, by Eoredlichia cf. ovetensis. At the Arroyo de Pedroche 2 section, Lemdadella linaresae is replaced by Lemdadella perejoni sp.nov. and Eoredlichia cf. ovetensis, while at the Puente de Hierro section Lemdadella linaresae, L. aff. linaresae and Serrania verae occurtogether. These new biostratigraphic data confirm that the Pedroche Formation, originally defined as a repetitive sequence of four members,contains only two members. The new trilobite discoveries permit the first tentative correlation between the Ovetian of southern Spain andLower Cambrian strata from the High Atlas (Morocco), Siberia, Antarctica and Carteret (France).© 2005 Elsevier SAS. All rights reserved.

Résumé

La succession du Cambrien inférieur basal dans la Sierra de Córdoba est l’une des meilleures en Europe ; elle se compose de faciès mixtesbien exposées et d’abondants assemblages de fossiles qui montrent de longues distributions stratigraphiques dans la Formation Pedroche. Cesassemblages sont constitués d’archaeocyathes, de trilobites, de Small Shelly Fossils, de calcimicrobes, de traces fossiles et de stromatolites del’Ovétien. Les trilobites sont encore très peu connus et sont le principal objectif de ce travail. Les trilobites étudiés proviennent de trois coupesdifférentes. Dans la coupe d’Arroyo de Pedroche 1, cf. Bigotinella et Bigotina bivallata sont remplacés vers le toit par Lemdadella linaresae,Lemdadella perejoni sp. nov. et, finalement, par Eoredlichia cf. ovetensis. Dans la coupe d’Arroyo de Pedroche 2, Lemdadella linaresae estremplacé par Lemdadella perejoni sp. nov. et ensuite par Eoredlichia cf. ovetensis ; dans la coupe de Puente de Hierro, Lemdadella linaresae,Lemdadella aff. linaresae et Serrania verae coexistent. Ces nouvelles données stratigraphiques confirment que la Formation Pedroche, origi-nalement définie comme une séquence répétitive de quatre membres, en possède seulement deux. Les nouvelles découvertes de trilobitespermettent pour la première fois d’établir une possible corrélation entre l’Ovétien du sud de l’Espagne et des strates du Cambrien inférieur del’Anti-Atlas (Maroc), de Sibérie, de l’Antarctique et de Normandie (Carteret, France).© 2005 Elsevier SAS. All rights reserved.

* Corresponding author.E-mail address: [email protected] (E. Liñán).

Geobios 38 (2005) 365–381

http://france.elsevier.com/direct/GEOBIO/

0016-6995/$ - see front matter © 2005 Elsevier SAS. All rights reserved.doi:10.1016/j.geobios.2003.11.007

Keywords: Spain; Lower Cambrian; Ovetian; Biochronology; Trilobites

Mots clés : Espagne ; Cambrien inférieur ; Ovétien ; Biochronologie ; Trilobites

1. Introduction

The Lower Cambrian rocks of the Sierra de Córdoba werefirst recognised by Hernández Pacheco (1907) through thediscovery of archaeocyaths from Cerro de las Ermitas withinthe lower part of the Pedroche Formation. During the follow-ing years new fossiliferous localities were found and studiedproviding not only archaeocyaths (Hernández Pacheco, 1918;Hernández Sampelayo, 1933; Simon, 1939; Cabanás, 1964;Liñán and Dabrio, 1974; Perejón, 1975a, 1975b, 1975c,1976a, 1976b, 1977, 1989; Zamarreño and Debrenne, 1977;Liñán et al., 1982; Moreno-Eiris et al., 1995) but also crusta-ceans (Richter and Richter, 1927; Liñán Guijarro, 1978), stro-matolites (Schmitt, 1983), small shelly fossils (Liñán Guijarro,1978; Fernández Remolar, 1996, 1999), brachiopods (Liñánand Mergl, 1984), trilobites (Liñán Guijarro, 1978; Liñán andGámez-Vintaned, 1993) and trace fossils (Cabanás, 1966;Fedonkin et al., 1985). Sampling for acritarchs was per-formed by Palacios (Liñán and Palacios, 1983) and also inthe sections studied herein (Palacios, oral com.) with nega-tive results. Because of the alternation of numerous levels ofarchaeocyaths and trilobites (Liñán et al., 1982), the Sierrade Córdoba is considered as a classic area for biostratigraphi-cal studies on the low Lower Cambrian.

The aim of this work is to study the trilobites of Member Iof the Pedroche Formation at three selected sections (Fig. 1),in order to learn new aspects about the biochronology of thelower Ovetian substage and to use these new data for inter-national correlations.

2. Geological setting

The Cambrian rocks of the Sierra de Córdoba follow asyncline trending NW–SE and dipping to the SE. The north-ern flank of this syncline is interrupted by the Cerro Murianofault while the southern part connects to an anticline whosecore is made up of Precambrian rocks of the Volcanic–Sedimentary Complex. This structure is covered by scarceTriassic outcrops, widespread Miocene materials and Qua-ternary terraces of the Guadalquivir river (Fig. 1). The stratig-raphy of the Sierra de Córdoba was established by LiñánGuijarro (1974,1978) in the southern flank of this synclinewhere the rocks are well exposed (Fig. 2). The Volcanic–Sedimentary Complex is the base of the sequence. Its higherlevels (San Jerónimo Formation) are made up of andesites,shales, sandstones and conglomerates (Pin et al., 2002) con-taining late Vendian acritarchs and trace fossils (Liñán andPalacios, 1983; Fedonkin et al., 1985). It is unconformablyoverlain by the Torreárboles Formation which is composedof conglomerate, sandstone, arkose and shale containing three

Corduban ichnoassemblages (Fedonkin et al., 1985). The firstpresents Skolithos ichnosp., Planolites ichnosp. and Cochli-chnus ichnosp. while the second contains Phycodes pedumSeilacher, 1955, Phycodes palmatum (Hall, 1852), Plano-lites cf. P. beverleyensis (Billings, 1862), Treptichnus ich-nosp.; all suggesting a lower Corduban age. The third ich-noassemblage is characterised by the first record ofRusophycus ichnosp., accompanied by Bilinichnus ichnosp.and the soft-bodied metazoan Tiernavia tiernae Fedonkin inFedonkin et al. (1985), indicative of an upper Corduban age.

The Torreárboles Formation is conformably overlain bythe Pedroche Formation. The latter is composed of limestoneand shale, with scarce sandstone and dolostone, and containsarchaeocyaths, trilobites (aff. Bigotinella, Bigotina, Lem-dadella, Serrania, Eoredlichia and Neoredlichiidae), brachio-pods (Paterina), algae, calcimicrobes, small shelly fossils,bradoriids, hyolithids, stromatolites (Charaulachia corduben-sis Schmitt, 1983 and Vetella cf. safartiae Schmitt, 1979, bothdefined at the Arroyo de Pedroche 1 section) and trace fossils(Bergaueria, Cochlichnus, Dactyloidites, Monocraterion,Palaeophycus, Phycodes, Planolites, Psammichnites, Ruso-phycus, Torrowangea, Treptichnus and Skolithos). Archaeo-cyath localities are numerous but the most important are LasErmitas and Arroyo de Pedroche. These palaeontologicalassemblages suggest an early Ovetian age for the PedrocheFormation.

The Santo Domingo Formation lies above the PedrocheFormation and is composed of red shale, dolostone and chert-bearing limestone. Only small shelly fossils, stromatolites(Stratifera), calcimicrobes (Renalcis) and brachiopods havebeen reported from the Orive Member (García-Hernández andLiñán, 1983). The Castellar Formation lies conformably abovethe Santo Domingo Formation and is composed of sandstoneand conglomerate with trace fossils. The Los Villares Forma-tion comprises sandstone and interbedded siltstone. TheMiddle Cambrian trilobites Agraulos cf. arenosus Sdzuy,1968, Badulesia tenera (Hartt in Dawson, 1868), Badulesiagranieri (Thoral, 1935), Bailiella sp., Conocoryphe (Para-bailiella) aff. matutina Sdzuy, 1968, Kymataspis bernardi(Liñán Guijarro, 1978), Jincella sp., Hydrocephalus cf.donayrei Liñán and Gozalo, 1986, Eccaparadoxides sdzuyiLiñán Guijarro, 1978, Eccaparadoxides cf. pusillus Bar-rande, 1846, Peronopsella prokovskajae Sdzuy, 1968, Condy-lopyge cf. rex (Barrande, 1846), Dolichometopus sp., Para-solenopleura aculeata (Angelin, 1851) and Skreiaspis sp.occur at the lower part of the Los Villares Formation, sug-gesting an age spanning from Leonian (Eccaparadoxidessdzuyi Zone) through Caesaraugustan (Badulesia granieriZone) (Liñán Guijarro, 1978; Liñán et al., 1995); the tracefossil Cylindrichnus concentricus Toots in Howard, 1966 andorganic-walled microfossils also occur.

366 E. Liñán et al. / Geobios 38 (2005) 365–381

3. Stratigraphy

The fossils studied belong to three sections located in mem-ber I of the Pedroche Formation (Figs. 3–5).

The Puente de Hierro (PH) section (Fig. 3) is the classiclocality where the first early Lower Cambrian trilobites ofSierra Morena were discovered (Liñán Guijarro, 1974, 1978).The palaeontological site is located in the intersection betweenthe Arroyo de Pedroche (Pedroche stream) and the bridge ofthe Córdoba-Cerro Muriano railway. The fossiliferous bedsapparently overly member II of the Pedroche Formation, butare separated from it by a space covered with alluvial mate-rials (Liñán Guijarro, 1974: p. 18, Fig. 2). That author con-

sidered the hypothetical existence of a stratigraphical repeti-tion of member I and II lithologies in the sequence, assigningthe levels containing the trilobites to member III of thePedroche Formation. In the present work, the discovery ofnew levels of trilobites within member I of the Pedroche For-mation (AP1 and AP2 sections) proves that the beds of thePuente de Hierro represent a tectonic repetition of member Iin the succession and thus the Pedroche Formation only con-sists of two stratigraphic members (Fig. 2).

The Arroyo de Pedroche 1 (AP1) section (Fig. 4) wasstudied by Liñán and Dabrio (1974); Liñán et al. (1982);Fernández-Remolar (1996, 1999). It is the type section of thePedroche Formation. The section begins in the junction

Fig. 1. Geological setting of the studied sections (after Liñán Guijarro in Perejón et al., 1996).Fig. 1. Situation géologique des coupes étudiés (d’après Liñán Guijarro in Perejón et al., 1996).

367E. Liñán et al. / Geobios 38 (2005) 365–381

Fig. 2. Cambrian stratigraphy of the Sierra de Córdoba (after Liñán Guijarro in Perejón et al., 1996).Fig. 2. Stratigraphie du Cambrien dans la Sierra de Córdoba (d’après Liñán Guijarro in Perejón et al., 1996).


between the new Badajoz-Córdoba road and the right side ofthe Arroyo Pedroche and follows along the abandonedCórdoba-Cerro Muriano road, which is located at the left sideof the Arroyo de Pedroche.

A new section (Fig. 5) called Arroyo de Pedroche 2 (AP2)is located near the Club Asland Córdoba and runs parallel tothe Arroyo de Pedroche 1 section.

4. Systematic palaeontology

4.1. Trilobita

The material studied herein is housed in the Museo Pale-ontológico de la Universidad de Zaragoza-Gobierno deAragón at Zaragoza, Spain, under references MPZ 2002/74 toMPZ 2002/708.

Order REDLICHIIDA Richter, 1932.Suborder REDLICHIINA Richter, 1932.Superfamily REDLICHIOIDEA Poulsen, 1927.Family BIGOTINIDAE Hupé, 1953.(nom. Transl. BIGOTININAE Hupé, 1953; emend Geyer,

1990b).

The subfamily Bigotininae was defined by Hupé (1953)who included only the genus Bigotina within it. Geyer (1990b)included five more genera (Bigotinops Hupé, 1953; Ouijja-nia Hupé, 1953; Pruvostina Hupé, 1953; Bigotinella Su-vorova, 1960; Hupetina Sdzuy, 1978) and considered thisgroup as a family.

Pillola (1993) revised this family on the basis of new mate-rial (including Serrania Liñán Guijarro, 1978), gave a diag-nosis and remarked on the problem of its assignment to aconcrete family and superfamily during the second half ofthe XX century by several authors (Hupé, 1953; Hen-ningsmoen, 1959; Sdzuy, 1959, 1961; Suvorova, 1960;Repina, 1960, 1966, 1969; Rasetti, 1972; Öpik, 1975; Geyer,1990b).

In our opinion, the fact that the material found is com-posed mainly of cranidia, makes an accurate taxonomic deter-mination at the level of family difficult for several of theincluded genera. For example, Bigotina and the redlichiidgenus Lemdadella Sdzuy, 1978 have very similar librigenae,thorax and pygidium, suggesting a common ancestor, whilethe librigenae of Serrania and Hupetina have significantlyobtuse genal angles and genal spines shorter than Bigotina.We have also provisionally included Bigotinella in this fam-ily due to the particular exoskeletal sculpture.

Genus Bigotinella Suvorova, 1960.Type species: Bigotina (Bigotinella) malykanica Suvorova,

1960.Discussion: Bigotinella was originally included by Su-

vorova (1960) as a subgenus of Bigotina, but Geyer (1990b)treated this taxon as a genus of the family Bigotinidae. Pillola(1993: p. 866) considered Bigotinella as a valid genus char-acterised by facial lines, a plectrum, very short anteriorbranches of facial sutures subparallel to the axis, and a sculp-ture of fine punctae of equal size distributed all over thecranidium, an opinion which we share. This author includedBigotina (Bigotina) angulata Suvorova, 1960, as a junior syn-onym of Bigotinella malykanica which differs mainly in theangulate anterior border. Close to Bigotina (Bigotinella) angu-lata is Bigotina (Bigotina) inornata Jegorova in Varlamovand Jegorova, 1986 which has a sculpture of small craters buthas no plectrum (smooth elevation in some specimens), norpreglabellar field. It has an elevated and arched anterior bor-der. Bigotinops privus Suvorova, 1960 also has a similar sculp-ture and no plectrum.

B. (Bigotinella) botomica Repina, 1966 has neither plec-trum nor facial lines while Bigotina (Bigotinella) rara Repinain Repina and Luchinina, 1981 has no facial lines and a sculp-ture of fine punctae which is more typical of Bigotina. Thisdiscussion is evidence of the poor knowledge of this genus,which includes species represented by a few incompletecranidia that probably should be not be included in the Fam-ily Bigotinidae because of their different exoskeletal sculp-ture.

cf. Bigotinella sp.Fig. 6(1, 2).

Fig. 3. Puente de Hierro (PH) section (after Liñán Guijarro, 1978).Fig. 3. Coupe de Puente de Hierro (PH) (d’après Liñán Guijarro, 1978).


Fig. 4. Arroyo de Pedroche 1 (AP1) section (after Liñán and Dabrio, 1974).Fig. 4. Coupe de Arroyo de Pedroche 1 (AP1) (d’après Liñán et Dabrio, 1974).


Material: One incomplete cranidium (MPZ 2002/74) andanother juvenile (MPZ 2002/75) preserved as the originalcarapace in blue-yellowish microbial limestones.

Description: Cranidium with a sculpture of fine craterssimilar in size but not equal, distributed all over its surface.When the craters are eroded they seem to be pits (Fig. 6(1)).Arcuate anterior margin slightly pointed in front of the gla-bella (Fig. 6(2)). Non-very homogeneous anterior border,more marked and slightly narrower exsagittally. Marked ante-rior border furrow. Depressed and wide (sag.) preglabellarfield. Strongly convex glabella, well delimited, subcylindri-cal in shape, with three pairs of marked non-transglabellarfurrows. Frontal lobe almost semicircular in shape. Glabellarlobes L1 to L3 similar in width (sag.). Wide (sag.) and deepoccipital furrow. Wide (sag.) occipital ring subtriangular inshape. Depressed preocular field at a lower level than thepalpebral field. Arcuate ocular ridge parallel to the anteriorborder, wider (sag.) towards the sagittal axis joining the gla-bella as wide as its frontal lobe. Long palpebral lobe, difficultto distinguish from the ocular ridge, starting in a constrictionplaced at the same exsagittal position as the beginning of theanterior branches of the facial suture. It then follows the samecurvature as the ocular ridge and finishes in the posterior bor-der furrow. Deep palpebral furrow. Convex palpebral fieldsubtriangular in shape. Deep and wide (sag.) posterior borderfurrow. The “S” parameter of the facial sutures (LiñánGuijarro, 1978: p. 158, Fig. 3(6)) is secant and slightly con-vergent. Straight anterior and posterior branches of the facialsuture, similar in length.Anterior branches of the facial sutureslightly divergent (Fig. 6(2)). Curved and long medialbranches of the facial suture.

Discussion: The slightly convex anterior margin and themarked anterior border of the cranidium which is placed at alower level than the glabella, brings our material near to thegenera Serrania, Bigotinops and Bigotinella. The cranidiastudied herein show a very developed sculpture with finepunctae distributed all over the carapace, characteristic ofsome primitive genera of trilobites such as Bigotinella. Ourspecimens differ from the type species of the genus Bigotina(Bigotinella) malykanica Suvorova, 1960 in the lack of a plec-trum.

At the present state of knowledge and because of the scar-city and poor preservation of this material we prefer to leavethis material in open nomenclature.

Stratigraphical distribution and biochronology:AP1 section, horizon 5 of trilobites (T5). Lower Ovetian. Inaddition, two poorly preserved cranidia from horizons T1 andT4 ofAP1 section have been doubtfully assigned to this taxon.

Genus Bigotina Cobbold, 1935.Type species: Bigotina bivallata Cobbold, 1935.Genus Bigotina has been widely revised by Pillola (1993)

who discussed its diagnostic characteristics and differencesclose genera such as Bigotinops Hupé, 1953; BigotinellaSuvorova, 1960; Hupetina Sdzuy, 1978; Serrania LiñánGuijarro, 1978; Ouijjania Hupé, 1953; Pruvostina Hupé,1953; and Tolbinella Egorova, 1983 among others of this age.

Fig. 5. Arroyo de Pedroche 2 section (AP2).Fig. 5. Coupe de Arroyo de Pedroche 2 (AP2).


Fig. 6. Scale bar = 1 mm. 1. cf. Bigotinella sp. Cranidium, calcitic exoskeleton. Note the sculpture with craters both in the exoskeleton and the internal mould.MPZ 2002/74a. AP1 section, horizon T5. 2. cf. Bigotinella sp. Cranidium, calcitic exoskeleton showing sculpture with craters. MPZ 2002/75. AP1 section,


In accordance with Pillola’s opinion, we include in Bigo-tina only the species B. bivallata Cobbold, 1935 and Bigo-tina copiosa (Egorova, 1983). Bigotina (Bigotina) egregicaRepina, 1960 has, among other features, a different anteriorborder and lacks of an ocular furrow, and is thus might beassigned to a new genus.

B. bivallata Cobbold, 1935Figs. 6(3–9) and 7(4, 5).1926. Ptychoparia – Bigot, p. 136, Pl. 4, Figs. 1–4.1935. B. bivallata n. sp. - Cobbold, p. 384, Pl. 17, Figs. 1–

10.1959. B. bivallata Cobbold - Sdzuy, pp. 396, Pl. 1, Fig. 6.1981. Bigotina sp. - Sdzuy, p. 396, Fig. 18.?1981. Gen. and sp. ind. - Sdzuy, p. 396, Fig. 13.1981. Gen. and sp. ind. - Sdzuy, p. 396, Fig. 14.1993. B. bivallata Cobbold - Pillola, pp. 162–163, Pl. 31,

Figs. 9–11.1993. B. bivallata Cobbold - Pillola, pp. 870–874, Pl. 2,

Figs. 1–6, Pl. 3, Figs. 1–9, Pl. 4, Figs. 1–7, Pl. 5, Figs. 1–3,5–7, 10.

v1993. B. bivallata Cobbold - Liñán and Gámez-Vintaned,p. 839–840, Fig. 6C, D.

Material: Thirty adult cranidia (MPZ 2002/76 to 105), onehundred juvenile cranidia and larvae (MPZ 2002/106 to 205),five librigenae (MPZ 2002/206 to 210), three rostral plates(MPZ 2002/211 to 213), and four incomplete thoracic seg-ments (MPZ 2002/214 to 217).

Description: Pillola (1993) describes accurately thecranidium, thorax, librigena, pygidium and larvae of B. bival-lata. We add the rostral plate description, unknown up to now.

Rostral plate (Fig. 7(5)): Anterior margin arcuate. Poste-rior margin more arcuate than anterior margin so that the ros-tral plate is slightly narrower in its middle part and becomeswider (exsag.) towards its lateral margins. Lateral margins(connective sutures) straight. The rostral plate shows terraceridges subparallel to its anterior and posterior margins.

The “S” parameter (Fig. 6(7)) is tangent or does not eventouch the palpebral lobe (external to eyes) as is present in theFrench material studied by Pillola (1993).

Discussion: The cranidia described by Pillola (1993) aresimilar to those presented herein including those where thefacial lines are not well marked. The only difference we havenoticed is that the plectrum is not always well marked in theSpanish specimens from Córdoba.

Stratigraphical distribution and biochronology: Arroyode Pedroche 1 section, horizons T3 to T7. Lower Ovetian.

Family REDLICHIIDAE Poulsen, 1927.Subfamily PARAREDLICHIIDAE Hupé, 1953.Genus Lemdadella Sdzuy, 1978.Type species: Lemdadella spectabilis Sdzuy, 1978.Sdzuy (1978: p. 92) defined Lemdadella as an opisthopar-

ian trilobite with features of the Pararedlichiinae, with wideto moderately wide preglabellar field, parafrontal band andocular furrows, plectrum and facial lines.

Liñán Guijarro (1978: p. 152) completed this diagnosisgiving the features of the librigenae, thorax and pygidium:semicircular cephalon with two long genal spines; fourteenor even fifteen thoracic segments with a pleural furrow andthe axis divided longitudinally into three parts. The pleuralfurrow of the thorax is wide and leaf-shaped and the axis hasan axial node. The pygidium has four segments and centralnode on each one. The hypostome has three pairs of smallspines in the posterior border.

Lemdadella has morphological similarities with the Boto-man genus Chorbusulina Lazarenko, 1962 from Siberia(Repina et al., 1974) in the cranidium, librigenae and py-gidium; differs in its characteristic furrow of the palpebrallobe which is absent in Chorbusulina. The flattened anteriorborder of the cranidium and the low angle of the pygidialpleurae with the axis are the only differences from BigotinaCobbold, 1935.

Lemdadella is known from the Ouneïn area in the HighAtlas (Morocco) by the species L. spectabilis Sdzuy, 1978and in the Anti-Atlas by the species L. tioutensis Sdzuy inLiñán and Sdzuy, 1978; both species occur in Zone 0 of ear-liest Cambrian trilobites (Sdzuy, 1981). The type species isrecorded in the lowest trilobite horizon of the Lemdad sec-tion.

horizon T5. 3. B. bivallata Cobbold, 1935. Cranidium, calcitic exoskeleton. MPZ 2002/76. AP1 section, horizon T5. 4. B. bivallata Cobbold, 1935. Cranidium,internal mould. MPZ 2002/77a. AP1 section, horizon T6. 5. B. bivallata Cobbold, 1935. Cranidium, internal mould. MPZ 2002/78. AP1 section, horizon T4.6. B. bivallata Cobbold, 1935. Cranidium, internal mould. MPZ 2002/79. AP1 section, horizon T7. 7. B. bivallata Cobbold, 1935. Cranidium, internal mould.MPZ 2002/80. AP1 section, horizon T6. 8. B. bivallata Cobbold, 1935. Cranidium, calcitic exoskeleton. MPZ 2002/81. AP1 section, horizon T5. 9. B. bivallataCobbold, 1935. Cranidium, internal mould. MPZ 2002/82. AP1 section, horizon T3. 10. L. perejoni Liñán sp. nov. Cranidium, internal mould. MPZ 2002/662.AP1 section, horizon T12. Note the external and convergent S parameter. 11. L. linaresae Liñán and Sdzuy, 1978. Cranidium, internal mould. MPZ 2002/230.AP2 section, horizon T11. See the tangential S parameter. 12, 13. L. linaresae Liñán and Sdzuy, 1978. Cranidium, internal mould. MPZ 2002/231 and MPZ2002/232. PH section, horizon 3. 14. L. linaresae Liñán and Sdzuy, 1978. Cranidium, internal mould. MPZ 2002/233. AP2 section, horizon T11.Fig. 6. Barre d’échelle = 1 mm. 1. cf. Bigotinella sp. Cranidium, exosquelette calcitique. Remarquer le relief cratériforme tant dans l’exosquelette que dans lemoule interne. MPZ 2002/74a. 5. Section AP1, horizon T5. 2. cf. Bigotinella sp. Cranidium, exosquelette calcitique montrant le relief cratériforme. MPZ2002/75. Section AP1, horizon T5. 3. B. bivallata Cobbold, 1935. Cranidium, exosquelette calcitique. MPZ 2002/76. Section AP1, horizon T5. 4. B. bivallataCobbold, 1935. Cranidium, moule interne. MPZ 2002/77a. Section AP1, horizon T6. 5. B. bivallata Cobbold, 1935. Cranidium, moule interne. MPZ 2002/78.Section AP1, horizon T4. 6. B. bivallata Cobbold, 1935. Cranidium, moule interne. MPZ 2002/79. Section AP1, horizon T7. 7. B. bivallata Cobbold, 1935.Cranidium, moule interne. MPZ 2002/80. Section AP1, horizon T6. 8. B. bivallata Cobbold, 1935. Cranidium, exosquelette calcitique. MPZ 2002/81. SectionAP1, horizon T5. 9. B. bivallata Cobbold, 1935. Cranidium, moule interne. MPZ 2002/82. Section AP1, horizon T3. 10. L. perejoni Liñán sp. nov. Cranidium,moule interne, MPZ 2002/662. Section AP1, horizon T12. Remarquer le paramètre S externe et convergent. 11. L. linaresae Liñán et Sdzuy, 1978. Cranidium,moule interne. MPZ 2002/230. Section AP2, horizon T11. Remarquer le paramètre S tangente. 12, 13. L. linaresae Liñán et Sdzuy, 1978. Cranidium, mouleinterne. MPZ 2002/231 et MPZ 2002/232. Section PH, horizon 3. 14. L. linaresae Liñán et Sdzuy 1978. Cranidium, moule interne. MPZ 2002/233. SectionAP2, horizon T11.


Fig. 7. Scale bar = 1 mm. 1. L. linaresae Liñán and Sdzuy, 1978. Moulting showing a meraspis on the border of librigena. MPZ 2002/218. AP2 section, horizonT11. 2. L. linaresae Liñán and Sdzuy, 1978. Complete specimen, internal latex mould. MPZ 2002/219. PH section, horizon 3. 3. L. linaresae Liñán and Sdzuy,1978. Cranidium, internal mould. MPZ 2002/234. AP2 section, horizon T11. 4. B. bivallata Cobbold, 1935. Librigena, internal mould. MPZ 2002/206.


In Sierra Morena (Spain), L. linaresae Liñán and Sdzuy,1978 occurs in the middle part of the Pedroche Formation,member I, corresponding to a lower Ovetian age.

Palmer and Rowell (1995) found Lemdadella antarcticaePalmer in Palmer and Rowell, 1995 in the Shackleton Lime-stone of Antarctica, corresponding to an early Atdabanian age(i.e. early Ovetian).

Another species is L. perejoni sp. nov. which is definedbelow.

Lemdadella linaresae Liñán and Sdzuy, 1978.Figs. 6(11–14) and 7(1–3).v1974. Dolerolenus formosus Sdzuy-Liñán Guijarro, p. 16.v1978. L. linaresae n. sp. - Liñán and Sdzuy, p. 391, Pl. 1,

Figs. 1–3.

v1978. L. linaresae Liñán and Sdzuy-Liñán Guijarro, p.153–169, Pl. 3, Figs. 1–10, Pl. 4, Figs. 1–12, Pl. 5, Figs. 1–13, Pl. 6, Figs. 1–13, Pl. 7, Figs. 1–12.

Material: Twenty-six almost complete specimens (MPZ2002/218-229, 289-299, 319, 324 and 571), sixty-fourcranidia (MPZ 2002/230-288, 300-302, 318 and 559), twohundred forty-five juvenile cranidia and larvae (MPZ2002/320-322, 330-558, 560-570, 572 and 573), thirty-fivelibrigenae (MPZ 2002/303-317 and 630 to 649), six pygidia(MPZ 2002/323 and 650-654), two rostral plates (MPZ2002/655 and 656) and five hypostomes (MPZ 2002/657-661); some of them preserved as original carapace in bluelimestones but most preserved as internal and external mouldswith limonite in green and yellow lutites.

Description: For the diagnosis and a complete descrip-tion see Liñán and Sdzuy (1978); Liñán Guijarro (1978). Therostral plate was previously unknown so we add here itsdescription. It is very similar to the rostral plate of B. bival-lata.

Rostral plate: Homogeneous width (sag.). Arcuate ante-rior margin slightly pointed in the sagittal axis. Strongly con-vex surface, with terrace ridges subparallel to the anterior mar-gin.

Remarks: Liñán Guijarro (1978) studied 50 completespecimens and five hundred cranidia from the typus locality

of Puente de Hierro (PH) showing the high population vari-ability by means of six morphotypes (A–F). We have nowstudied new material from the type locality and from two newsections:Arroyo de Pedroche 1 (AP1) andArroyo de Pedroche2 (AP2).

In AP1 section, L. linaresae is represented by two speci-mens. One is a juvenile cranidium from horizon T7, pre-served as external and internal mould and assigned to mor-photype E. The other is an adult cranidium (morphotype A)from level 8 preserved as a calcite exoskeleton. Two librige-nae from horizon T9 could be of this species.

The material studied from the AP2 section belongs to themorphotypes A and B (Fig. 6(11, 14)). In this section, wehave noticed that morphotype B appears before morphotypeA, then we can find both morphotypes occurring together and,some metres above, morphotype B is statistically replaced bymorphotypeA. This fact agrees with the stratigraphical recordof these morphotypes in the Puente de Hierro section (LiñánGuijarro, 1978: p. 74, Fig. 12). The replacement of the mor-photypes shows the evolutionary trend of the Lemdadellapopulation towards a narrower glabella (smaller glabellawidth/glabellar length relationship) and the palpebral lobeapproaching towards the glabella.

Discussion: L. linaresae, morphotype E, strongly re-sembles L. spectabilis Sdzuy, 1978 which has a narrowerpalpebral area. Further studies are needed to know if they arereally two geographic species or two ecotypes of the samespecies.

Stratigraphical distribution and biochronology: Puentede Hierro locality, Arroyo de Pedroche 1 section (horizonsT7 and T8) and Arroyo de Pedroche 2 section (horizons T10,T11, and perhaps T12). Lower Ovetian.

Lemdadella aff. linaresae Liñán and Sdzuy, 1978.Fig. 7(11).v1974. Lunolenus lotzei Sdzuy - Liñán Guijarro, p. 16.v1978. Lemdadella sp. 1 - Liñán Guijarro, p. 169–170,

Pl. 7, Figs. 13 and 14.?v1978. Lemdadella sp. indet. - Liñán Guijarro, p. 170–

171, Pl. 7, Fig. 15.

AP1 section, horizon T7. Note the granulose and bulky border and compare with librigena of L. linaresae (Fig. 7(2)). 5. B. bivallata Cobbold, 1935. Rostralplate. MPZ 2002/211. AP1 section, horizon T7. 6, 7. L. perejoni Liñán sp. nov. Cranidium, internal mould. MPZ 2002/663 and MPZ 2002/664. AP1 section,horizon T12. Note the external and convergent S parameter. 8. L. perejoni Liñán sp. nov. Hypostome, internal mould. MPZ 2002/699. AP1 section, horizon T12.9, 10. L. perejoni Liñán sp. nov. Cranidium, internal mould. MPZ 2002/665 and MPZ 2002/666 (holotype). AP1 section, horizon T11. Note the external Sparameter 11. Lemdadella aff. linaresae Liñán and Sdzuy, 1978. Cranidium, internal mould. MPZ 2002/704a. AP1 section, horizon T6. 12, 13. Eoredlichia cf.ovetensis (Sdzuy in Liñán and Sdzuy, 1978). Cranidium, internal mould. MPZ 2002/705 and MPZ 2002/706. AP2 section, horizon T15. Note the narrow andbulky eye and the absence of ocular furrow. 14. Eoredlichia cf. ovetensis (Sdzuy in Liñán and Sdzuy, 1978). Cranidium, internal mould. MPZ 2002/707.AP1 section, horizon T12. Note the narrow and bulky eye, the absence of ocular furrow and the insertion point in the glabella. Compare with Fig. 7(10).Fig. 7. Barre d’échelle = 1 mm. 1. L. linaresae Liñán et Sdzuy, 1978. Mue montrant une larve meraspide au bord de la librigène. MPZ 2002/218. Section AP2,horizon T11. 2. L. linaresae Liñán et Sdzuy, 1978. Spécimen complet, moule interne en latex. MPZ 2002/219. Section PH, horizon 3. 3. L. linaresae Liñán etSdzuy, 1978. Cranidium, moule interne. MPZ 2002/234. Section AP2, horizon T11. 4. B. bivallata Cobbold, 1935. Librigène, moule interne. MPZ 2002/206.Section AP1, horizon T7. Remarquer le bord granuleux et bombé et comparer le avec la librigène de L. linaresae (Fig. 7(2)). 5. B. bivallata Cobbold, 1935.Plaque rostrale. MPZ 2002/211. Section AP1, horizon T7. 6, 7. L. perejoni Liñán sp. nov. Cranidium, moule interne. MPZ 2002/663 et MPZ 2002/664. SectionAP1, horizon T12. Remarquer le paramètre S externe et convergent. 8. L. perejoni Liñán sp. nov. Hypostome, moule interne. MPZ 2002/699. Section AP1,horizon T12. 9, 10. L. perejoni Liñán sp. nov. Cranidium, moule interne. MPZ 2002/665 et 666 (holotype). Section AP2, horizon T11. Remarquer le paramètreS externe. 11. Lemdadella aff. linaresae Liñán et Sdzuy, 1978. Cranidium, moule interne. MPZ 2002/704a. Section AP1, horizon T6. 12, 13. Eoredlichia cf.ovetensis (Sdzuy in Liñán et Sdzuy, 1978). Cranidium, moule interne. MPZ 2002/705 et MPZ 2002/706. Section AP2, horizon T15. Remarquer l’oeil étroit etbombé et l’absence de sillon oculaire. 14. Eoredlichia cf. ovetensis (Sdzuy in Liñán et Sdzuy, 1978). Cranidium, moule interne. MPZ 2002/707. Section AP1,horizon T12. Remarquer l’oeil étroit et bombé et l’absence de sillon oculaire. Comparer avec la Fig. 7(10).


Material: One cranidium preserved in yellow lutites asinternal and external mould with limonite. MPZ 2002/704a,b.

Description: Cranidium with a granulose sculpture dis-tributed all over its surface.

Arcuate anterior margin. Homogeneous anterior border,almost flat and as wide as the preglabellar field. Faint ante-rior border furrow. Preglabellar field occupied by a slightlymarked plectrum. Plectrum limited laterally by very faint fur-rows that become deeper near the glabella. Strongly convexglabella subcylindrical in shape. This has three pairs of fur-rows deep and not transglabellar. Frontal lobe subsemicircu-lar. L3 is shorter (sag.) than L1 and L2. Occipital furrow fadestowards the sagittal axis and curves anteriorly. Non-homogeneous occipital ring as wide as L2. This presents afaint furrow, almost parallel to the occipital furrow, thatdivides the occipital ring into two unequal parts. The anteriorpart of the occipital ring is w-shaped and the posterior part issemicircular in shape, narrower (sag.) than the anterior oneand presents an occipital node. Preocular field flat and trian-gular in shape. This has straight facial lines parallel to theanterior branches of the facial suture. The parafrontal bandthickens in front of the glabella. Straight ocular ridge parallelto the anterior margin. Palpebral lobe parallel to the sagittalaxis, thicker in its middle part than in its edges. Ocular fur-row present, more marked in the ocular ridge, near the gla-bella, than in the palpebral lobe. Deep palpebral furrow. Flatpalpebral areas subtriangular in shape and occurring at higherlevel than the preocular field. Posterior border very narrow(approximately 1/3 of the occipital ring width) and homoge-neous.Anterior branches of the facial suture straight and diver-gent. Middle branches of the facial suture curved and subpar-allel. Posterior branches of the facial suture slightly curvedtowards the sagittal axis and strongly convergent. “S” param-eter secant and subparallel.

Discussion: L. aff. linaresae resembles morphotype B ofL. linaresae Liñán and Sdzuy, 1978 within which it could beincluded, but it shows a glabella with a bigger glabellarwidth/glabellar length relation. This character is also presentin L. linaresae sp. 1 from the Puente de Hierro locality (LiñánGuijarro, 1978). Sdzuy (1981) figured one undescribedcranidium of Zone 0 from Amouslek, Anti-Atlas (p. 396,Fig. 14) very similar to the one here described; because of itspoor preservation we do not include it in the synonymy. Somespecimens of B. bivallata with non-prominent anterior bor-der resemble L. aff. linaresae and could be interpreted in thesense that B. bivallata is the ancestor of Lemdadella. Never-theless, the ontogenetic study of Lemdadella (see LiñánGuijarro, 1978) combined with the presence of Lemdadellaspecies with long eyes in Morocco (Sdzuy, 1978; Liñán andSdzuy, 1978) may support the view that B. bivallata is part ofa lineage parallel to the Lemdadella species, both with a prob-able common ancestor.

Stratigraphical distribution and biochronology: Arroyode Pedroche 1 section, horizon T6. Lower Ovetian. Oneincomplete cranidium of Lemdadella sp. indet. found in hori-

zon T5 might correspond to L. aff. linaresae. Puente de Hierrosection, horizon 3 (material figured in Liñán Guijarro, 1978).

Lemdadella perejoni sp. nov.Figs. 6(10) and 7(6–10).Material: Thirty six cranidia (MPZ 2002/600-615 and 662-

681), four juvenile cranidia and larva (MPZ 2002/616-619),fifteen librigenae (MPZ 2002/627-629 and 682-693), sevenpygidia (MPZ 2002/625, 626 and 694-698), four hypos-tomes (MPZ 2002/620 and 699-701), two rostral plates (MPZ2002/702 and 703) and four thoracic segments (MPZ2002/621-624).

Derivatio nominis: Dr. A. Perejón from the UniversidadComplutense of Madrid and CSIC.

Locus typicus: Arroyo de Pedroche 1 section, level 12.Sierra de Córdoba, Spain.

Holotype: MPZ 2002/666 (Fig. 7(10)).Paratypes: MPZ 2002/662-665 (Fig. 7(6–9)).Diagnosis: Species of Lemdadella characterised by pre-

ocular, ocular and posterior portions of the facial suture ofquite similar length, “S” parameter markedly external to theeye and slightly convergent, and a smooth palpebral furrow.

Description: Cranidium with a granulose sculpture. Arcu-ate anterior margin. Homogeneous and almost flat anteriorborder, narrower (sag.) than the preglabellar field. This is sepa-rated from the preocular and preglabellar fields by a shallowfurrow. Preglabellar field with a flat plectrum that is narrower(trans.) in its middle part. Strongly convex glabella slightlynarrower forward. It has three pairs of non-transglabellar fur-rows, backward directed. Occipital furrow deeper exsagit-tally than sagittally. Occipital ring convex and homoge-neous. Flat preocular field with straight facial lines parallelto the anterior branches of the facial suture. Parafrontal bandanteriorly distinguished from the preocular field by a deepfurrow. Arcuate eye-ridge joins the glabella in the base of thefrontal lobe and has a shallow ocular furrow that fades back-wards. Palpebral lobe arcuate, wide, homogeneous. It showsshort and very faint ocular furrow located in the posteriorhalf of the palpebral lobe. Deep palpebral furrow. Flat palpe-bral area subtriangular in shape. Posterior border very nar-row (half of the palpebral lobe width) and non-homogeneous.Straight posterior margin. External and slightly convergentforwards, almost parallel, “S” parameter. Anterior branchesof the facial suture straight and divergent, medial branches ofthe facial suture arcuate and slightly convergent. Posteriorbranches of the facial suture straight and strongly conver-gent. The angle between the posterior margin and the poste-rior branch of the suture is very low (less than 45°). Hypos-tome and rostral plate isolated. Rostral plate typical forLemdadella (see L. linaresae above). Hypostome subellipti-cal in shape. It has a granulate sculpture distributed all overits surface. Anterior border of the hypostome unknown.Middle body strongly convex, subelliptical in shape andbecoming narrower backwards instead of having a middlefurrow dividing it into two lobes. Homogeneous posterior bor-der.


The librigena, thorax and pygidium are typical of the genusLemdadella.

Discussion: The evolutionary trend in Lemdadella spe-cies from Córdoba seems to be reflected in the progressiveshallowing of the ocular furrow together with a more roundedpalpebral lobe and an enlargement of the anterior and poste-rior portions of the facial sutures.

L. perejoni differs from its assumed ancestor L. linaresaefrom Sierra Morena and L. spectabilis from the High Atlas inhaving longer posterior branches of the facial suture, an exter-nal “S” parameter and more divergent anterior branches ofthe facial suture. It differs from L. tioutensis Sdzuy in Liñánand Sdzuy, 1978 from the Anti-Atlas in having a relativelywider palpebral area; from L. antarcticae Palmer in Palmerand Rowell, 1995 in the homogeneity of the anterior border.In the youngest specimens, the ocular furrows are not alwayswell preserved and we have some intermediate specimenswhich we have difficulty in assigning finally to either Lem-dadella or Eoredlichia.

Stratigraphical distribution and biochronology: Arroyode Pedroche 1 section (horizon T12) and Arroyo de Pedroche2 section (horizon T11). Lower Ovetian.

Genus Eoredlichia Chang in Lu and Dong, 1952.Type species: Redlichia intermedia Lu, 1940.

Eoredlichia cf. ovetensis (Sdzuy in Liñán and Sdzuy,1978).

Fig. 7(12–14).Material: Four incomplete cranidia preserved in yellow

lutite as internal and external mould with limonite. MPZ2002/705 to MPZ 2002/708.

Description: Cranidium with an arcuate anterior margin.Anterior border homogeneous, slightly convex and dippingbackwards. It has terrace ridges. Anterior border furrow shal-low and less marked in front of the glabella. Preglabellar fielddepressed. Subcylindrical glabella with three pairs of non-transglabellar furrows well marked. Semicircular frontal lobe.Deep occipital furrow only distally. The occipital ring seemsto be as wide as L1. Convex preocular field triangular in shape.This is placed at a higher level than the palpebral area. Pre-ocular furrow deeper near the glabella. Ocular ridge and palpe-bral lobe wide and homogeneous. Straight ocular ridge thatfits the glabella in its anterior part. Curved palpebral lobe sub-parallel to the sagittal axis. Narrow (sag.) postocular fieldslightly wider than the occipital ring. Posterior border nar-row and homogeneous. Anterior branches of the facial suturestraight and strongly divergent, medial branches of the facialsuture curved and slightly convergent.

Discussion: The specimens from the Arroyo de Pedroche1 and 2 sections show the typical features of Eoredlichia: noocular furrow, the eyes are narrower next to the glabella andthe anterior branches of the facial suture forms an acute anglewith the eye (Fig. 7(13)). A librigena found in horizon T14 ofthe AP1 section is quite similar to Lemdadella species andmay be included in Eoredlichia cf. ovetensis. There are somedifferences in the width of the preglabellar field and in the

interocular field of the fixigenae considered herein as intraspe-cific variations.

Our material resembles E. ovetensis (Sdzuy in Liñán andSdzuy, 1978) from the Cantabrian Mountains but there aresome differences in the curvature of the anterior border, inthe length of the eye and posterior branches of the facialsuture. E. ovetensis has the ocular ridges extending up thesides of the glabella forward from about S3, almost to meetin front of the glabella, which is not shown in E. cf. ovetensis.Because we do not have an appropriate number of specimensto know the taxonomic relation between material from bothSpanish localities in terms of population variability, we haveleft the material from Córdoba in open nomenclature. Ourspecimens also resemble E. pulchella Hupé, 1953 of the Anti-Atlas but the anterior border of that species is wider than inthe material from Sierra de Córdoba.

E. cf. ovetensis also has similarities with L. perejoni Liñánsp. nov. which is its probable ancestor but it cannot be placedin Lemdadella because of the absence of an ocular furrow.

Stratigraphical distribution and biochronology: Arroyode Pedroche 1 section, horizons T12b and probably T14.Arroyo de Pedroche 2 section, horizon T15. Lower Ovetian.

5. Biochronology

5.1. Lower Cambrian stages in the Mediterraneansubprovince

Using selected trilobite assemblages as chronomarkers,Sdzuy (1971) proposed the following regional stages for theIberian Peninsula from the base up: Ovetian, Marianian andBilbilian. For beds bearing Cambrian trace fossils that areunderlying rocks containing the first Ovetian fossil assem-blages, Liñán (1984) proposed the Corduban stage. These fourLower Cambrian regional stages were revised by Liñán et al.(1993) with new stratigraphical and palaeontological data (tri-lobites, archaeocyaths and trace fossils). Recent works on acri-tarch biochronology (Gámez et al., 1991; Palacios and Vidal,1992; Palacios, 1993) and small shelly fossils (Fernández-Remolar, 1996) enable a more complete characterisation ofsome of these stages. These chronostratigraphic units haveusually been cited in Cambrian successions of Portugal(Gonçalves, 1982), Sardinia (Pillola et al., 1995), Germany(Elicki, 1997), France (Álvaro et al., 1998) and Jordan (Rush-ton and Powell, 1998). They may also be applied to Moroccowhere a high number of trilobite species are common.

The Ovetian is widely represented in Iberia where manywell exposed sequences can be found in its northern, south-ern and central parts. All these sequences are well character-ised by archaeocyath, ichnofossil and trilobite assemblages.At the moment, the lower Ovetian biostratigraphy has beenmainly established by archaeocyaths (Zones I–III of Perejón,1986); thus the new complementary data on trilobite will beuseful for a better intercontinental correlation of these earlyCambrian strata.


The Ovetian lower boundary is defined by archaeocyathsof Zone I of Perejón (1986) which appear together with unde-termined Bigotinidae. Using archaeocyaths, a tentative cor-relation of the Ovetian with the Atdabanian of Siberia hasbeen made (Perejón, 1986; Debrenne et al., 1990; Zhuravlev,1995). This correlation is in agreement with data from smallshelly fossils (Fernández Remolar, 2001) and with the pres-ence of Bigotinidae both in Iberia (Ovetian) and Siberia(reported from the LowerAtdabanian by Repina, 1966). Smallshelly fossils found in the Pedroche Formation of Sierra deCórdoba also permit a correlation between the lower Ovetian(Zones I–III) and the Upper Meishucunian stage of China(Fernández Remolar, 1996, 2001). On the other hand, trilo-bites permit a good correlation between the Ovetian andHupé’s Amouslek and lower Timghit stages from Morocco(Sdzuy, 1971), and the Issendalenian stage proposed by Geyer(1990a).

5.2. Biozones

The Lower Cambrian zonation of Iberia is based on tracefossils for the Corduban, on archaeocyaths for the Ovetian-Marianian and on trilobites for the Bilbilian (Fig. 8). Smallshelly fossils have been used to analyse the Precambrian-Cambrian boundary in Iberia (Vidal et al., 1995; Palacios etal., 1999a, 1999b), which is well established by the Phy-

codes pedum-Monomorphichnus lineatus trace fossils assem-blage zone (Gámez-Vintaned and Liñán, 1995). An archaeo-cyath zonation containing seven assemblage zones wasproposed by Perejón (1986) and subsequently modified byPerejón (1994); Moreno-Eiris et al. (1995). The acritarchzonation proposed for the East European Platform (Moczy-dłowska, 1991) has also been applied to some Ovetian rocksof Northern Iberia (Palacios and Vidal, 1992), thus allowinga good correlation with Baltica.

The new trilobite discoveries permit us to propose herethree trilobite interval biozones for the lower Ovetian of SierraMorena: B. bivallata Biozone, Lemdadella linarease Bio-zone and L. perejoni Biozone. They are limited by the firstappearance datum (FAD) of the guide species of each bio-zone, following an evolutionary lineage (i.e. interval phylo-zones). The upper boundary of the L. perejoni Biozone isplaced at the first occurrence of Eoredlichia cf. ovetensiswhich is a young descendant of this lineage.

The B. bivallata Biozone contains the trilobites B. bival-lata, Lemdadella aff. linaresae and aff. Bigotinella; thearchaeocyaths Aldanocyathus carteretensis, Aldanocyathusvaldegrajensis, Agastrocyathus gregarius, Rotundocyathuskhemtschikensis, Aldanocyathus notabilis and Reticoscinussp.; the brachiopod Paterina sp.; the calcimicrobes Epiphy-ton sp., Renalcis sp. and Girvanella sp.; the stromatoliteVetella nodosa and the trace fossils Cochlichnus ichnosp.,

Fig. 8. Lower Cambrian chrono- and biostratigraphic units in the Iberian Peninsula, including the new trilobite biozones proposed here.Fig. 8. Unités chrono- et biostratigraphiques du Cambrien inférieur dans la Péninsule Ibérique, avec les nouvelles biozones de trilobites proposées dans cetravail.


Phycodes ichnosp., Planolites montanus, Psammichnites ich-nosp. and Treptichnus bifurcus; and the probable soft-bodiedcoelenterate cf. Tiernavia.

The L. linaresae Biozone contains the trilobites L. linare-sae, B. bivallata (in the lower part only), Serrania verae andL. aff. linaresae; the archaeocyaths Aldanocyathus carter-etensis and Aldanocyathus virgatus; the calcimicrobes Epi-phyton sp., Renalcis sp., Girvanella sp., Kordephyton sp. andTubomorphophyton sp.; the brachiopod Paterina sp; and thetrace fossils Bergaueria perata, aff. Bergaueria, Bilinichnusichnosp., Cochlichnus ichnosp., Dactyloidites cabanasi, Phy-codes pedum, Phycodes ichnosp., Planolites annularis, P.montanus, Psammichnites gigas, Psammichnites ichnosp.,Rusophycus ichnosp., Torrowangea ichnosp., Treptichnusbifurcus and T. ichnosp.

The L. perejoni Biozone contains the trilobites L. pere-joni, L. linaresae (only in the lower part) and Neoredlichidaeindet., indetermined archaecyaths, the brachiopod Paterinasp., and the trace fossils Planolites montanus and Torrowan-gea ichnosp.

The Eoredlichia cf. ovetensis assemblage contains the tri-lobites L. perejoni (in its lower part) and Neoredlichidae sp.,and the archaeocyaths Aldanocyathus notabilis and Robus-tocyathus moori.

5.3. Correlation

The B. bivallata Biozone almost coincides with the lowerpart of the archaeocyath Zone III at the AP1 section (Fig. 4).Nevertheless, since trilobites are lacking in the lower part ofthis section, an extension of the B. bivallata Biozone into theuppermost part of archaeocyath Zone II cannot be excluded.The B. bivallata Biozone is also recognised in the LowerCambrian Saint-Jean-de-la-Rivière Formation at Carteret inNormandy, France (lower Ovetian; Pillola et al., 1994), wherethe index taxon shows a longer stratigraphical range. Thisobservation, together with the presence of Aldanocyathus car-teretensis within archaeocyath Zones II and III at section AP1(Perejón, 1989) noted above, supports the probable exten-sion of the B. bivallata Biozone into archaeocyath Zone II.On the other hand, the species of Bigotinella from Siberia(Lena river) are Atdabanian in age (Pagetiellus anabarusZone), which agrees with the archaeocyath correlation madeby Perejón (1986). Furthermore, Pillola (1993) included onespecimen from Zone 0 of Morocco (figured by Sdzuy, 1981)in B. bivallata.

The L. linaresae and L. perejoni Biozone correspond tothe upper part of archaeocyath Zone III. The two guide spe-cies of these biozones are close to L. spectabilis and L. tiouten-sis (see above, trilobite systematic palaeontology), respec-tively, from Zone 0 and Zone I (F. tazemmourtensis Zone) ofMorocco (Sdzuy, 1978,1981; Liñán and Sdzuy, 1978).

In summary, if our conclusions are correct, the B. bival-lata and the L. linaresae Biozones correlate to Zone 0 ofMorocco, and the L. perejoni Biozone to Zone I (F. tazem-mourtensis Zone). Finally, the three established biozones

probably correspond to the Atdabanian stage of Siberia andcorrelative beds on the East European Platform and in Ant-arctica Bigot, (1926).

Acknowledgements

We thank commentaries on the original manuscript fromDr. Perejón, Dr. Palmer, Dr. Rushton and Dr. Geyer. Finan-cial support was given by the Dirección General de Investi-gación, Spanish Ministerio de Educación y Cultura (ProjectsBTE2000-1145-C02, BTE2003-04997) and by the Diput-ación General de Aragón (Grupo Consolidado “Paleon-tología del Paleozoico”). The Servicio de Fotografía Cientí-fica de la Universidad de Zaragoza is thanked for acces to itsfacilities and for photographic work with trilobites.

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