Vol. LEIDSE GEOLOGISCHE MEDEDELINGEN, 39, pp. 129—192, published separately 5-4-1967

Devonian and Lower Carboniferous Conodonts of the Cantabrian Moun- tains (Spain) and their stratigraphic application

BY

H.A. van Adrichem Boogaert

Summary

of the literature the of the Devonian and the Lower Carboniferous ofthe Cantabrian Mountains A short review on stratigraphy

precedes the report of the author's stratigraphic and palaeontologic observations in León: the Río Esla area (Gedinnian to

central Cantabrian and the Gildar-Montó area in Asturias: Viséan), the area (Famennian to Viséan), (Eifelian to Viséan);

the coastal area (Frasnian to Viséan); in Palencia: the Arauz-Polentinos area (Gedinnian to Givetian), the Cardaňo-Triollo

and San Martín-Valsurvio area and Famennian and in Santander: the area (Eifelian to Viséan), the (Givetian to Viséan);

Liébana area (Eifelian to Viséan).

which extracted from calcareous could be in the zonal succession Most of the conodont faunas, were formations, arranged

established in Germany, and thus supplied new data about several formations in the Cantabrian Mountains.

The of the Ermita Formation in Asturias and Palencia is demonstrated. The of this unit presence transgressive age ranges

from Famennian Tournaisian. The Cardaňo Formation from middle maximally uppermost to lowermost ranges or upper

Givetian Frasnian. The Vidrieros Formation from the of the lower Famennian the lowermost to upper ranges upper part to

Tournaisian.

A synthesis of the stratigraphic data delimits the Palentine facies area, which is clearly separated from the Asturo-Leonese

The units the Asturo-Leonese the Palentine facies area by positive areas. following palaeogeographic are distinguished: Basin,

Basin, and the Asturian Geanticline. The development of these units from the Middle Devonian to the Lower Carboniferous is

The sedimentation the Asturian Geanticline limited and incom- demonstratedby eight facies-pattern maps. on was probably plete. An epeirogenetic uplift of this geanticline took place in late Frasnian to early Famennian times. This uplift is correlated sheltered Palentine with the deposition of the quartzitic Murcia Formation in the Basin. The uplifted area was covered by the Ermita transgression in the late Famennian to early Tournaisian.

resulted in the Formation in the Tournaisian. In After a break in the sedimentation, a local transgression Vegamian upper

in the Tournaisian lower Viséan. In the Palentine Basin the most ofthe area the Alba transgression began uppermost or deposi-

started in the Viséan. tion of the Alba Formation upper

with the zonal forms of the conodonts. Three elements The chapter onsystematics deals mainly most important guide new are from the middle from the described: Icriodus eslaensis n.sp. to upper Givetian, Siphonodella? n.sp. a, probably upper Tournaisian, and A conodont from the Gedinnian lower n.gen. n.sp. a, a simple compound upper or Siegenian.

Contents

130 III. Zonation and of the Introduction dating stratigraphic sequence

by means of conodonts 155

I. General outline ofthe stratigraphy ofthe Devonian The biostratigraphic framework 155

and Lower Carboniferous of the Cantabrian Conodont zones demonstrated in the Canta- 131 Mountains brian Mountains 155

Leon 131

Asturias 131 IV. Discussion of the new data with respect to the

Palencia 132 Devonian and the Lower Carboniferous forma-

tions in the Cantabrian Mountains 158 General remarks 133

Devonian lowermost 159 (and Tournaisian). ...

134 The Asturo-Leonese facies area 159 II. Stratigraphic and palaeontologic observations . .

Leon 135 The Ermita Formation 159

135 The facies 161 The Rio Esla area Palentine area

138 The Abadia Formation 161 The central Cantabrian area

141 The Formation 161 The Gildar-Monto area Gustalapiedra

145 The Cardafio Formation 161 Asturias, coastal area

Palencia 146 The Murcia Formation 162

146 The Vidrieros Formation The Arauz-Polentinos area 162

149 Lower Carboniferous 162 The Cardafio-Triollo area

The Formation 163 The San Martin-Valsurvio area 153 Vegamian

154 The Alba Formation 164 Santander, the Liebana area 130 H. A. Adrichem Conodonts Cantabrian van Boogaert: of the Mountains

V. Palaeogeographic synthesis of the stratigraphic The Lower Carboniferous development of the

data 166 sedimentary basins 177

General remarks 166 VI. Conodont systematics 177 The Asturian Geanticline 169 Samenvatting 188

The and of the location, nature, development Sumario 188 Devonian sedimentary basins 173 References 189

The Asturo-Leonese Basin 173 3 Plates

The Palentine Basin 175 6 Enclosures

INTRODUCTION

The formed of San Isidro and Tarna Adrichem Boo- present study part a geological investiga- pass pass (van of the tion of the southern Cantabrian Mountains which gaert et al., 1963). Sampling and surveying sur-

to has been carried out since 1950 by staff members and roundings of the Pico Gildar and, more the east, basis for students of the University of Leiden, starting in the Cardano-Triollo area, provided the a northern Palencia. This study was gradually extended conodont stratigraphy of these regions. In 1965 a westwards, and in 1964 reached the westernmost final field trip was made to obtain additional data to in the southern of of the observations occurrence of the Devonian, part complete a synthesis stratigraphic "Asturian Knee". The the provisional geological map already compiled. of the southern slope of the Cantabrian Mountains Acknowledgements. — During the preparation of this of the thesis the author received and advice (de Sitter, 1962) provides a general picture area. special help

As of detailed and from Sincere thanks are extended part a more stratigraphic palaeon- many persons. to of directed A. Dr. W. Landesamt Nordrhein- tologic investigation the area by prof. Dr. Ziegler (Geologisches

Brouwer (Department of Stratigraphy and Paleonto- Westfalen, Krefeld, Germany), who guided the logy), the present author started a study of conodonts author's first step in conodont techniques and system- in 1959. atics. Thanks are also due Prof. Dr. O. H. Walliser

of Initially, conodont faunas were obtained from Devo- (Gottingen, Germany) for the discussion problems the author with nomenclature and Lower Devonian nian rocks from the Rio Esla area, where respect to carried out stratigraphic fieldwork in the Esla conodontstratigraphy. Dr. P. Bender (Marburg/Lahn, the conodont autochthone from 1959 to 1961, but they were not Germany) kindly showed the author this collections of the of The author very abundant in area. Samples provided by University Marburg. is A. Mr. I. B. H. M. Rubbens in 1959 from the Pico also very grateful to Dr. J. G. Binnekamp, Mr. J. indicated Mr. A. Mr. R. W. Lan- Gildar, situated more to the north, a more van Hoeflaken, J. Kutterink, promising area for the study of conodonts. The results ting, Dr. N. Sjerp, and Dr. J. van Veen for providing his of sampling to the west of this area, near the water- samples or putting field data at disposal. Mrs.

contributed I. assisted with the text. shed, to a new stratigraphic interpretation Seeger-Wolf English of Palaeozoic sections in the region between the

1. Situation of the Cantabrian Mountains The shaded shows the Fig. map (1 : 6,000,000). quadrangle general region of the study. General outline of the stratigraphy 131

CHAPTER I

GENERAL OUTLINE OF THE STRATIGRAPHY OF THE DEVONIAN AND LOWER

CARBONIFEROUS OF THE CANTABRIAN MOUNTAINS

The Cantabrian Mountains (Fig. 1 ) have long Special attention has been paid by Wagner (1957,

attracted the attention of geologists, to whom they 1963), Wagner-Gentis (1963), and Kullmann (1961, offer excellent fossil and mineral and Alba exposures, riches, 1962, 1963a) to thecephalopodsofthe Formation,

interesting structures. Many papers on this mountain which indicate that this formationranges in age from chain been in of the last Lower Visean Visean. Namurian have published the course to Upper Lower

Publications with modern strati- were found in the lowermost beds of the century. dealing a goniatites of of it of direct Caliza transitionalbeds between graphic analysis the area or parts are de Montana or in the

importance for this paper. A review of these publica- the Alba Formation and this limestone.

tions is given below. Higgins (1962) reported on conodonts of the Alba

Formation in Leon and Palencia. In an investigation

LEÓN of the Devonian-Carboniferous transition in the southern of the Rio Bernesga region, Higgins The stratigraphy of the Lower Palaeozoic and the part et al. identified Tournaisian conodonts. Devonian in the northern of with the (1964) part Leon, New data the of near the of the northeasternmost is based on stratigraphy Leon, exception part, on al. watershed, reported by van Adrichem Boogaert et the work of Comte (1959), who distinguished as revealed the of the Ermita Formation Devonian and Lower Carboniferous units: (1963), presence overlying Ordovician quartzites. The Lower Palaeo- Griotte de Puente de Alba zoic of these had been age quartzites already correctly Couches de Vegamian Viseen estimated by Julivert (1960) and Martinez Alvarez lacune (1962) in Asturias. Sjerp (1967) has recently presented Gres de l'Ermitage Strunien lacune further details about the stratigraphy of this area. of Schistes de Fueyo Famennien Rupke (1965) published data on the structure the Rio Esla in which he dealt with the area, stratigraphy Calcaires de Valdore Galcaire. Portilla gasmen of the "Esla nappe", the "Esla autochthone", and the Gres et Schistes de Huergas _,,., "Las Salas zone" the northern of the (in part area Calcaires de Santa Lucia studied). Calcschistes et Calcaires de La Vid „. the of in the Siegenien In northeasternmost area Leon, Valdeon, Gres de San Pedro Gedinnien there is Devonian which differs (partie superieure) a sequence strongly

from the sequence known from the Rio Luna-Rio Esla mentioned The importance of the hiatus below the transgressive region. Julivert (1960) some cephalo- from this northeastern where the Devonian "Gres de l'Ermitage" gradually increases in a north- pods area, limestones, northeasterly direction to such an extent that finally consists of nodular and argillacous shales,

this sandstone directly overlies Cambrian formations. and quartzites. This author also assigned the forma-

Comte traced these units in northern Leon from the tions south of the Pico Gildar to the Devonian. Fossil

has shown these rocks Rio Luna, in the west, to the Rio Esla, in the east. evidence, including foraminifers, established to be of Westfalian Ginkel, The age of the units has been mainly by age (van 1965). theirbrachiopod fauna. Kullmann(1960, 1961) described a section in a part of the Valdeon which he called the Monto Most of the units introduced by Comte are now ac- area, ranging

formationsin the ofthe stated in from lower Famennian to Visean and cepted as sense principles age upper dated He named the Devonian nodu- by the International Subcommission on stratigraphic by cephalopods. nomenclature (Hedberg, 1961). A few slight changes lar limestones the "Monto Schichten". Budinger &

have been introduced, however. The "Calcaires de Kullmann(1964) traced this section into the Frasnian. Adrichem mentioned Givetian Valdore", mentioned only for the Rio Esla area, are Van Boogaert (1965) of Pico and considered the of the La Portilla conodonts from a section west Gildar as upper part called the "Gres from Formation. Van Ginkel (1965, p. 183) a spot sample. de l'Ermitage" the Ermita Formation. He redefined

the Vegamian Formation and the Alba Formation ASTURIAS

"Griotte de Puente de Alba"), and recognized these (= The framework of Devonian and the Lower Carboni- formations in Leon, Palencia, and Asturias. For the ferous stratigraphy in Asturias was provided by Barrois historic but confusing names "Caliza de Montana" (1882). He distinguished the following "zones", which Ginkel and "Calcaire des Canons", Brouwer & van can be considered as formations: formal of (1964, p. 309) proposed the name Escapa

Formation. Smits (1965) described an aberrant devel- Assise du Griotte (Marbre a Carbonifere inferieur Couvinian Goniatites) opment of the upper Emsian and the lower

de Cue near Caldas de Luna as the Caldas Formation. Gres Famennien 132 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

de Frasnien Calcaire Candas avec Spirifer (Rouault) are typical for the Ordovician accord-

Verneuli ing to Seilacher, 1962). Hernandez Sampelayo & Kin-

Gres Gosseletia Givetien delan Devonian a (1950, p. 60) proposed a Lower age

de for the of the formationin north- Calcaire Moniello avec Eifelien upperpart quartzitic Calceola sandalina eastern Asturias, which is locally conglomeratic and

Calcaire d'Arnao a Sp. cultrijugatus ferruginous.

de Ferrones Further to the west in in the Cabo Penas area Calcaire a Athyris Coblenzien sup. Asturias, where the Alba Formation is underlain somewhat Coblenzien inf. by a Schistes et calcaires de Nieva a there have also been uncertain- Sp. hystericus ferruginous sandstone, ties about the of this unit. To avoid confusion with Gres ferrugineux de Furada Taunusien age other stratigraphic units, Radig (1961) named this

sandstone Pifieres Sandstone. Altevogt (1963) and The subsequent work of Delepine (1928a, 1932a,

Radig (1964) found fossil evidence for an 1932b) and Comte (1959) provided new palaeontolo- upper Frasnian age of this sandstone. Delepine (1928b, 1937, gic evidence, which led to a revised stratigraphic 1943) studied the Alba Formation, from which he de- classification as presented by Comte (1959, p. 298). scribe dupper Visean goniatites. Comte supplied a comparison with the Devonian Lys & Serre (1958) published the first report on cono- formations of northern Leon, as follows: donts from the Cantabrian Mountains. They described Gres culminants de Candas ? Gres de l'Ermitage from the Alba Formation a fauna which they assigned

Calcaires de Candas Calcaires de Valdore the Visean. Kullmann to upper Buddinger & (1964) l'assise (sauf terminale) showed by means of conodonts and cephalopods that Calcaires de La Portilla from in Perlora this formation ranges in age lower to

Gres du Naranco Gres et Schistes de Huergas upper Visean. They mentioned the occurrence of Calcaires de Moniello (5.1.) Calcaires de Santa Lucia Tournaisian conodonts together with conodonts lower in section Complexe de Rafieces Complexe de La Vid generally recognized as Visean, a the transition Gres de Furada Gres de San Pedro incorporating Devonian/Carboniferous this (partie superieure) (partie superieure) near Entrago. They assigned a Tournaisian age to but Adrichem fauna, van Boogaert (1965, p. 168)

Llopis Llado (1958a, 1962) also used a classification suspected that it contains a stratigraphic admixture. based on the units of Barrois, and Radig (1961, 1964) and Poll modified version of Bar- (1963) employed a PALENCIA rois's units in their on the stratigraphy of the papers In the northeastern ofthe of de part province Palencia, Asturian Devonian. Almela et al. (1956) and Garcfa- Sitter (1955) distinguished Devonian successions re- Fuente short ofthe Devonian (1959) gave a description ferred to as "la serie de Ventanilla", "la serie de stratigraphy in the explanation of their maps (1: Polentinos", and "la serie de San Julian" (Barruelo- 50,000). Llopis Llado (1960) published a stratigraphic Muda area). analysis of the area south of Oviedo, in which he The Serie de Ventanilla was discussed in more detail described facies changes in the Emsian and Eifelian by Kanis (1956), who described small bioherms rang- showing continental influences in the east. from Emsian Frasnian. Brouwer ing in age to The same author presented new evidence concerning (1964a) presented a description of these reefs and of the Silurian-Devonian boundary in the Furada sand- the biostromes of the Devonian in Leon. Other data stone and the distribution of the Lower Devonian in on the Serie de San Julian were provided by Wagner Asturias (Llopis Llado, 1965). (1955), Wagner & Wagner-Gentis (1963), Frets The ofthe Devonian has been the upper part sequence (1965), and de Sitter & Boschma (1966). subject of much controversy. In the eastern part of Kullmann described Devonian in (1960) a sequence Asturias the Alba Formation (= the Griotte), of the valley of the Arroyo Arauz, northwestofPolentinos, Visean is underlain thick of age, by a sequence which he dated by means of cephalopods. These rocks, quartzites. The ages assigned to this quartzitic forma- which consist mainly of shales, argillacious limestones, tion have varied from Carboniferous (Schulz, 1858; nodular in from and limestones, range age the Siege- Hernandez-Pacheco & Hernandez-Pacheco, 1935) nian to Frasnian. Part of the Emsian strata is called or Devonian (Barrois, 1882), to Silurian (5.1.) (Hernan- "Arruz Schichten". dez Sampelayo, 1928; Delepine, 1932a; Llopis Llado, Binnekamp (1965) published data on the stratigraphy 1958b; Julivert, 1960; Martinez Alvarez, 1962). Lot- of the Lower Devonian of the Polentinos and Arauz

ze (1957) postulated a Tournaisian (lower age described dated of areas, and as formations, by means limit not mentioned) for this quartzitic formation, brachiopods: which he called "Tina Serie".

Hernandez and Sampelayo Delepine, however, pro- Lebanza Formation (limestone) Siegenian duced palichnologic evidence, consisting of bilobites, Gedinnian Silurian for these pointing to a (5.1.) age quartzites (the Carazo Formation Silurian

trace fossils Cruziana furcifera d'Orbigny and Cruziana(quartzite, sandstone and shale) rogosa General outline of the stratigraphy 133

Frets (1965) recognized these formations in the Palencia and the Monto-Gildar area in northeastern

Barruelo-Muda area. Van Veen (1965), in tracing the Leon do not correlatewell with those of the Rio Luna- stratigraphy of the region between Polentinos and Rio Esla region in Leon and the Valsurvio area in

Cardano de Arriba, distinguished as formations lying Palencia. This led Brouwer (1964b) to distinguish two above described facies in the Devonian ofthe Cantabrian Moun- those by Binnekamp: types tains: Alba Griotte Formation (nodular, spotted Visean referred limestone) a) The Asturo-Leonese facies (elsewhere also

Vegamian Formation (black shale and chert) Tournaisian to as Leonide, Asturo-Leonian, Asturo-Leonesean, or

Vidrieros Formation (nodular limestone) _ Asturo-Leonesian of sandstones, Famennian facies), consisting and detrital limestones, and biostromes, containing a Murcia Quartzite Formation fauna of Cardano Formation (nodular limestone) Frasnian predominantly brachiopods, bryozoans, This facies Gustalapiedra Formation (dark limestones Givetian crinoids, corals, and stromatoporoids. and Couvinian in the of and shales) occurs Asturias, greater part Leon, Abadia Formation (shales and limestones) ., the Valsurvio and Ventanilla areas in Palencia. Emsian

b) The Palentine facies (elsewhere also referred to as Palencian Palentian of East of the Ventanilla area, Koopmans (1962) studied or facies), consisting mainly the stratigraphy of the Valsurvio region. He distin- shales and argillacious and nodular limestones. Only unit is i.e. the Mur- guished: one prominent quartzitic present,

cia Formation. The fauna has a more pelagic charac- Red Lower Carboniferous Griotte shale-griotte ter than the Asturo-Leonese facies. As characteristic

horizon elements it contains cephalopods, conodonts, tenta- Camporredondoformation Cremenes This facies in culites, and a few solitary corals. occurs (quartzite and sandstone) sandstone the northwestern of the northeastern part Palencia, Upper Devonian of Leon, and the southwestern of Santander. Valcovero formation Portilla part part The facies of the Barruelo-Muda in (shale and limestone) limestone area corresponds the Lower and Middel Devonian with the Hornalejo formation Middle Devonian Huergas mostly Palentine facies In the how- (shale and sandstone) shale and type. Upper Devonian, sandstone affinities the Asturo-Leonese facies be ever, to can formation Sta Otero Lucia observed.

limestone (limestone) In the Lower Devonian the differencebetween the two Compuertoformation Lower Devonian La Vid is but in the facies types not very pronounced, higher (shale and limestone) shale and of the Devonian a differentiationof the sedimen- limestone part basin becomes evident. Brou- tary into two facies areas

wer (1964b) argued that this difference in facies The third column shows Koopmans' correlation of his between corresponds more or less with the difference units with those of the Rio Esla area. (The term Cre- the Rhenan facies and the Bohemian (or Hercynian) menes sandstone was provisionally used for the sand- facies in northwestern Europe. Kullmann (1965) gave stone between the La Portilla and the Alba Formation an interesting account of the difference between the in this area.) In the northern part of the Valsurvio coral faunas ofthese two areas. He pointed out that the area the sequence becomes thinner and less complete cephalopod and coral fauna of the Palentine facies than in the south because of the progressive develop- area (his "Monto-Arruz-Zone") are impoverished ment of stratigraphic gaps. faunas.

A general review ofthe structural history of the Canta- GENERAL REMARKS brian Mountains and their relationship with the The Middle and Upper Devonian formations of the Pyrenees and the Celtiberic fold-belt has been given

Arauz-Polentinos and the Cardano-Triollo areas in by de Sitter (1965). 134 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

CHAPTER II

STRATIGRAPHIC AND PALAEONTOLOGIC OBSERVATIONS

central Fig. 2. Map (1 : 1,000,000) showing the location of the investigated areas. 1: the Río Esla area (Fig. 5); 2a—c: the

Gilder-Montó b: coastal area Cantabrian area (Figs. 14, 16, 21); 3: the area (Fig. 23); 4a, Asturias, (Figs. 32,

the Arauz-Polentinos 6: the 7: the 34); 5: area (Fig. 37); Cardaño-Triollo area (Fig. 42); Liébana area (Fig. 51).

Fig. 4. Legend to the distribution charts. The samples

from sections are consistantly indicated by capital

letters and sample numbers and spot samples by

lower-case letters and sample numbers, with the

exception of the Río Esla and the Liébana areas.

Fig. 3. Legend to the stratigraphic sections. Stratigraphic and palaeontologic observations 135

5. of the Río Esla area. The arrows the section sites toward the Fig. Locality map (1 : 100,000) indicating point

of the youngest part sequence.

LEÓN icriodids often had broken of the was damaged or off, The Rio Esla area which made their determination difficult if not im-

5 locality map, Fig. possible. Fig. 6 composite section, The Lower Carboniferous Alba Formation, on the sections, Figs. 7—12 contrary, yielded rich, well-preserved faunas. distribution chart, Enclosure no. 1 Sections VIII (Fig. 7), Ilf (Fig. 9) and X (Fig. 10) Table 1 ' sections AG were taken in the Esla autochthone, and and OL in the Esla The Devonian of the Rio Esla area is of the Asturo- (Fig. 11) (Fig. 12) nappe (Rupke,

facies contains rich This contains more Leonese type. It a macrofauna, 1965). nappe a complete develop- of the Devonian than the Esla autoch- among which brachiopods, corals, and stromatopo- ment Upper

roids elements. An extensive thone. Above the La Portilla Formation there is a are the most conspicious

ofthis has been of about 275 m of sandstones and shales in description area given by Rupke (1965). sequence into which detrital limestone band is inter- The dating of the stratigraphic sequence takes a prominent, i.e. the Cremenes Limestone. 1 account the work of Comte (1959) and of Dr. Th. F. calated, Samples rup and 2 taken from this limestone. The Krans (pers. comm.) who is making a study of the rup were sharp spiriferids from this region. contact between the Alba Formation and the karst

The conodont content of the Devonian limestones is surface of the Ermita Formation in section AG is

the low. The conodonts recovered from the shown in 13. Section SAL (Fig. 8) was taken in on average Fig.

detrital limestones were in general badly preserved. the Las Salas zone (Rupke, 1965), where the Ermita

The delicate ofthe Formation lies on of the La Vid Formation. posterior part expanded pulp cavity top 136 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Fig. 6. Composite section of the Río Esla autochtone. Fig. 7. Section VIII. Fig. 8. Section SAL.

Fig. 11. Section AG.

Fig. 9. Section IIf.

Fig. 12. Section OL.

Fig. 13. Contact between the Ermita and Alba Formations

in section AG. Karst crevices in the top of the

calcareous Ermita Formation are filled with red

detrital material belonging to the base ofthe Alba

Fig. 10. Section X. Formation. 138 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Table 1 Alphabetic index of the distribution chart of the Río Esla area (Enclosure 1)

45 Elictognathus lacerata (Branson & Mehl) 43 Pol. communis carina Hass

55 Gnathodus antetexanus Rexroad & Scott 44 Pol. cf. P. communis communis Branson & Mehl

59 G. bilineatus (Roundy) 24 Pol. decorosa Stauffer 58 G. commutatus commutatus (Branson & Mehl) 36 Pol. inornata E. R. Branson

65 G. commutatus cruciformis Clarke 11 Pol. linguiformis linguiformis Hinde

(> 62 G. commutatus homopunctatuss Ziegler Pol. linguiformis linguiformis Hinde (with large pulpcavity)

63 G. commutatus multinodosus Higgins 32 Pol. longipostica Branson & Mehl

64 G. commutatus nodosus Bischoff 35 Pol. nodomarginata E. R. Branson

51 G. cuneiformis Mehl & Thomas 25 Pol. normalis Miller & Youngquist

54 G. cf. G. cuneiformis Mehl & Thomas 42 Pol. pura subplana Voges

56 G. delicatus Branson & Mehl s.l. 21 Pol. varca Stauffer

60 G. cf. G. girtyi Hass 33 Pseudopolygnathus dentilineata E. R. Branson

49 G. punctatus (Cooper) 38 Ps. fusiformis Branson & Mehl 52 G. semiglaber Bischoff 37 Ps. multistriata Mehl & Thomas

53 G. typicus Cooper 40 Ps. triangula inaequalis Voges

23 Icriodus alternatus Branson & Mehl 48 Ps. triangula pinnata Voges

26 I. cf. I. alternatus Branson & Mehl 57 Scaliognathus anchoralis Branson & Mehl

16 I. corniger Wittekindt 46 Siphonodella cooperi Hass 15 I. cf. I. corniger Wittekindt 47 Si. obsoleta Hass

9 I. curvatus Branson & Mehl 39 Si. sulcata (Huddle)?

22 I. Branson & 41 Si. ? cymbiformis Mehl n.sp. a

20 I. eslaensis 27 aculeatus & n.sp. Spathognathodus (Branson Mehl)

1') I. expansus Branson & Mehl 18 Sp. bipennatus Bischoff & Ziegler

17 I. latericrescens latericrescens Branson & Mehl 61 Sp. campbelli Rexroad

4 I. latericrescens latericrescens Branson & Mehl 28 Sp. costatus costatus (E. R. Branson)

Usensu Ziegler, 1956) 2') Sp. costatus spinulicostatus (E. R. Branson) 8 I. latericrescens bilatericrescens Ziegler 30 Sp. costatus (E. R. Branson) subsp. indet.

10 I. nodosus 1 indet. (Huddle) Sp. sp.

14 I. symmetricus Branson & Mehl 2 Sp. inclinatus wurmi Bischoff & Sannemann

13 /. stabilis & sp. a 31 Sp. (Branson Mehl)

5 Neoprioniodus excavatus (Branson & Mehl) 12 Sp. steinhornensis steinhornensis Ziegler 3 Pelekysgnathus serrata ,Jentzsch 7 Sp. steinhornensis Ziegler subsp. indet.

34 Polygnathus communis communis Branson & Mehl 50 Sp. cf. S. strigosus (Branson & Mehl)

The central Cantabrian area 15, 19, 20, 22) were taken in the "Isidro-Tarna- Ponton subarea" 21 (Sjerp, 1967, Fig. 41). locality maps, Figs. 14, 16,

sections, Figs. 15, 17—20, 22

distribution chart, Enclosure no. 2

Table 2

The central Cantabrian area lies near the watershed

of the Cantabrian Mountains. Sections FE and OS

lie in southern discussed actually Asturias, but they are

here because they link up directly with the strati- graphy of Leon. little Devonian is the central Although very present,

Cantabrian area is considered to belong to the Asturo-

Leonese fades area because of the detrital lithology of

the Ermita Formation.

The sections investigated in this region show that the Palaeozoic. Ermita Formation rests directly on Lower

For a detailed description of this area, the reader is

referred to Sjerp (1967).

Sections RI (Fig. 17) and MA (Fig. 18) were taken in

the "Mampodre-Fontasguera-Ten subarea" (Sjerp,

1967, Fig. 41). The Ermita Formation is calcareous

in section RI. The Formation throughout Vegamian of the of Fig. 14. Locality map (1 : 100,000) western part in section MA contains two calcareous beds; this is the the central Cantabrian area. The arrow indicating

where we were able to extract conodonts section site toward the only place the points youngest part

and OS of the from this formation.Sections FE, U, PO, (Figs. sequence. Slratigraphic and palaeontologic observations 139

Fig. 17. Section RI.

Fig. 15. Section FE.

16. of the central of the central Cantabrian The the section Fig. Locality map (1 : 100,000) part area. arrows indicating

toward the of the sites point youngest part sequence. Adrichem Conodonts the Cantabrian Mountains 140 H. A. van Boogaert: of

Fig. 18. Section MA.

Fig. 20. Section PO.

19. Section U. Location of the of Fig. Fig. 21. map (1 : 100,000) eastern part

the central Cantabrian area. The arrow indicating

the section site points toward the youngest part of

the sequence. Stratigraphic and palaeontologic observations 141

Table 2 Alphabetic index of the distribution chart of the central Cantabrian area (Enclosure 2)

41 Cavusgnathus unicornis Youngquist & Miller 11 Pol. inornata E. R. Branson 21) Gnathodus antexanus Rexroad & Scott 8 Pol. longipostica Branson & Mehl

22 G. cf. G. antetexanus Rexroad & Scott 9 Pol. nodomarginata E. R. Branson

35 G. bilineatus (Roundy) 25 Pol. symmetrica E. R. Branson

33 G. commutatus commutatus (Branson & Mehl) 20 Pol. pura subplana Voges 38 G. commutatus cruciformis Clarke 13 Pseudopolygnathus dentilineata E. R. Branson

34 G. commutatus homopunotatus Ziegler 7 Ps. fusiformis Branson & Mehl

36 G. commutatus multinodosus Higgins 14 Ps. multistriata Mehl & Thomas

39 G. commutatus nodosus Bischoff 23 Ps. triangula pinnata Voges 27 G. Mehl & Thomas cuneiformis 16 Siphonodella cooperi Hass 30 G. cf. G. cuneiformis Mehl & Thomas 17 Si. isosticha (Cooper) 32 G. delicatus Branson & Mehl s.l. 18 Si. obsoleta Hass 40 G. girtyi Hass Si.? 19 n.sp. a 20 G. punctatus (Cooper) 1 Spathognathodus aculeatus (Branson & Mehl) 28 G. semiglaber Bischoff 37 Sp. campbelli Rexroad 31 G. typicus Cooper 2 Sp. costatus costatus (E. R. Branson) 24 Hindeodella segaformis Bischoff 3 costatus R. 10 Polygnathus communis communis Branson & Mehl Sp. spinulicostatus (E. Branson) 4 R. indet. 12 Pol. cf. P. communis communis Branson & Mehl Sp. costatus (E. Branson) subsp.

21 Pol. delicatula Ulrich & Bassler 5 Sp. stabilis (Branson & Mehl)

15 Pol. flabella Branson & Mehl 6 Sp. strigosus (Branson & Mehl)

The Gildar-Montó area

23 locality map, Fig.

composite section, Fig. 24 sections, Figs. 25 —30

Enclosure distribution chart, no. 3

Table 3

This area is situated on the southern slope of the

Valdeon valley. The Devonian of this area belongs to

the Palentine facies type, in which cephalopods and characteristic locally lamellibranchs are the macro-

OS. 23. of the Gildar-Montó Fig. 22. Section Fig. Locality map (1 : 100,000)

area. The arrows indicating the section sites point

toward the of the youngest part sequence. 142 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Fig. 26. Section CU I.

Fig. 24. Composite section of the Gildar-Montóarea.

Fig. 28. Section CU II.

Kutterink Leidse Geol. will fossils. (in prep., Med.)

give a structural description of the area.

The oldest lithostratigraphic unit in this area is a lime-

stone band which, because of its lithologic properties

and its place in the sequence, is considered to be the

Polentinos Memberofthe AbadiaFormation. Six ofthe

seven samples taken fromthis limestone did not contain

conodonts; the seventh, sample cal 1, yielded two

specimens, one ofwhich could be tentatively identified

as a juvenile specimen ofIcriodus corniger. Higher in the

succession the nodular limestones generally yielded conodont faunas. 31 rich, well-preserved Fig. shows a slump of dark-grey, laminated, quartzitic sandstone in

Fig. 25. Section CAL I. the Vegamian Formation near section CAL II. Stratigraphic and palaeontologic observations 143

Fig. 27. Section CAL II.

Fig. 30. Section POS. Fig. 31. Slump of dark-grey, laminated, quartzitic sand- Formation stone in the Vegamián near section CAL II (Photo by Kutterink). 144 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

32. of the of Fig. Locality map (1 : 100,000) western part

Asturias, coastal area. The arrow indicating the

section site toward the of points youngest part

the sequence.

Fig. 33. Section PE.

29. Section CU III. 34. of the of Fig. Fig. Locality map (1 : 100,000) eastern part

Asturias, coastal area. The arrow indicating the

section site points toward the youngest part of the

sequence. Stratigraphic andpalaeontologic observations 145

Table 3 index of the distribution chart of the Gildar-Montó Alphabetic area (Enclosure 3)

26 Ancyrodella curvata (Branson & Mehl) 20 Pol. asymmetrica ovalis Ziegler & Klapper

25 Ancyrodella lobata Branson & Mehl 53 Pol. communis communis Branson & Mehl

28 Ancyrognathus asymmetrica (Ulrich & Bassler) 66 Pol. cf. P. communis communis Branson & Mehl

27 Ancyrognathus triangularis Youngquist 17 Pol. cristata Hinde 77 Cavusgnathus unicornis Youngquist & Miller 23 Pol. decorosa Stauffer

71 Gnathodus bilineatus (Roundy) 62 Pol. delicatula Ulrich & Bassler

70 G. commutatus commutatus (Branson & Mehl) 21 Pol. dengleri Bischoff & Ziegler

75 G. commutatus cruciformis Clarke 45 Pol. diversa Helms

72 G. commutatus homopunctatus Ziegler 38 Pol. glabra glabra Ulrich & Bassler

74 G. commntatus nodosus Bischoff (if) Pol. inornata E. R. Branson

67 G. cuneiformis Mehl & Thomas 1 Pol. linguiformis linguiformis Hinde

69 G. delicatus Branson & Mehl s.l. 10 Pol. linguiformis mucronata Wittekindt

76 G. girtyi Hass 63 Pol. longipostica Branson & Mehl

68 G. typicus Cooper 46 Pol. nodocostata nodocostata Branson & Mehl 31 Icriodus alternatus Branson & Mehl 61 Pol. nodomarginata E. R. Branson

1 I. corniger Wittekindt 29 Pol. normalis Miller & Youngquist 12 I. curvatus Branson & Mehl 22 Pol. ordinata Bryant

Mehl 14 I. cymbiformis Branson & 8 Pol. pennata Hinde

3 I. eslaensis 47 Pol. n.sp. pennatuloidea Holmes

15 I. expansus Branson & Mehl 9 Pol. pseudofoliata Wittekindt 11 I. nodosus (Huddle) 6 Pol. cf. P. robusticostata Bischoff & Ziegler

13 I. symmetricus Branson & Mehl 39 Pol. subserrata Branson & Mehl 78 Mestognathus beckmanni Bischoff 36 Pol. cf. P. triphyllata (Ziegler)

41 Mehl Palmatolepis distorta Branson & 5 Pol. varca Stauffer 40 Pal. glabra glabra Ulrich & Bassler 7 Pol. xyla Stauffer

43 Pal. glabraelongata Holmes 64 Pseudopolygnathusfusiformis Branson & Mehl

42 Pal. glabra pectinata Ziegler 18 Schmidtognathus peracuta (Bryant) 49 Pal. gracilis gracilis Branson & Mehl 55 Spathognathodus aculeatus (Branson & Mehl)

(,() Pal. gracilis sigmoidalis Ziegler 48 Sp. amplus (Branson & Mehl) 34 Pal. hassi Miiller & Miiller 73 Sp. campbelli Rexroad

35 Pal. minuta minuta Branson & Mehl 56 Sp. costatus costatus (E. R. Branson) 44 Pal. perlobata schindewolfi Miiller 57 Sp. costatus spinulicostatus (E. R. Branson)

37 Pal. quadrantinodosa marginifera Ziegler 58 Sp. costatus ultimus Bischoff

32 Pal. subrecta Miller & Youngquist 54 Sp. jugosus (Branson & Mehl) 24 Pal. transitans Miiller 52 Sp. inornatus (Branson & Mehl) 33 Pal. unicornis Miller & Youngquist 16 Sp. cf. S. sannemanni sannemanni Bischoff & Ziegler

Paltodus 2 sp. 50 Sp. stabilis (Branson & Mehl) 30 Miiller Miiller Polygnathus amana & .",! Sp. strigosus (Branson & Mehl)

10 Pol. Bischoff & 59 asymmetrica asymmetrica Ziegler Sp. supremus Ziegler

ASTURIAS, COASTAL AREA gether with light-brown to amber, undamaged speci-

34 mens. is a mixed fauna so locality maps, Figs. 32, Only rarely distinctly repre- sections, Figs. 33, 35 sented.

distribution 36 chart, Fig. The main quartzite body in section LL (Fig. 35) on Table 4 the Piedras Luengas beach is considered to be the

Ordovician "Ojiartcita Armoricana" (= Barrios the Asturian Along coast a few sections were studied Formation). We follow here Hernandez Sampelayo for comparison with the formations in Leon. The (1928), Delepine (1932a), Llopis Llado (1958b), and Devonian in this area belongs to the Asturo-Leonese others. We do not accept the concept of the Tina facies Section PE from the de Serie for which type. (Fig. 33) Playa (Lotze, 1957), no palaeontologic proof

Carranques near Perlora contains a clearly distinguish- has been put forward (see also Ch. I, p. 132). The detrital limestone band the Pineres 12 ofthe which able, separating uppermost m quartzite, consists ofvery Sandstone and the Alba Formation. In analogy with coarse-grained quartzite, is assigned to the Ermita other of the Cantabrian Mountains in parts we assign Formation, analogy with the northernpart of Leon. this limestone band to the Ermita Formation, which in Further arguments for this classification are given in this is almost case completely calcareous. From the Chapter IV (p. 160). The completely unconsolidated baseof the Alba Formation(sample PE 4) an abundant, sandy and argillacious beds intercalated between the mixed fauna obtained. Attrited conodonts and was coarse-grained quartzite and the Vegamian Formation,

to- conodonts encrusted with iron oxide were found are remarkable. This unit yielded no macrofossils, and 146 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

palynological examination gave negative results. PALENCIA this unit the Ermita Provisionally, we assign to Forma- The tion. The Alba Formation, which overlies the Vega- Arauz-Polentinos area

mian Formation, begins with a lens of soft, grey clay. 37 locality map, Fig.

composite section, Fig. 38

sections, Figs. 39, 40

distribution chart, Fig. 41

Table 5

The Devonian of the Arauz-Polentinos area is of the

Palentine facies type. In the Gedinnianand Siegenian but in brachiopods are abundant, the younger stages

cephalopods and trilobites are the most characteristic

macrofossils. The area has been described by Binne-

kamp (1965), van Veen (1965), and de Sitter &

Boschma (1966).

The oldest conodont fauna (sample ar 1) comes from

detrital limestone lens in the a very locally developed,

upper part of the Carazo Formation (Member c).

Samples bk 13 and pol 23 were taken from a nodular

limestone lens or tongue in the upper part of the

Abadia Formation. This limestone occurs only 3.5 km

west-northwest of Polentinos in a tectonically very

disturbed area. The Polentinos Member of the

Abadia Formation yielded few conodonts. Of the

eleven samples studied, only two (pol 9 and pol 11) Formation yielded a usable fauna. The Gustalapiedra

in this area is in general badly exposed, which hampers

its investigation.The locality ofsection POL II (Fig. 40) is the of the Cardano For- only well-exposed outcrop

mation in this area. Fig. 35. Section LL.

coastal Fig. 36. Distribution chart of Asturias, area. Stratigraphic and palaeontologic observations 147

Table 4 index of the distribution chart of coastal Alphabetic Asturias, area (Fig. 36)

27 Gnathodus antetexanus Rexroad & Scott 15 Pol. inornata E. R. Branson

23 G. cuneiformis Mehl & Thomas 11 Pol. longipostica Branson & Mehl

26 G. cf. G. cuneiformis Mehl & Thomas 13 Pol. nodomarginata E. R. Branson 25 G. delicatus Branson & Mehl s.l. I)! Pseudopolygnathus dentilineata E. R. Branson

22 G. punctatus (Cooper) 20 Ps. triangula pinnata Voges 24 G. semiglaber Bischoff 21 Scaliognathus anchoralis Branson & Mehl

28 G. typicus Cooper 19 Siphonodella obsoleta Hass 1 Icriodus alternatus Branson & Mehl 5 Spathognathodus aculeatus (Branson & Mehl)

2 I. cymbiformis Branson & Mehl (. Sp. costalus costatus (E. R. Branson)

12 Polygnathus communis communis Branson & Mehl 7 Sp. costalus spinulicostatus (E. R. Branson)

17 Pol. communis carina Hass 8 Sp. costalus ultimus Bischoff

16 Pol. cf. P. communis communis f Branson & Mehl 9 Sp. costalus (E. R. Branson) subsp. indet.

3 Pol. decorosa Stauffer 10 Sp. inornatus (Branson & Mehl) 4 Pol. delicatula Ulrich & Bassler 11 Sp. stabilis (Branson & Mehl)

37. of the Arauz-Polentinos and San Martín The the section sites Fig. Locality map (1 : 100,000) areas. arrows indicating

toward the of the point youngest part sequence. 148 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Table 5 Alphabetic index of the distribution chart of the

Arauz-Polentinos area (Fig. 41)

f> Hindeodella priscilla Stauffer

18 Icriodus corniger Wittekindt

23 I. cymbiformis Branson & Mehl

22 I. eslaensis n.sp

15 I. expansus Branson & Mehl

3 I. latericrescens latericrescens Branson & Mehl {:sensu

Ziegler, 1956)

4 I. latericrescens bilatericrescens Ziegler

14 I. nodosus (Huddle)

1 I. woschmidti Ziegler

8 Neoprioniodus bicurvatus (Branson & Mehl)

9 Ozarkodina typica denckmanni Ziegler

7 Pelekysgnathus serrata I Jentzsch

12 indet. Plectospathodus sp.

19 Polygnathus angusticostata Wittekindt

20 Pol. angustipennata Bischoff & Ziegler

17 Pol. linguiformis linguiformis Hinde

21 Pol. varca Stauffer

16 Pol. webbi Stauffer

10 Spathognathodus inclinatus wurmi Bischoff & Sannemann

11 Sp. steinhornensis steinhornensis Ziegler

2 Sp. steinhornensis Ziegler subsp. indet.

13 Trichonodella symmetrica (Branson & Mehl)

5 A n.gen. n.sp. a

Fig. 38. Composite section of the Arauz-Polentinos area.

Fig. 39. Section POL I. Fig. 40. Section POL II. Stratigraphic and palaeontologic observations 149

Fig. 41. Distribution chart of the Arauz-Polentinos area.

T he Cardaño-Triollo area cleavage (Fig. 48). Conodonts recovered from the

cleaved limestones are sometimes strongly deformed Reality map, Fig. 42 Some of deformed faunas (Plate 2, Figs. 5, 6). these are composite section, Fig. 43 bleached to a light-grey colour. Because of tectonic sections, Figs. 44—47 complications it was impossible to obtain a complete,

attribution no. 4 chart, Enclosure reliable section ofthe Vidrieros Formation. The basal

Table 6 of this unit is little calcareous in the northern part so of the that conodont faunas The part area no adequate TV 0nian of the Cardano-Triollo also could be extracted there. In the of the bel area uppermost part 0 tQ l^e P a'ent:ine facies The area has been Vidrieros Formation in section TR, samples were taken desc 'K type. m detail van Veen ( 1 965 The formations close together to obtain dataconcerning the Devonian/ are by )' of° te n folded and affected Carboniferous Ch. 157—158). strongly isoclinally by boundary (see Ill, pp. 150 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

42. Location of the Cardaño-Triollo Fig. map (1 : 100,000) area. The arrows indicating the section sites point toward

the of the youngest part sequence.

Table 6 index of the distribution chart of the Cardaño-Triollo Alphabetic area (Enclosure 4)

30 Ancyrodella buckeyensis Stauffer 2 I. cymbiformis Branson & Mehl

25 curvata & 8 I. eslaensis Ancyrodella (Branson Mehl) n.sp.

20 Ancyrodella lobata Branson & Mehl 18 I. expansus Branson & Mehl

14 Ancyrodella rotundiloba rotundiloba (Bryant) 17 I. nodosus (Huddle) 32 Ancyrodella asymmetrica (Ulrich & Bassler) 5 I. cf. I. obliquimarginatus Bischoff & Ziegler 26 Ancyrognathus triangularis Youngquist 21 I. symmetricus Branson & Mehl

4 Belodus sp. 48 Palmatolepis distorta Branson & Mehl 73 Gnathodus bilineatus (Roundy) 28 Pal. gigas Miller & Youngquist

74 G. commutatus commutatus (Branson & Mehl) 41 Pal. glabra glabra Ulrich & Bassler 80 G. commutatus cruciformis Clarke 40 Pal. glabra elongata Holmes

77 G. commutatus homopunctatus Ziegler 35 Pal. gracilis gracilis Branson & Mehl

79 G. commutatus multinodosus Higgins 31 Pal. hassi Muller & Muller

78 G. commutatus nodosus Bischoff 42 Pal. minuta minuta Branson & Mehl

72 G. delicatus Branson & Mehl s.l. 53 Pal. minuta schleizia Helms

75 G. girtyi Hass 44 Pal. perlobata schindewolfi Muller

64 G. kockeli Bischoff 49 Pal. perlobata sigmoidea Ziegler

G. A Scott & Rexroad 43 Pal. 65 sp. Collinson, quadrantinodosa marginifera Ziegler

29 Icriodus alternatus Branson & 50 Pal. Mehl rugosa ampla Muller

1 I. corniger Wittekindt 27 Pal. subrecta Miller & Youngquist

22 I. curvatus Branson & Mehl 23 Polygnathus amana Muller & Muller Stratigraphic andpalaeontologic observations 151

Table 6 Continued

36 Pol. communis communis Branson & Mehl 34 Pol. subserrata Branson & Mehl

Pol. Stauffer 12 Pol. cristata Hinde 7 varca

19 Pol. decorosa Stauffer 52 Scaphignathus velifera Ziegler

l(, Pol. dengleri Bischoff & Ziegler 9 Schmidtognathus hermanni Ziegler

38 Pol. diversa Helms 10 Sch. peracuta (Bryant)

6 Pol. cf. P. eiflia Bischoff & Ziegler II Sch. wittekindti Ziegler

71 Pol. flabella Branson & Mehl 66 Pseudopolygnathus dentilineata E. R. Branson

39 Pol. glabra glabra Ulrich & Bassler 60 Ps. fusiformis Branson & Mehl

47 Pol. glabra bilobata Ziegler (17 Ps. multistriata Mehl & Thomas

70 Pol. inornata E. R. Branson 55 Spathognathodus aculeatus (Branson & Mehl)

3 Pol. linguiformis linguiformis Hinde 45 Sp. amplus (Branson & Mehl) 62 Pol. longipostica Branson & Mehl 69 Sp. anteposicornis Scott

33 Pol. nodocostata nodocostata Branson & Mehl 76 Sp. campbelli Rexroad

51 Pol. P. nodocostata Branson & Mehl 56 R. ex. gr. Sp. costatus costatus (E. Branson)

61 Pol. nodomarginata E. R. Branson 57 Sp. costatus spinulicostatus (E. R. Branson) 24 Pol. normalis Miller & Youngquist 63 Sp. costatus ultimus Bischoff

68 Pol. cf. P. obtecta Branson & Mehl 58 Sp. costatus (E. R. Branson) subsp. indet.

13 Pol. pennata Hinde 54 Sp. inornatus (Branson & Mehl)

37 Pol. cf. P. pennatuloidea Holmes 59 Sp. stabilis (Branson & Mehl) 15 Pol. cf. P. Huddle 46 (Branson & rugosa Sp. strigosus Mehl)

Cardaño-Triollo 44. Section CAR I. Fig. 43. Composite section of the area. Fig. 152 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Fig. 45. Section VID.

Fig. 47. Section TR.

Fig. 48. Cleaved nodular limestone of the Vidrieros For-

mation in section CAR II. The handle of the

Fig. 46. Section CAR II. hammer lies perpendicular to the cleavage plane. Stratigraphic andpalaeontologic observations 153

metres The San Martín-Valsurvio area few of quartzitic sandstone of the Ermita For-

mation. The shale is followed 4 m of nodular 42 by red, locality map, Figs. 37, limestone 1 from the section, Fig. 49 (griotte). Sample cam comes distribution chart, Fig. 50 basal part of this limestone. The succession continues with of nodular limestone which the 4 m grey, on dark,

Only a few observations were made in this region, fine-grained, well-bedded limestone of the Escapa Formation follows. which is part of the Asturo-Leonese facies area. The of the transition We have taken this revised properties Devonian/Carboniferous opportunity to present a

were studied near San Martin de los Herreros and version of the conodont fauna mentioned by Koop- around the Pantano de for full from E 81 in the Valcovero Camporredondo; a mans (1962, p. 146) locality

description, the reader is referred to Kanis (1956) and Formation.

Koopmans (1962). The Alba Formation in section MAR (Fig. 49) begins limestone. South of almost directly with Triollo, near the Pantano de Camporredondo, this formation starts with 2 m of red, somewhat cherty shale overlying a

49. Section MAR. 51. of the Liébana Fig. Fig. Locality map area.

Fig. 50. Distribution chart of the San Martín-Valsurvio area. 154 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

SANTANDER, THE LIÉBANA AREA

51 locality map, Fig. composite section, Fig. 52

distribution chart, Fig. 53

Table 7

The Devonian in the Liebana is exposed in a tectoni- anticlinal cally strongly disturbed, structure, sur- rounded by Upper Carboniferous formations. It be- forma- longs to the Palentinefacies type. The Devonian tions deformed and the are too outcrops too scarce to allow reliable measurement of complete stratigraphic sections. Mr. R. W. who the Lanting, mapped area,

provided a number ofsamples from which the forma- tions could be dated and strati- present roughly a graphic column could be reconstructed (Fig. 52). The oldest from this derives sample area, sample 85,

from a succession of dark-grey limestones and black

shales. Lithologically, this unit shows affinitieswith the

Gustalapiedra Formation, but its isolated occurrence

makes it uncertain whether it belongs to this forma- tion. The Alba Formation is absent in the Liebana.

Only a few, thin, detrital limestone bands occur in the

Potes Formation, i.e. the rock unit that overlies the

Vegamian Formation.

52. Fig. Composite section of the Liébana area.

53. Distribution chart of the Liébana Fig. area. Stratigraphic and palaeontologic observations 155

Table 7 Alphabetic index of the distribution chart of the Liébana area (Fig. 53)

7 Ancyrodella curvata (Branson & Mehl) 16 Pal. perlobata schindewolfi Muller 10 Ancyrognathus asymmetrica (Ulrich & Bassler) 3 Pal. subrecta Miller & Youngquist 33 Gnathodus bilineatus 12 (Roundy) Polygnathus amana Muller & Muller 32 G. commutatus commutatus (Branson & Mehl) 27 Pol. communis communis Branson & Mehl

34 G. commutatus homopunctatus Ziegler 13 Pol. decorosa Staufler

G. nodosus Bischoff 35 commutatus 17 Pol. diversa Helms

5 Icriodus alternatus Branson & Mehl 2 Pol. linguiformis linguiformis Hinrle 11 I. cf. I. alternatus Branson & Mehl 18 Pol. nodocostata nodocostata Branson & Mehl 1 I. corniger Wittekindt 4 Pol. normalis Miller & Youngquist 6 I. cymbiformis Branson & Mehl 19 Pol. cf. P. pennatuloidea Holmes 14 Palmatolepis distorta Branson & Mehl 26 Scaphignathus velifera Ziegler 8 Pal. gigas Miller & Youngquist 28 Spathognathodus aculeatus (Branson & Mehl) 20 Pal. glabra glabra Ulrich & Bassler 21 Sp. amplusr (Branson & Mehl) 15 Pal. glabrapectinata Ziegler 29 costatus costatus (E. R. Branson) 23 Pal. gracilis gracilis Branson & Mehl Sp. R. 9 Pal. hassi Muller & Muller 30 Sp. costatus spinulicostatus (E. Branson)

& 24 Pal. minuta minuta Branson & Mehl 31 Sp. inornatus (Branson Mehl)

25 Pal. minuta schleizia Helms 22 Sp. strigosus (Branson & Mehl)

CHAPTER III

ZONATION AND DATING OF THE STRATIGRAPHIC SEQUENCE BY MEANS OF CONODONTS

THE BIOSTRATIGRAPHIC FRAMEWORK & erous of this basin is in preparation (Mauvier Lys,

Geol. Univ. in the The conodont zones established in Europe by Boger Mem. Inst. de Louvain, prep.), from the (1962), Voges (1959, 1960), Walliser (1962, 1964), correlation of this part of the Carboniferous

Kulm facies have to be Wittekindt (1966), and Ziegler (1962b, 1966), proved Ardennes and the area may be valuable revised to a basis for arranging and dating the to some extent. into conodontfaunas obtainedfrom the Cantabrian Moun- The bilineatus Range Zone is split up two assem- tains. Enclosure bilineatus-delicatuss.l. Zone is character- 5 shows the conodont zones referred blage zones.The

to in this A few remarks about this chart ized the of Gnathodus bilineatus and Gna- paper. are by concurrence required. Apart from the woschmidti Zone, well-defined thodus delicatus s.l. without Gnathodus commutatus nodosus,

conodont have established in the and be divided into lower and an zones not yet been can a upper part by the absence of Gnathodus Lower Devonian. A rough subdivisioninto "Conodont- presence or semiglaber and/or Bereiche" has been given by Walliser (1962). The Gnathodus typicus and/or Gnathodus cuneiformis and/or bilineatus Zone semi- upperGivetian transversa Zone could not be demonstrat- Gnathodus antetexanus (compare + Ed in the Cantabrian Mountains. Polygnathus lingui- glaber + texanus of Boger, 1962). defined the formis transversa Wittekindt, 1966 was never found in The bilineatus-commutatus nodosus Zone is by

faunas with number of which of Gnathodus bilineatus and Gnathodus com- our Polygnathus varca, a concurrence and must an It is mutatus nodosus; it can be divided into a lower an certainly have upper Givetian age.

the absence of Gnathodus conceivable that the distribution of Polygnathus lingui- upper part by presence or formis transversa is restricted to northern Europe. We delicatus s.l.

refer to the combined of the Zone and the The relationofthese to the Visean and Namurian range varca zones

transversa Zoneof Wittekindt(1966) as the varca Zone s.l. Stages is tentatively derived from the literature mentioned Enclosure The comparison of the Lower Carboniferous zones (see 5). with the Tournaisian and Visean stages, as presented DEMONSTRATED IN THE here, is partly after Voges (1959) and Boger (1962) CONODONT ZONES CANTABRIAN MOUNTAINS and partly after Conil, Lys, and Mauvier (1964). The is latter authors showed that the anchoralis Zone is The lowermostGedinnian woschmidti Zone represent-

1 from limestone lens in the present in the Franco-Belgian Basin in assise Tn 3b ed in sample ar a upper Since detailed of the Carazo Formation (upper Tournaisian). a more report on part (Arauz-Polentinos the repartition of conodonts in the Lower Garbonif- area) by Icriodus woschmidti. 156 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Icriodus latericrescens latericrescens From the Middle Devonian well-defined (sensu Ziegler, 1956) on, conodont

and Icriodus latericrescens bilatericrescens are generally zones are available. The corniger Zone is represented by in the Devonian dis- Icriodus well represented Lower samples corniger in the uppermost part of the Santa cussed below. Lucia Formation in the Rio Esla the Polentinos area, conodont faunas from lower of Vid The the part the La Member of the Abadia Formation in the Arauz- Formation Esla and the Lebanza Forma- Polentinos Cardano-Triollo and (Rio area) area, the area, proba-

tion (Arauz-Polentinos area) do not contain conodonts bly in the Gildar-Montoarea. The oldest sample from for of the Conodont-Bereiche. the Liebana diagnostic one They area, sample 85, (Gustalapiedra For-

are characterized by Pelekysgnathus serrata, which is mation?), also belongs to this zone. The lower be especially abundant in the lower part of the above- part ofthe bidentatus Zone may represent- mentioned units. Its is Ed in in which Icriodus and exact range not known; sample ar 3, corniger

Jentzsch (1962) indicated a Lower Devonian age in Polygnathus angustipennata were questionably identified. Thuringen, whereas Le Fevre (1965) mentioned it The robusticostata Zone is probably represented in

from the "Devonian basal" (Gedinnian) in France sample CAR 34 from the Gustalapiedra Formation and the Sahara. showed the which contains cf. Binnekamp (1965) on (Cardano-Triollo area), Polygnathus basis of that the lower ofthe Lebanza P. with This is brachiopods part eiflia together Polygnathus varca. zone

Formation has an upper Gedinnian to lower Siegenian probably also represented in spot sample cu 20 from and that the has middle the Formation in the Gildar-Monto age upper part a Siegenian Gustalapiedra

So serrata age. Pelekysgnathus ranges well into the area, which yielded Polygnathus cf. P. robusticostata and

Siegenian. Polygnathus varca.

Conodont-Bereich X is in the The Zone s.l. is demonstratedin thebasal of probably represented varca part from the samples the uppermost part of the basal lime- La Portilla Formation (Rio Esla area) by Polygna-

stones of the La Vid Formation Esla be- thus Icriodus latericrescens latericrescens and (Rio area), varca, Spatho-

cause in this level Polygnathus linguiformis linguiformis gnathodus bipennatus. Wittekindt (1966) reported Icriodus Plate latericrescens from lower bank (with a large pulp cavity; see 3, Fig. 1) appears the with ”Terebratula” for the first time. Bar-type and blade-type conodonts pumilio in the discoides-Kalk in the Hercynian Givetian,

are absent, however. just above the ”walliseri Horizont”. Spathognathodus The faunas from the Requejada Member of the bipennatus is known only from the ”Sparganophyllum-

Abadia Formation which forms the basal of the Rhenan (Arauz-Polentinos area) may Kalk”, part either Conodont-Bereich X Conodont- belong to or to upper Givetian ("obere Stringscephalenstufe"). The Bereich XI. lack fauna collected from basal of the They Polygnathus linguiformis lingui- brachiopod the part but show few formis, bar-type conodonts. The cepha- La Portilla Formation points to a middle Givetian of the Th. lopod content Requejada Member, according age (Dr. F. Krans, pers. comm.).

to Kullmann (1960, Arruz Schichten), points to a The varca Zone s.l. is also well represented in the lower Emsian after age. However, visiting the Arauz upper part of the Gustalapiedra Formation and the

area Prof. Dr. O. H. Walliser lower of the (pers. comm.), came to part Cardano Formation in the Gildar- the conclusion that the from this unit Monto in the of the cephalopods area, upper part Gustalapiedra indicate Emsian final Formation an upper age. Until a solution in the Cardano-Triollo area, and in the of this problem has been reached, we shall indicate the Cardano Formation in the Arauz-Polentinos area. In age of the Requejada Member as lower? Emsian. In these areas this zone is characterized only by Poly- 38 and Enclosure 6 this member is Fig. still placed in gnathus varca. It seems probable that the distribution

the lower Emsian. of Icriodus latericrescens latericrescens and Spathognathodus

Conodont-Bereich XI is represented in the limestone bipennatusis controlled by the facies. Theabundance of band in the lower of the Icriodus eslaensis in the part predominantly shaly varca Zone s.L is remarkable.

portion of the La Vid Formation (Rio Esla area). It is The hermanni-cristata Zone comprises the transition remarkable that from this only two samples level from Middle to Upper Devonian. It is most proba-

yielded Polygnathus linguiformis linguiformis. Bar-type bly represented in sample CAR 30 from the lowermost and blade-type conodonts are absent. This Conodont- part of the Cardano Formation (Cardano-Triollo

Bereich is possibly also represented in the nodular area), which contains various species ofSchmidtognathus. limestone lens in the Abadia Formation or tongue Although sample CU 3a from the Cardano Formation

(Arauz-Polentinos in the Gildar-Montoarea lacks it area). Schmidtognathus, may The ofthe La Vid this upper part Formation and the basal belong to zone. part of the Santa Lucia Formation (Rio Esla area) The Lower asymmetrica Zone is demonstratedin samples characteristic yielded as species Icriodus cf. I. corniger. cu 6 and cu 18 from the lower to middle part of the This Dr. O. Walliser species, according to H. (pers. Cardano Formation in the Gildar-Monto area and in in the Rhenan Emsian lower CAR from the basal of comm.), occurs upper to sample 29 part the Cardano Couvinian. Therefore, the upper part of the la Vid Formation (Cardano-Triollo area). Formation and the basal of Santa The well in part the Lucia For- gigas Zone is represented the upper part mation be in the of the Cardano Formation in the Gildar-Monto may probably placed uppermost area, Emsian in the Hercynian sense. in the Cardano-Triolloarea, and in the Liebana area. donation and dating by conodonts 157

The faunas from this zone contain ancyrodellids, It is difficult to locate the Devonian/Carboniferous

Ancyrognathus triangularis, Palmatolepis gigas, and Palma- boundary exactly by conodonts. The upper and lower The of limits of the of the Devonian and tolepis hassi. presence Ancyrognathus assymmetrica ranges Upper the

makes it possible to delimit the Upper gigas Zone. Lower Carboniferous zonal guide forms in Europe are In the facies Frasnian known The detailed work of Asturo-Leonese area aberrant not precisely. Voges conodont faunas recovered from the and leaves small in were coarse- (1959) Ziegler (1962b, p. 42) a gap for grained, detrital limestones from the uppermost part the sequence not tested conodonts, corresponding

of the La Portilla Formation (Rio Esla area) and the to the uppermost Devonian Hangenbergschiefer.

limestone from the uppermost part of the Pineres Henningsen (1965) describes a continuous Upper

Sandstone (Asturias, coastal area). These faunas lack Devonian — Lower Carboniferous succession in

the conodonts diagnostic for the Frasnian zones, such calcareous rocks from the Lahn syncline (Germany). the This is ”Gatten- as genera Ancyrodella, Ancyrognathus, and Palmato- section, however, highly condensed, the icriodids and few 8 and therefore lepis. Only a polygnathids are pre- dorfia-Stufe” being only cm thick,

sent. The distribution of the first of these Frasnian a certain degree of condensation is to be expected be facies. A middle in the faunas. generaappears to dependent on the

Frasnian age must be assigned to the upper part ofthe The conodont zonation of the Upper Devonian and Portilla Formation in of the of the La the Rio Esla area because Mississippian Upper Mississippi Valley

the occurrence in it of Mucrospirifer bouchardi (Mur- given by Collinson el al. (1962) cannot be applied the Cantabrian faunas because chison) (Dr. Th. F. Krans, pers. comm). directly to several of

The Cremenes Limestone (Rio Esla area) also yielded the zonal forms mentioned are not present in the

a similarly aberrant fauna. Westbroek (1964) suggests, latter. The important species Gnathodus kockeli, original- the basis ofits of from on rhynchonellid fauna, that the age ly reported Germany (Bischoff, 1957), occurred

this limestone could fall around the boundary between in only one of our lowermost Carboniferous faunas.

the Frasnian and Famennian. Dr. Th. F. Krans It is difficult to explain to what extent this absence is

favours Frasnian related to the because even in the nodular lime- (pers. comm.) an uppermost age on facies, basis of its the spiriferid contents. stones of the Vidrieros Formation this gnathodid was The quadrantinodosa Zone is well represented in the generally missing.

lower part of the Vidrieros Formation in the Gildar- An additional problem in the Asturo-Leonese facies Monto the Cardano-Triollo is the and area, area, and the area presented by transgressive regressive

Liebana area. Apart from the occurrence of Palma- nature of the thin Ermita Formation, as a result of which be tolepis quadrantinodosa marginifera the concurrence of mixed faunas would expected. Stratigraphic Palmatolepis glabra glabra and Palmatolepis distorta points admixtures could be caused by reworking of older, to this zone. pre-Ermita strata, or stratigraphic leaks may have The Zone is in occurred the of the Ermita which velifera poorly represented sample at top Formation,

CAR 22 from the Cardano-Triollo area. A good fauna locally shows karst features (Fig. 13). Krebs (1964) this is has discussed the of mixed belonging to zone found in sample LV 435 from frequent occurrence cono- the Liebana area. The presence of Palmatolepis glabra dontfaunas in sequences with hiatusses and in condens- with glabra together Scaphignathus velifera in this sample ed sequences. suggests that it belongs to the Lower velifera Zone. Section TR (Fig. 47) in the Vidrieros Formationin the Cardano-Triollo suitable The Upper styriaca Zone to Lower costatus Zone is area is highly for the study probably represented in sample CAL 12 from the ofthe Devonian/Carboniferous transition in the Canta- Vidrieros in brian Mountains. TR 15 TR Formation the Gildar-Monto area with Samples 20 yielded

amount Spathognathodus jugosus. The Lower to Middle costatus an ample of conodonts of the Spathognathodus

Zone is indicated in the same area by sample kutt 77. costatus group and Palmatolepis gracilis gracilis, and with

The Middle to Upper costatus Zone is well represented the exception of sample TR 15, they also contain in the of the Vidrieros Formation in the fusiformis. Sample TR 21 upper part Pseudopolygnathus provided of the Gildar-Montoarea, the Cardano-Triolloarea, and the two specimens Spathognathus costatus group. Liebana The from this contain Gnathodus kockeli and Gnathodus A Rex- area. samples part as sp. Collinson, which indicate either the diagnostic elements Spathognathodus aculeatus, Spathogna- road & Scott, 1961, very thodus costatus costatus, Spathognathodus costatus spinuli- uppermost Devonian (Gnathodus n. sp. A. Assemblage the costatus, Polygnathus longipostica, and/or Polygnathus Zone of Collinson et al., 1962) or, more probably, nodomarginata. The Middle to Upper costatus Zone is lowermostTournaisian kockeli-dentilineata Zone. Sample but shows also represented in the detrital limestone in the upper TR 22 contains no gnathodids, Pseudopoly- of Ermita Formation in sections and dentilineata and fair number of part the AG, Ilf, gnathus a Spathognathodus

X in the Rio Esla sections and OS in the it lacks inornata. TR 23 area; FE, U, aculeatus; Polygnathus Samples in the TR 25 also lack all contain central Cantabrian area; and section PE gnathodids. They as

Asturian coastal area. It should be noted that Palma- conspicious elements Polygnathus inornata and Poly- in whereas it The of tolepis gracilis gracilis is absent these areas, gnathus longipostica. occurrence one juvenile is in the Palentine facies This of in frequent area. appears specimen Spathognathodus anteposicornis sample again to be controlled by the facies. TR 23 is remarkable. It is clear from their position in 158 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

the succession above sample TR 21 that samples TR 22 sented in sample SJ/H6 from section MA (central

TR 25 should all be placed in the kockeli-dentilineata Cantabrian area), which yielded an abundant fauna

Zone, in spite of the absence of zonal guide forms. including gnathodids and siphonodellids. This fauna

communis communis is well contains broken them a Polygnathus represented many conodonts, among great the of section TR. throughout upper part number of reworked specimens. this shall consider the faunas with The anchoralis Zone is well in the basal In paper we represented in Polygnathus inornata, Polygnathus longipostica, numerous part of the Alba Formation in the Rio Elsa area, the communis and few central Cantabrian in Asturias Polygnathus communis, occasionally a area, (coastal area),

Spathognathodus aculeatus and/or Spathognathodus costatus and in section MAR (San Martin-Valsurvio area). Tournaisian in costatus as being of lowermost age and be- Scaliognathus anchoralis is not always present the longing to the kockeli-dentilineata Zone. The higher samples from this zone, but the concurrence of with Tournaisian zones are again well defined by the genus Gnathodus semiglaber and Gnathodus delicatus Pseudo- inornata communis Siphonodella. Polygnathus already occurs scantily polygnathus triangula pinnata and/or Polygnathus absence in the costatus Zone. communis are also diagnostic for this zone. The is of in these faunas The kockeli-dentilineata Zone so interpreted represent- or scarcity siphonodellids suggests ed in of the Vidrieros Formation that the of the anchoralis the uppermost part they belong to upper part in section CAL II in the Gildar-Monto area and in Zone. sections CAR II and TR in the Cardano-Triollo area. The anchoralis-bilineatus Interregnum is represented in

This is also demonstrated in the the lower of the Alba Formation in the same areas zone uppermost part part of the detrital limestone of the Ermita Formation in as those in which the anchoralis Zone is demonstrated. sections AG and Ilf (Rfo Esla area). The few speci- These areas belong to the Asturo-Leonese facies area. of bilineatus-delicatus s.l. Zone is in the mens ofGnathodusI in sample Ilf223 and GnathodusI and The represented in AG 5 assumed have leaked of the Alba Formation in the sections Siphonodella sample are to upper part This in from the base of the overlying Alba Formation. studied in the above-mentioned area. zone was Tournaisian lower also demonstrated in the basal and lower of the These conodonts have a late to part

but in the Alba Formation in section POS and Visean age, they only occur uppermost (Lower Upper bilineatus-delicatus s.l. in the Gildar-Monto samples from the sections and no trace ofthe conodont Zone) area

TR zones between the kockeli-dentilineata Zone and the and section (Upper bilineatus-delicatus si. Zone) in Cardano-Triollo Siphonodella crenulata Zone was found. The kockeli- the area.

dentilineata Zone is also represented in section U (cen- The bilineatus-commutatus nodosus Zone is well repre- Alba tral Cantabrian area) and section PE (Asturias, coastal sented in the upper to uppermost part of the Formation in of the where this area). In sections MA and RI (central Cantabrian most areas part was in area) all the samples from the limestone in the Ermita investigated. The basal part of the Alba Formation and CU III Formation appear to belong to the kockeli-dentilineata sections CAR II (Cardano-Triollo area) Zone. Unfortunately, these conodont faunas from the (Gildar-Monto area) belongs already to the Lower and

Ermita Formation are and contain bilineatus-commutatus nodosus Zone, poor many Upper respectively. damaged specimens. The bilineatus-commutatus nodosus Zone is also present in of the anchoralis is detrital limestone band in the Liebana The probably lower part zone repre- a area.

CHAPTER IV

DISCUSSION OF THE NEW DATA WITH RESPECT TO THE DEVONIAN AND THE LOWER CARBONIFEROUS

FORMATIONS IN THE CANTABRIAN MOUNTAINS

in the literature and our observations are The new stratigraphic data mentionedin the preceding the data

have increased about the collected. The ruled of the columns on chapter our knowledge age vertically parts

and distribution of the Cantabrian formations. This the chart indicate stratigraphic gaps. authors whose data into this material will be discussed on the basis of the time- The are incorporated

which chart indicated in Table 8. stratigraphic correlation chartof Enclosure 6, in are 159 Discussion of the new data

Table 8 Literature consulted for the construction of the time-stratigraphic correlation chart (Enclosure 5)

Llado column Adrichem et al. column 1. Barrois (1882); Comte (1959); Llopis 10. van Boogaert (1963); Sjerp (1958a, 1962); Radig (1961, 1962). (1967).

Llado column 2. Delepine (1928a); Llopis (1960). column 11. Delepine (1932a); Hernandez Sampelayo column 3. Almela et al. (1956); Budinger & Kullmann ' (1928); Llopis Llado (1958b). (1964); Delepine (1928a); Garcia-Fuente column 12. van Adrichem Boogaert (1965); Budinger & (1959); Grosch (1912). Kullmann (1964); Kullmann (1960, 1961). column 4. Smits (1965).

column 13. van Adrichem Boogaert (1965); van Veen column 5. Comte (1959) ;Higgins ttal.{ 1964); Lotze( 1961); (1965). Wagner (1963). column 14. Mr. R. W. column 6. Comte (1959); Rupke (1965). Lanting (unpublished). column 15. Kullmann de Sitter column 7. Koopmans (1962). Binnekamp (1965); (1960); column 8. Brouwer (1964a); Kanis (1956). & Boschma (1966). column 16. & column 9. van Adrichem Boogaert et al. (1963); Julivert Frets (1965); Wagner (1962); Wagner (1960); Martinez Alvarez (1962); Sjerp (1967). Wagner-Gentis (1963).

DEVONIAN (and Lowermost Tournaisian) This section consists of 2 to 3 metres of calcareous,

yellowish to white sandstone (decalcified in the out-

The Asturo-Leonesefacies area crop).

Few new data can be added to the fundamental work

— of Comte (1959) on the Devonian formations of this Lithology and thickness. The lithology ofthe forma- facies area. The conodont faunas recovered from the tion is very variable. The main constituent is medium-

La Vid, Santa Lucia, and La Portilla Formations in to coarse-grained, quartzitic sandstone. Decalcifica-

to the Rio Esla area do not lead essential changes con- tion has often caused a pronounced porosity. The colour from via cerning their age, the only modification being that the can vary white, rose, to a true ferrugi-

Siegenian/Emsian boundary is placed somewhat lower nous red.

in the sequence. The correlation of the Asturian with Lenses and layers of coarse-grained, rose or grey the Leonese Lower and Middle Devonian formations limestone often in the of the are developed upper part in has not yet been completely unravelled, but detailed Ermita Formation the western and southwestern of the and of investigation lithology a modern study part of the CantabrianMountains. The sandstone pass-

the brachiopod and microfossil contents will solve this es gradually into this limestone. Locally, the detrital of problem. limestone forms the dominant rock type the forma-

The calcareous part of the Ermita Formation yielded tion (section RI, Fig. 17; section PE, Fig. 32). A small conodont faunas which view interesting permit a new layer of dispersed pebbles is often found at the base of the definition and distribution of this unit. of the Ermita Formation (section U, fig. 19; section

LL, Fig. 35). The Ermita Formation. The Ermita For- The thickness is also variable. In the transgressive type area which Comte studied the Comte and the mation, (1959) throughout designated by (1959) near upper course

Rio - Rio Esla and in the of of Rio Curueno and the Rio Porma Bernesga area uppercourse (Mr. H.J. Evers, the Rio is in various other of thickness of than 100 is Tuiza, recognized parts pers. comm.) a more m the Cantabrian mountain chain (see Enclosure 5). attained. In most other areas the Ermita Formation

has a thickness of 1 to 40 m. The bedding is usually

Definition and section. — This formation massive type was not distinct, giving a appearance to the forma- originally defined by Comte as "Gres de l'Ermitage". tion. Occasional crossbcdding is observed. The section chosen located type was on the eastern bank of the Rio Bernesga near the Ermita del Buen Age. Comte (1959) described fossiliferous beds at in Suceso, the vicinity ofthe village ofHuergas. Accord- the top of the Ermita, containing as diagnostic ele- ing to Comte's description, the formation consists here ments: Spirifer verneuili Murchison, Cleiothyris royssii of 140 m of yellowish or rose quartzitic sandstone, Leveille, Camarotoechia letiensis Gosselet, and Pugnax in beds of variable thickness moresnelensis Koninck. He that (Comte, 1959, p. 193). pointed out especially

recent of this i.e. the moresnetensis indicates an Famennian However, investigation area, Pugnax uppermost Alba showed that its and sedi- In lower of syncline, stratigraphic (= Strunian) age. the part the Ermita, in

mentological properties make it unsuitable as type area places where this formation attains considerable of the Ermita Formation. that found Therefore, we suggest thickness, Gomte only Spirifer verneuili and the section of this unit be the section Camaroloechia which do type changed to letiensis, not permit an equally south of from which Comte determination. Camplongo, (1959, p. precise age In the southern part of the his first ofthe Ermita Formation. Alba he with 166) gave description syncline, near Peredilla, identified, some Conodonls 160 H. A. van Adrichem Boogaert: of the Cantabrian Mountains

reservations, Cleiothyris lamellosa Leveille, which might A thin pebble bed, passing into a more coarse-grain- indicate Tournaisian affinities even (Comte, 1959, ed and less quartzitic sequence in the uppermost 20 m, often indicates the of the Ermita Formation. p. 315). presence Ermita The brachiopods from the Ermita Formation near This kept in mind, the distribution of the can

the watershed of the Cantabrian Cordillera, identified be investigated beyond the Rio Bernesga-Rio Esla

by Krans (van Adrichem Boogaert et al., 1963), also region into Asturias. Comte (1959, p. 295) already Famennian clue indicate an uppermost age. A subsequently gave a by stating: "Dans la region d'lnfiesto,

fauna from the de dans tout le bassin sampled, well-preserved brachiopod pres Rozapanera et meme

Riosol area (section RI, Fig. 13)), however, points to a inferieur du Rio Sella, il est fort possible, a en juger

Tournaisian Th. F. Krans, les caracteres des des possible lower age (Dr. par lithologiques, que gres famenniens les pers. comm.). quartzites transgressifs accompagnent Ermita Conodont faunas from the basal part of the quartzites siluriens. Dans le meme ordre d'idees, des

Formation in section PE (Fig. 23) from the Asturian quartzites roses qui couronnent les quartzites siluriens

indicate Famennian coeur meme des Picos de meritent de retenir coast an uppermost age (upper- au Europa — most to V to VI). The basal fauna from the Ermita l'attention"(In the region ofInfiesto, near Rozapanera

Formation in section RI (Fig. 17) located in the Riosol and even throughout the lower basin of the Rio Sella, which is from the area points to a lowermost Tournaisian age, it highly possible, judging lithological indicates that the base of the Ermita Formation is properties, that transgressive Famennian sandstones

diachronous. This feature is difficult to study in more and quartzites accompany the Silurian quartzites. In of Ermita Formation that the detail, because the basal part the the same sense, the rose quartzites crown

is rarely calcareous, making it impossible to extract Silurian quartzites in the very heart of the Picos de conodonts from it. Europa should be kept in mind).

of the studied, the ofthe difficult to on the basis of fossils be- In most areas uppermostpart Although prove in Ermita Formation extends into the lowermost Tour- cause they rarely occur in the rocks question, for Comte's be Where lime- naisian. Higgins el al. (1964) even concluded the top assumption seems to correct. Aviados section late Tournaisian beds the Ermita Formation could oftheir a age. Sample stone were present,

AG 5 from section AG (Fig. 11) in the Rio Esla area be ascertained with conodonts. Tournaisian but In the northeastern of Hernandez also seems to point to an upper age, part Asturias, for the this & Kindelan reasons already explained (Ch. 111, p. 158), Sampelayo (1950, p. 60) distinguished 6—20 sample is assumed to be oflowermost Tournaisian age, on the Lower Palaeozoic quartzites m conglom- material leaked in from the and sandstones with thin some younger having erates coarse-grained a overlying Alba Formation. bed of coal containing distinguishable plant remains.

They assigned this formation to the Lower Devonian

Distribution and the underlying and overlying forma- on lithological grounds. Our opinion is, however, that

tions. — Because of its transgressive character, the these rocks, too, belong to the transgressive Ermita Formation Ch. and Ermita Formation overlies rocks of widely differing (see II, p. 145 Fig. 35). In the southern of the Cantabrian Moun- The conodont faunas mentioned & Kull- ages. part by Budinger in Asturias tains it lies on Famennian rocks of the Nocedo Forma- mann 423 from Entrago (1964, pp. —425) tion. A hiatus between these rocks and the Ermita is revealed the of the Ermita Formation with presence a re-inter- here often difficult if not impossible to establish. In all calcareous development in this area. Our that probability the hiatus disappears completely in the pretation of the section from this locality suggests southern and southwestern directions. samples 1 —3 correspond to the Ermita Forma- Formation. It is therefore not surprising that in the lithologic suc- tionand samples 4—7 to the Alba

cession of the southern Alba syncline the Ermita For- Sample 4, taken from the base of the Alba Formation

Devonian in which lies the Ermita mation cannot be discerned. The Upper disconformably on Formation,

this consists of of sandstone and shale contains a mixed fauna llf224 and OL 1 area a sequence (cf. samples with calcareous intercalations. We attribute this kind from the Rio Esla area and PE 4 from section PE, of lithology to the Nocedo Formation. The Ermita Fig. 29). Pseudopolygnathus dentilineata and Pseudopoly- forms with thick Formation grades into this lithology in the northern gnathus triangula inaequalis, which are considered be reworked part of the Alba syncline, where we suggest that an platforms, are to specimens arbitrary cut-off be drawn between the Ermita from older strata (stratigraphic admixture). The

Formation and the Nocedo Formation. The hiatus rest of the fauna points to the anchoralis Zone of upper-

its in of Tournaisian lower Visean This leaves the reaches maximum the northernmost part most to age. the where the Ermita overlies established of Gnathodus Gnathodus Leon, near watershed, ranges delicatus, the Lower Cambrian dolomites of the Lancara girtyi, Gnathodus semiglaber, and Gnathodus texanus

Formation (section RI, Fig. 17). All formations in unchanged, and explains why "das Unter-Tournai. . . between can be overlain by the Ermita. Where the auf Grand von Goniatiten trotz grosster Bemuhungen

Ermita overlies the Ordovician Barrios Quartzite nicht festzustellen war" (Budinger & Kullmann, 1964,

Formation it is often difficult, and sometimes very p. 416). thickness of about even impossible, to distinguish the disconformity plane. The limestone band with a 20 m 161 Discussion of the new data

in be omitted here. few remarks are which underlies the griotte of the Alba Formation may Only a required 1960, 50, with to their and distribution. the Barranco de Las Xanas (Llopis Llado, p. respect age

PI. 5) is thought to be the calcareous part of the

Ermita Formation. The Abadia Formation. This unit consists ofshale and

with limestone members, There are indications that the Ermita Formation also sandy shale two prominent

in the Asturo-Leonese facies of Palencia. The and in thickness from about 100 to 300 m. Van occurs ranges is based section sections of Kanis (1956) and Koopmans (1962) show Veen's description (1965) on a along and the Rio which be considered section. that the Camporredondo Formation (quartzite Arauz, can as type

Ventanilla and the northern The formation there has a thickness of 180 and its sandstone) in the area part m, is to Near of the Valsurvio dome becomes thinner towards the shaly to sandy part usually grey dark-grey.

of the the Abadia itself the structures to north. Koopmans mentioned that at the top are too complicated reliable of section. Camporredondo Formation, locally, microconglom- permit measurement a complete of Another section is located the northern erates or coarse sandstones with a thickness 1 —2 m complete along

Besande and south of the border of the Pantano de 2.75 km south- occur, for example near Requejada, A. Pantano de Camporredondo. In analogy with the southeast of Polentinos, for which Mr. J. van ofabout 295 for the development of the Upper Devonian in the Rio Hoeflaken reported a thickness m of Esla it be assumed that the Abadia Formation. Here, the non-calcareous part Bernesga-Rio area, may the formation consists of to brown shales uppermost part of the Camporredondo Formation fine, yellow intercalations. corresponds to the Ermita Formation. without sandy

the lower i.e. the bulk of the The lowest limestone member is the Mem- Consequently, part, Requejada of "Arruz-Schich- Camporredondo Formation, (Camporredondo For- ber. This unit is the equivalent the mation be correlated with the Nocedo ten" of Kullmann 465 We consider s.s.) can (1960, pp. —466). lower? Emsian Formation. Where the quartzitic sandstone that the age ofthe Requejada Member to be Member in underlies the Visean griotte (Alba Formation) is only (see Ch. Ill,p. 156). The Requejada occurs

at the Rio Arauz-Polentinos area and the Barruelo- a few metres thick, as in the section taken San and in the Muda Martin de los Herreros (Kanis, 1956) sec- area (Kullmann, 1960, pp. 466, 469). The limestone unit of the Abadia Formation is tions east of Pena Negra, Alba de los Cardanos, and upper of Triollo (Koopmans, 1962), the whole quartzitic unit the Polentinos Member, which is the equivalent considered Ermita Formation. Schicht 5 of Kullmann and Schicht 5 can be as (1960, p. 465) of and Schicht 6 of Kullmann The Ermita Formation is overlain in most parts ("Encimero Schichten")

ofan eastern Asturias and northern Leon by the Vega- (1965, p. 45), which, according to Kullmann, are

contact Eifelian The conodont content of two samples mian Formation. The between these forma- age. spot

tions is sharp and often slightly erosional. Where the point to a lower Eifelian age. The Polentinos Member is found in the Arauz-Polen- Vegamian Formation does not occur, the Ermita For- the Gildar-Monto mation is overlain by the Alba Formation. Since the tinos area, the Cardano area, and

in the Barruelo-Muda is not contact between these formationsis disconformable, a area. Its occurrence area

rocks oflower Eifelian can small amount of erosion took place locally. known. In the Liebana, age these should be to the To summarize, it may be stated that the Ermita be traced, but possibly assigned of Formation. Formation is widely distributed over a large part Gustalapiedra

the Cantabrian Mountains in Leon, Asturias, and been de- Palencia. Although the Ermita Formation cannot The Gustalapiedra Formation. This unit has Veen from the Car- always be demonstrated with palaeontologic evidence, scribed and denned by van (1965)

consists of to black shales and lithologic criteria are very often available to confirm dafio area. It dark-grey

section. The Ermita Formation limestones with thickness of 30 to 70 its presence in a may a m. follows the lower Eifelian Polentinos Member vary in age from upper Famennian to lowermost Since it the lower ofthe Gustala- Tournaisian, and is overlain disconformably by the ofthe Abadia Formation, part be of later Eifelian Vegamian Formation or the Alba Formation. The piedra Formation must a age. Veen in the Cardano Ermita Formation does not occur in the Palentine Cephalopods collected by van identified Kullmann from facies area. area and by (1963b) range

lower to Givetian. The conodont faunas from upper of the Formation in the upper part Gustalapiedra The Palentinefacies area this Givetian section CAR I agree well with macro-

Conodonts from the Carazo Formation (Member c) fauna.

and from the Lebanza Formation confirm, or at least The Gustalapiedra Formation occurs in the Cardano

of formations de- the Arauz-Polentinos area (Schicht 6 of Kull- do not contradict, the age these as area, and the Gildar-Monto This fined by Binnekamp (1965). The distribution of these mann, 1960, p. 465) area.

demonstrated unit has not been for the Barruelo-Mudaarea. two formations has been adequately reported and Veen de- by Frets (1965) van (1965). Lithologic formations of the Palentine The Cardano Formation. The Cardano Formation has scriptions of the younger

defined and described from the Cardano area facies have been published by van Veen (1965) and been by 162 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Veen It consists of and brownish The Vidrieros van (1965). grey Formation. This unit has been defined calcareous shale, nodular limestone, and bedded and described by van Veen (1965). It consists of thickness of limestone, attaining a 10 to 40 m. Cepha- nodular limestone, calcareous shale, and pure shale

found van Veen in the and deter- with thickness of 10 lopods by type area a to 100 m. mined by Kullmann (1963b) point to a Frasnian age. This unit is the equivalent of the "Monto Schichten" Schicht 7 ofKullmann the Car- of Kullmann the Gildar- (1960, p. 465) represents (1960, pp. 472—473) from dano Formation in the northern part ofthe Arauz area. Monto area. On the basis of their cephalopod contents, conodontfaunas recovered from the Cardano For- Kullmann The established their range in the Famennian mation permit a precise age determinationofthis unit. fromWocklumeria-the Cheiloceras-Stufe (to 11(3) to the The Cardano Formation in section CAR I in the Kalloclymenia-Stufe (to VI). Budinger & Kullmann

Cardanoarea shows a range in age from the lowermost (1964) confirmed this by conodonts.

to upper Frasnian (in the German notation: lower Our oldest conodontfaunas from the Vidrieros Forma- to la lower to IS). The Frasnian conodont faunas tion in the Cardano-Triollo area and the Gildar- from the Cardano Formation in the Liebana fit well Monto from the of the lower area range upper part In the Gildar-Monto in the same range. area the Famennian to the lowerpart ofthe middleFamennian

Cardano Formation starts earlier and ranges from (upper to lip — lower to Ilia). The conodont faunas Givetian Frasnian middle to upper to upper (lower mentioned by Frets (1965, p. 121) for localities 3 and The Cardano Formation in the 11 Santibanez de well extend down to IS). western near Resoba, may Arauz-Polentinos POL is into the of the lower Famennian. area (section II) exclusively uppermost part middle Givetian This be lo- of to upper age. may only a The upper part of the Vidrieros Formation yields cal conodont faunas of Famennian development. an uppermost age Formation is known from — The Cardano not the Bar- (uppermost to V to VI). The uppermost beds of ruelo-Muda A shale of Frasnian is indicated the Vidrieros Formation in Cardano-Triollo area. age the area

the of Aulatornoceras bicostatum and in the northern of the Gildar-Monto by presence (Hall) men- part area tioned & by Wagner Wagner-Gentis (1963). range into the lowermost Tournaisian.

The Vidrieros Formation can be traced from the Car-

The Murcia Formation. This described formation, by dafio-Triolloarea to the Gildar-Monto area. South of Veen consists of van (1965), mainly light to dark-grey the Pico Gildar and west of Barniedo, the Vidrieros quartzitic sandstones with a variable but mostly small Formation is sometimesnon-calcareous locally (section ofshales intercalated. Some beds amount graded occur. CU III, Fig. 29; Mr. H. Teer, unpublished internal

The thickness of the formationranges from 40 to pro- report). The Vidrieros formation is also demonstrated bably 200 m. in the Liebana area.

The lower of the Murcia Formation small The of Vidrieros in part yields possible occurrence the Formation lamellibranchs which of Buchiola the Barruelo-Muda is evidence among specimens area supported by identified were provisionally by van Veen (1965). reported by Kullmann (1960), who described nodular Frasnian Because of the limestones with fauna They suggest an upper age. a cephalopod pointing to an lack of the of the limitof the Murcia Famennian from 500 fossils, age upper upper age (to V) a locality m

Formation can be deduced from the of the west of Muda. A conodont fauna found 700 north only age m which the Vidrieros of Verbios Frets shows directly-overlying beds, belong to by (1965, p. 121) an upper to

Formation. This unit in — starts the upper part of the upper Famennian age (upper to V to VI). Older lower Famennian the lowermost of the middle Famennian limestones or part are not known for this region; Famennian. The Murcia Formation therefore ranges possibly no full development of the Vidrieros Forma- between the Frasnian and lower Famen- uppermost tion exists in the Barruelo-Muda area. Wagner &

nian to IS — to Its of existence of (upper Up). rate deposition was Wagner-Gentis (1963) suspect the a hiatus about 5 to 10 times faster than that of the underlying between the Moradillo quartzite and the Famennian and overlying nodular limestone formations. limestone, which they call the Verbios Formation. The

The Murcia Formation be traced from the Car- ofrock and the biofacies ofthis Verbios Formation can type dano-Triollo the Gildar-Monto The area to area. pre- differ strongly from the Ermita Formation. in sence this area oftwo quartzite formations separated band ofnodular limestone of by a to lip age, as report- LOWER CARBONIFEROUS ed & Kullmann is by Budinger (1964, p. 419), not confirmed by our observations. The Murcia Formation The sharp lithological and palaeontological difference also occurs in the Liebana, though in reduced thick- between the Asturo-Leonese facies and the Palentine ness (40—60 m, Mr. R. W. Lanting, pers. comm.). In facies disappears in the Lower Carboniferous. None- this the area underlying and overlying Devonian theless, a different development can be traced, as indi- formations also show areduced thickness, but without cated by the following discussion of the formations. evidence of breaks in the sedimentation. The Vegamian Formation occurs in the Palentine It that the MoradilloFormation of seems probable the facies area and in a part of the Asturo-Leonese facies

Barruelo-Muda can be correlated with The Alba Formation almost area partially area. occurs everywhere the Murcia Quartzite. throughout the Cantabrianmountain chain. A revision Discussion of the new data 163

Alba based the of the Vegamian and Formations, on the upper part near Genicera, Lingula, Orbiculoidea,

literature and our observations, is presented below. trilobites, ostracods, lamellibranchs, and squashed

goniatites which seem to belong to Pericyclus. The upper of the black shales in section OS (Fig. 22) yielded The Vegamián Formation part small brachiopods, a few trillobite pygidia, ostracods,

Definitionand — The Formation type area. Vegamian and badly-preserved cephalopods. Some of the Comte was originally designated by (1959, p. 330) as cephalopods belong to the genus Ammonellipsites "Couches de Vegamian", consisting of about 15 m (= Pericyclus auct.), pointing to Tournaisian or "anthracitic" shales few upper black, containing a phosphate lower Visean, according to Budinger and Kullmann nodules in the which is situated 1 km type locality, (1964, p. 421, footnote). south-southwest of Vegamian. Van Ginkel (1965) The conodont faunas from the basal sandy bsds of the presented a revision of this formation, which he also de las Villas and Vegamian Formation near Santiago distinguished in the Palentine facies area. Genicera middle Tournaisian point to a to upper age,

according to Higgins et al. (1964). In th: uppsr part of Lithology and thickness. The main constituent of the Vegamian Formation they found Pseudopolygnathus

the formation is black shale, but a variety of accessory triangulapinnata, which ranges from upper Tournaisian

rock occur. nodules are common in the types Phosphate to lower Visean.

formation. In the Cardano area van Veen (1965) ob- The calcareous beds in the lower part ofthe Vegamian served thin "phosphate bearing lenticular laminae". Formation in section MA (Fig. 18) point to an upper- The occurrence of black, thin-bedded chert is also Visean most Tournaisian to lower age (lower cu common in the Vegamian Formation, in which very np/y). radiolarians have been found. Gildar-Monto In the section CAR In the Palentine facies area, II (Fig. 46) the of the formation is area part cherty, upper very located near Cardano de Arriba shows a gradual lighter in colour, and often finely laminated. Locally, transition of the underlying Vidrieros Formation into

the cherts have a greenish or reddish tinge. At the top the black shale of the Vegamian Formation. The of the formation, near Barniedo, concretions of iron- highest nodularlimestones of the Vidrieros Formation stone occur (Mr. J. F. Savage, oral. comm.). Ironstone which contain conodonts, already have a very dark concretions and laminated ironstone beds are present colour; it follows from this that the Vegamian Forma- in the Vegamian Formation in the Liebana. Some tion here begins early in the Tournaisian. Other

localities show the development of pyrite or marcasite sections located in the Palentine facies area (e.g. TR, concretions, e.g. sections U (Fig. 19) and OS (Fig. 22). Fig. 47; CAL 11, Fig. 27; and CU 11, Fig. 28) show a Higgins el al. (1964) described from the base of the clear lithologic break between the Vidrieros Forma-

Vegamian Formation near Genicera and Olleros de tion and the black shales of the Vegamian Formation. thin bed with nodules Alba, a sandy phosphatic and determinationsof For the latter, we have no age the its in the of de las near top, neighbourhood Santiago basal The direct indication for the portion. only age Villas, a sandstone with a conglomeratic base. Section of the black shales in the Palentine facies area is given MA contains basal bed with (Fig. 18) a glauconitic Kullmann who by Budinger & (1964, p. 421), sandstone. Van Veen (1965, p. 63) observed two mentioned a few shell fragments from Schicht 6 in the lenses of in the Formation. quartzitic pebbles Vegamian of Goniatites Monto area (Goniatites) ex gr. striatus, Some slumped beds of laminated quartzitic sandstone belonging to the upperVisean (cu Illa-P (?)). It is not are intercalated (Fig. 31). known whether this upper Visean age of the Vegamian The black shales often have a brilliant, coally appear- to Formation also applies the Cardafio-Triolloarea. in the Gildar-Monto and in ance, e.g. area eastern Formation The overlying Alba presents a sharp con- Asturias. When this type of black and very brittle tact with the Vegamian Formation in the Palentine shale is it releases a combustible heated, vapour the facies area, clearly indicating a break in sedimen- which smells of bitumen and burns with a yellow tation. Its lower has Visean part already an upper age flame. In black general, the shales are not calcareous. (cuIII).

Only in section MA (Fig. 18) do a true argillacious In the Asturo-Leonese facies area the Vegamian limestone bed and a calcareous, sandy, breccious bed Formation is also overlain by the Alba Formation, occur. in this in the Tournaisian which begins area uppermost The Vegamian Formation is generally thin but shows to lower Visean (cu lip/y). Often, there exists a thin wide variation, the maximum development being zone of gradual transition between these formations found in the Palentine facies area. Near Cardano de 1959, there (e.g. Comte, p. 331). Locally, however, are Veen found thickness of Arriba, van (1965) a 30 m. with clear Olleros sharp contacts a disconformity (e.g. South ofPico Gildar, a thickness of 50 m was measured de Alba and Santiago de las Villas, Higgins et al., (section CU III, Fig. 29). 1964). These considerations show that the and lower upper and of the Formation Age underlying and overlying formations. The boundaries Vegamian are diachro- black shales of the Formation and that its fullest took in the Vegamian generally nous, development place show few fossils. for Wagner (1963, p. 210) mentioned Palentine facies area. 164 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

— Distribution. The distribution of the Vegamian show that this unit is a thin, marine, diachronous for-

Formation in the Asturo-Leonese facies area is rather mation deposited in a shallow basin of limited extent.

irregular. It has been reported from the central part It starts in a rather small area in the lower Tournaisian, Cantabrian Mountains be observed Arriba. It of the near the Tarna Pass as can near Cardano de greatest and the Ponton Pass and from southwest and distribution occurred in the Tournai- an area probably upper northeast of this central region, in Leon and Asturias. sian, when it occupied not only the Palentine facies Formation does in the San but also ofcentral northeastern Asturias The Vegamian not occur area part and

Martin-Ventanilla area, the Valsurvio area, or the and part of the Leonese area. In the lower Visean and

has it been for the lower of the Visean the Rfo Esla area, nor reported western part upper deposition of the

ofLeon and Asturias. 65 ofthe Formation seems have been restricted part Fig. gives a picture Vegamian to

possible maximum distribution of the Vegamian to the Palentine facies area. The formation is underlain

Formation as compiled from the literatureand our ob- by the Ermita Formation in the Asturo-Leonese facies

servations. The variation in thickness and the local area and by the Vidrieros Formation in the Palentine of formation have been caused in facies The formation is absence the may area. overlying the Alba local erosion of this unit the is in the central part by prior to deposition Formation, which, however, not present of the Palentine facies of the overlying Alba Formation. In the larger areas part area.

where no trace of the Vegamian Formation is found,

it was probably never deposited. The Alba Formation

The Vegamian Formation is present throughout the Definitionand — The Alba Formation type area. was Palentine facies and is absent in the central and area, Comte originally designated by (1959, p. 40) as eastern part of the Barruelo-Muda area. "Griotte de Puente de Alba" or "Griotte a (Goniatites

Frets and van Veen (1965) observed that in the (1965) crenistria” (op. cit., p. 330). In the type section near Cardano-Triolloarea near the Leon Lineand Cardano Puente de Alba in the Rio Bernesga valley, the forma- Line, both the Vegamian and Alba Formations are tion consists of about 30 m of red nodular limestone often not developed. Even the Vidrieros Formation is (griotte). The formation has been often referred to as locally absent, in which case the Escapa Formation "the Griotte" or "Alba Griotte". The present author directly overlies the MurciaFormation. In the Liebana, prefers the name Alba Formation, because ofthe varia- the Vegamian Formation passes gradually into an ble lithology of the unit here described. undated shale-graywacke sequence, the Potes Forma- lens R. W. tion, containing a conglomerate (Mr. Lithology and thickness. — The most diversifieddevel- Lanting, comm.). pers. of the Alba Formation shows from opment top to bottom: Depositional environment. — The old belief that

relatively deep water is always necessary for the devel- red, nodularlimestone (griotte), of black shales been opment has disproved by the becoming upwards more calcareous observations of black mud accumulation in and thus into the many very grey, passing literature — shallow water (see review by Conant& Swan- Escapa Formation: 8 22 m cherts and shales: son, 1961). The Vegamian Formation is undoubtly a red bedded 2 —12 m

marine sediment, as shown its fossil contents. It is nodular or by grey, wavy-bedded,

clearly a transgressive unit, and must have spread finely crystalline limestones: 0.5 — 4 m flat This character rapidly over a area. transgressive shallow-water environment for this forma- This is found in the central Cantabrian suggests a sequence

tion, a view supported by the occurrence of several Mountains, e.g. sections FE, MA, U, PO, and OS

lenses in the formation and the fact 18 — Similar sections also found conglomerate by (Figs. 15, 20, 22). are

that the top of the Vegamian Formation can also be a in the Rio Bernesga, Rio Torio, Rio Curueno, and Rio

disconformity plane. The anoxic bottom conditions Porma valleys and section MAR (Fig. 49) in the San in the Sea be Martin The Alba Formation with Vegamian might explained by some- area. begins directly

what restricted conditions, combined with high plank- red marls and red nodular limestonesin the type area, the surface and Rio Esla and in of northern ton production in water consequent the area, parts Asturias,

high oxygen consumption in the waters below. e.g. section PE (Fig. 33) and LL (Fig. 35). The basal Formation in of formation consist of red shales and red The Vegamian many respects closely beds the resembles the Shale de- cherts in other the northernValsurvio Chattanooga extensively some areas, e.g.

scribed by Conant & Swanson (1961), who proposed area (locality cam 1), in the Monte del Naranco and

as environment for the Chattanooga Shale a shallow the Playa de San Pedro (Radig, 1964) in Asturias. The

than 100 but fossil of the lower of the Alba quiet sea (probably not deeper feet, content part Formation, indicates that base of this unit in the locally, close to the shore, only a few feet deep). The se- however, the

diment is believed to have been transported by slow Asturo-Leonese facies area has approximately the

currents over the bottom ofthe sea. According to these same age (cu II(3/y) everywhere. Accurate mapping sedimentation basal of the Alba Formation authors, the rate of was extremely slow. of these layers might give

To summarize, the data on the Vegamian Formation an interesting palaeogeographical picture. Discussion of the new data 165

The Alba Formation generally lacks the typical red Distribution and the underlying and overlying forma-

colour in the Palentine facies area. In the Cardano- tions. — The Alba Formation is a well-known marker

Triollo area, for example, the formation consists of a bed throughout the Cantabrian mountain chain. It is light-grey, cream and rose spotted, styliolithic lime- not deposited in the central and eastern part of the stone Veen, in the northern Palentine facies the Liebana and (van 1965). Only part area, e.g. some places of the Gildar-Monto area is the red colour present. In near the Leon line and the Cardano line. In the in the Asturo-Leonese facies northern Cardano-Triollo the Alba Formation one area area a grey area in development of the Alba Formation is known, i.e. wedges out locally. It is also absent in some localities in

area Asturo-Leonese the Riosol (section RI, Fig. 17.) the facies area, but whether this is due

The thickness of the Alba Formation varies from about to tectonic, sedimentologic, or erosional events is not

5 to 40 m, 20 to 30 m being the most common. clear. The Alba Formation is underlain by the Vegamian — Fossil content and age. Commonly found in the Formation or by the Ermita Formation in those parts

Alba Formation are cephalopods, small solitary corals, of the Asturo-Leonese facies area where the Vegamian and crinoid ossicles. Formation is absent. In that area it is overlain by the Radiolarians from in the southern the red cherts and shales were Escapa Formation, with an exception mentioned by Delepine (1937), Kanis (1956) and Albasyncline, where it is followed by a detricalforma-

Koopmans (1962). Conodonts are abundant. tion of shales and sandstone with conglomerate inter- The conclusion be from calations C. G. der Meer general to drawn cephalopod (Mr. van Mohr, pers. comm.). Palentine facies studies is that the base of the Alba Formation in the The overlying formationin the area is facies lies in middle the detrital Cervera Formation Asturo-Leonese area the Pericyclus- (van Ginkel, 1965,

Stufe (cu lip/y). The conodont faunas from the basal p. 198). Only in the Triollo area and near Santibanez part of the Alba Formation in that area invariably de Resoba is the Alba Formation overlain by the Esca- the anchoralis which correlated Formation. belong to Zone, was pa with the has cu lip/y by Voges (1959). The cu II(3/y

generally been correlated with the lower Visean, but Depositional environment. — The Alba Formation is

and & Mauvier formation which Boger (1962) Conil, Lys (1964) clearly a transgressive spread very evidence produced that this zone extends down into rapidly over the flat Asturo-Leonese facies area. A very the Tournaisian Enclosure environment be forsuch upper (see 5). deep-water cannot envisaged a

The base ofthe Alba Formation in the Palentinefacies deposit. Koopmans (1962, p. 152) reported breccias

area is of Visean (cu Ilia — cu the Alba Formation which indeed to indi- upper age Illy; see from seem Ch. 158. shallow-water environment. The radiolarian Ill, p. cate a In the Barruelo-Muda Alba is contradict area, the Formation content of the shales and cherts does not a

nodular Villa- shallow-water environment developed as a grey, limestone, e.g. near (Shackleton Campbell, bellaco (Villabellaco Formation of Wagner & 1954, pp. D17, D18). The development of a bottom

Wagner-Gentis, 1963). The base of this formation is fauna was not inhibited by anoxic conditions, but was of lower Visean of here age, according to Wagner & perhaps limited by fine mud. The rate deposition

Wagner-Gentis (1963); Kullmann (1961), however, was very slow. did not find fossils older than Visean in this To summarize the data the Alba this upper area. on Formation, of in The highest part the Alba Formation is dated as unit is a thin, diachronous formation deposited a Visean the basis of in the lower upper (cu Illy) on cephalopods. shallow, transgressive sea; Visean, or The from it beds, transitional the Alba Formation to the possibly already in the uppermost Tournaisian,

Escapa Formation, have in some places yielded Na- spread rapidly and widely over the Asturo-Leonese murian A (Ej and Eg) goniatites described by Kull- area. and Gonodonts Palentine facies mann (1962) Wagner-Gentis (1963). In the upper Visean it extended to the have not sufficient information define the in the The Alba Formation is supplied to area except centre. Visean-Namurian boundary with certainty. underlainby the Ermita or the Vegamian Formation It is evident from the above-mentionedobservations in the Asturo-Leonese facies and in the Palentine area, that the base of the Alba facies both Formation is diachronous. area only by the Vegamian Formation. In The also be diachronous. disconformable formations upper boundary can slightly areas contacts with these The fullest development of the Alba Formation took are observed.

place in the Asturo-Leonese facies area. 166 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

CHAPTER V

PALAEOGEOGRAPHIC SYNTHESIS OF THE STRATIGRAPHIC DATA

GENERAL REMARKS Gomez (1963), Rupke (1965), de Sitter (1962), de Sitter & Boschma (1966), Sjerp (1967), and van Veen The and sections Almela geologic maps stratigraphic by (1965), cover most of the Cantabrian Mountains. To- et al. (1956), Comte (1959), Garcia-Fuente (1959), gether with our sections and the time-stratigraphic Hernandez & Kindelan correlation chart basis Sampelayo (1950), Julivert of Enclosure 6, they provide a Kanis (1960), (1956), Koopmans (1962), Llopi's Llado on which a be palaeogeographic reconstruction can (I960, 1962), Martinez Alvarez (1962, 1966), Pastor attempted.

Fig. 54. Schematic cross-section stratigraphic from west to east through the region of the study. Palaeogeographic synthesis 167

Fig. 55. Schematic stratigraphic

cross-section from south

to north through the

eastern part of the re- gion ofthe study.

Fig. 56. Schematic stratigraphic cross-section from south to north through the western part of the region of the study. 168 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Fig. 57. Map (1 : 1,000,000) showing the location of the stratigraphic cross-sections in Figs. 54—56.

Fig. 58. Map (1 : 1,000,000) showing the main Devonian palaeogeographic units. Palaeogeographic synthesis 169

Table 9 Literature consulted for the construction of the schematic cross-sections (Figs. 54—56).

columnar section 1. Garcia-Fuente 20 and "cor- columnar section 14. Llado (1959, p. Llopis (1958b, p. 11). tes geologicos"). columnar section 15. Hernandez Sampelayo & Kindelan columnar section 2. Almela etal. (1956, Fig. 2). (1950, pp. 58—61).

columnar section Llado columnar section 16. Comte 3. Llopis (1960, p. 31). (1959, pp. 186—195).

columnar section 4. Mr. H. F. Rouffaer columnar section 17. Comte Sjerp (1967); E.J. (1959, pp. 174—181). (unpublished). columnar section 18. Mr. J. J. K. Poll & Mr. J. L. Liezen-

columnar section 5. this paper (Fig. 17). berg (unpublished).

columnar section 6. columnar section 19. Mr. E. Oele & Mr. Julivert (1960, p. 35). J. van Dillewijn columnar section 7. this paper (Fig. 24). (unpublished).

columnar section this columnar section 20. Martinez Alvarez 8. paper (Fig. 52). (1962, p. 35). columnar sections 9--11. Koopmans (1962, Fig. 10). columnar section 21. Delepine (1928a) and Llopis Llado

columnar section 12. van Veen (1965, Enclosure 2) and (1960, Fig. 6). Mr. Hoeflaken columnar section 22. PI. and this J. A. van (unpublish- Radig (1961, 11; 1964)

ed). paper (Fig. 33).

columnar section 13. Veen Enclosure columnar sections 22 b. van (1965, 2). a, Radig (1962, Fig. 1).

Two important elements attract special attention: De Sitter (1966) showed that the Leon line and the

Pardomino-Pontonline divide the Palaeozoic core of The of hiatus in the Asturo- a. presence a pre-Ermita the CantabrianMountains intofour blocks, each having Leonese facies area, that reaches its maximum its own tectonic and epeirogenetic development. in the central of the mountain chain part (Riosol area).

THE ASTURIAN GEANTICLINE b. The existence of facies two areas, i.e. the Asturo- in Leonese and the Palentine. The Asturian Geanticline did not arise suddenly the Famennian. From the stratigraphic picture of the

Three schematic stratigraphic sections (Figs. 54—56), formations around the geanticline it is clear that this the locations Cambrian. of which are indicated in Fig. 57, show area must already have been a high in the the of these elements. Some of the colum- "Kantabrisch- importance Lotze (1961, p. 485) postulated a

sections to — z.T. Abtra- nar have been shifted along the strike give Iberischer Trog" versus a "Schwellen section straighter lines. No palinspastic correction of gungsgebiet" of Cambrian age. Comte (1959, pp. these sections has been attempted. 147—155) and Radig (1962) showed that from west The the Ordovi- area with the big hiatus, where the pre-Ermita to east towards the Asturian Geanticline,

Devonian is absent, we shall call the Asturian Geanti- cian and the Silurian become thinner and less com- cline and the (Fig. 58). The term "geanticline" is used here plete. In the Rio Bernesga area (Leon) in the broad and Castro sense of "a zone in a mobile belt of the Bufaran anticline (Asturias), the Luarca indicated earth's crust which tends to rise" (Schieferdecker, Formations are missing. Llopis Llado (1965)

and not the of full scale that the and Ferrones Formations 1959), as counterpart a Furada, Nieva,

geosyncline. (Silurian to Lower Devonian) near Oviedo decrease

The Asturian geanticline is fringed in the Asturo- in thickness considerably from west to east. In this Leonese facies direction ratio of the Furada For- area by a zone in which the pre- the sandstone/shale mentioned Ermita hiatus becomes gradually smaller, followed by mation increases. Rupke (1965, pp. 19, 38) Vid a zone in which there is no indentifiable break in the a remarkable difference in thickness for the La sedimentation. In Formation in the Rio Esla where this formation the Palentine facies area a complete, area, Salas though fairly thin, Devonian is present. is much thinner in the northerly situated Las Because the the Van Veen filling of a sedimentary basin is determin- zone than in the autochthone or nappe. ed of the from the Palentine facies area primarily by epeirogenetic movements (1965, p. 53) reported basin and north- and the source area, it is not surprising that these a decrease in thickness, in the northern Forma- isopical zones and facies areas are often bordered by western direction, of the Carazo and Lebanza "fundamental lines" or "tectonic hinge lines", which tions (Silurian and Lower Devonian). have been demonstrated in the Llado 44— by the geologists who map- Higher Devonian, Llopis (1960, pp. MonielloLime- ped the area in detail. The most important of these 46) demonstrated that the Arnao and lines is stones into red sandy the Leon line (de Sitter, 1962; de Sitter & wedge out and pass laterally a

Oviedo. He com- Boschma, 1966). Other fundamental lines are the and shaly formation southeast of

Sabero-Gordon line (Rupke, 1965), the Porma fault pared this deposit with the Old Red Sandstone. It is, the Pardomino line of de Sitter whether this sediment (Rupke, 1965; = however, questionable was

(1966), the Cardano line (van Veen, 1965), and the deposited under continental conditions. But it is quite Geanti- Ponton line (de Sitter, 1966; Sjerp, 1967). probable that the western part of the Asturian 170 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

in cline was an emerged source area that time. The Silurian, and locally the whole Ordovician and part Caldas Formation of Smits (1965), in which terrigi- of the Cambrian. During this interval the Murcia material is fits well into nous important, this picture. Formation was deposited in the Palentine facies area,

It can be inferred from Almela el al. (1953), Garcfa- where it terminated the sedimentation of nodular

Fuente (1959), and Llopis Llado (1960) that this facies limestone (Cardano Formation). At the same time, has the of Pineres with a larger terriginous component an ample higher part the Nocedo Formation, the distribution in Asturias. and of the Forma- Sandstone, part Camporredondo

The sections of Koopmans (1962, p. 169) from the tion were deposited in the Asturo-Leonese facies area, Valsurvio in the southeastern of the Canta- where sedimentation then restricted area part was to a peripher- brian Mountains show thinning and the development al belt around the Asturian Geanticline (Fig. 62). In Devonian south the southern of the sedimentation of stratigraphic gaps in the from to part area the con- north. Besides the pre-Ermita hiatus, Koopmans (1962, tinued only south of the Sabero-Gordon line. hiatus This pp. 173—174) described a pre-upper Valcovero uplift caused by the Bretonic phase (de Sitter, 1962)

in some places cutting down as deep as the Lower did not lead to discontinuities in the Palentine facies

Devonian. Not the Asturian Geanticline proper but area. In the San Julian region of the Barruelo-Muda branch is in northern of this hiatus also in a of it, present the part area a developed, some places reaching

area. We shall call this, in the purely palaeogeo- as far down as the Silurian (Wagner, 1962; Frets, 1965, de Sitter graphic sense, the Santibanez Ridge (see Fig. 58). p. 150; & Boschma, 1966, pp. 230—231). Another, but less conspicious branch of the Asturian

Geanticline is the Pardomino High (Rupke, 1965, pp. 39—41). The question arises whether the Asturian Geanticline

emerged during the Devonian. We observe that:

of a. red sandstones and shales upper Lower Devonian and Middle Devonian southeast of age occur Oviedo (Llopis Llado, 1960); b. material with of coarse erosion is, the exception the

Murcia absent quartzitic Formation, or very scarce

in the Palentine facies area, which abuts on the

Asturian Geanticline.

c. the Huergas Formationin the Asturo-Leonese facies

area contains terriginous detrital material, with an

increasing sandstone/shale ratio towards the Astu- rian in the Rio Geanticline, e.g. Bernesga area and the Rio Esla (Comte, 1959, p. 188) area and (Comte, 1959, pp. 211—212, our observations).

Taking these points into account, we conclude that

the Asturian Geanticline emerged locally from time to time during the Lower, the Middle, and the lower of the but sub- part Upper Devonian, that strong aerial erosion generally did not take place (Fig. 60).

A limited amount of terrigenous, detritalmaterial was supplied by the Asturian Geanticline towards the

Asturo-Leonese facies Eifelian to lower area in upper Givetian times and probably also in lower to middle

Frasnian times (Fig. 61), contributing to the sandy formations in that area.

We assume that the hiatus on the Asturian Geanticline was mainly the result of non-deposition and possibly submarine erosion, (sublevation, Dunbar & Rodgers,

1957, pp. 12—13), both due to little or no subsidence.

The sea floor was probably often above the base level of agradation. Thus, only a thin and incomplete

Devonian deposit could be formed. An epeirogenetic uplift of the Asturian Geanticline causing active, sub-aerial erosion took place in the uppermost Frasnian to lower Famennian, which led to the removal of the inferred thin the 59. the Devonian, Fig. Legend to facies-pattern maps (1 : 1,000,000). Palaeogeographic synthesis 171

60. for the Emsian Eifelian. Fig. Facies-pattern map upper to lower

61. for the middle Frasnian. Fig. Facies-pattern map 172 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

62. for the lower Fig. Facies-pattern map Famennian.

63. for Fig. Facies-pattern map the middle to upper Famennian. Palaeogeographic synthesis 173

The discovery of pebbles of San Pedro ferruginous In the southeast it continued south of the Santibanez

sandstone in and below the A of the basin in a just probably uppermost Ridge. curving southeasterly the in that direction Frasnian Cremenes Limestone (Rupke, 1965, p. 38) direction is suggested by curving

in the Esla confirm of the ofthe Leon line in the The structures nappe may our dating Pisuerga region. this with the Celtiberic fold-belt uplift. We do not, however, accept Rupke's assumption of region link up

that the reached a climax in the 1965; movements upper (de Sitter, 1957, Frets, 1965).

Famennian. Most of the strong erosion caused by the We have seen that the Asturian Geanticline already

uplift ceased at the end ofthe lower Famennian, when existed in the Cambrian. It is possible that on its

the deposition of the Murcia Formation came to an eastern flank there was a land mass on the Ebro block

end and the deposition of the nodular limestone from Cambrian to Devonian times, bounding the

(Vidrieros Formation) in the Palentine facies area was sedimentary trough in the northeast and acting as a

resumed (Fig. 63). source area of varying importance.

The whole Asturian Geanticline must have been More positive information about this supposed land Ermita peneplained before deposition of the post-hiatus, mass can be obtained from the Formation in Ermita the northeast of the Cantabrian Mountains transgressive Formation started in the upper (Sierras

Famennianbecause: Planas), where it lies disconformably on the Quarcita in Ch. Amoricana, as suggested IV, p. 160. In this a. ravinement at the base of the Ermita Formation is it is of region composed predominantly coarse, absent very or very slight; sandy, and partly conglomeratic sediments, in which b. no conglomerate is found in the basal part of the a thin bed with plant remains occurs. According to Ermita Formation, only a few local beds with Hernandez Sampelayo (1928), it can even contain a scattered, small pebbles being known; bed of coal with "tallos vegetales hasta de seis a ocho

c. the Ermita Formation spread fairly rapidly over centimetros de diametro mas de un metro de y longi- the Asturian Geanticline in the Famen- uppermost tud" (reeds as much as six to eight centimetres in dia-

the Riosol to have been nian; only area seems meter and with a length of more than one metre).

reached as late the lower Tournaisian as (Fig. 64). This points to the proximity of a continent along which coastal could swamps develop. The question arises whether the inferred thin, incom- The extension of the Asturo-Leonese Basin in the

Devonian on the Asturian Geanti- plete, pre-Ermita western and southern directions is not well known. The cline belonged to the Asturo-Leonese facies or to the of the Cantabrian Mountains lifted western part was Palentine facies. In view of the situation the during an extent up to such in pre-Stephanian times that the when clastic uppermost Devonian, coarse-grained, western limit of the Devonian and Lower Carbonif- sediments of the Ermita Formation, clearly belonging a erous is boundary fixed by erosion. The great thick- to the Asturo-Leonese the facies, were deposited over of ness the Lower Devonian and, where present, the whole Asturian Geanticline while in the adjacent Middle Devonian formations, as established by Poll

Palentine facies area the nodular limestones and cal- (1963) near Belmonte (Asturias), indicates that the

careous shales of the Vidrieros Formation were being outer limit of the Devonian basin lay beyond that

formed (Fig. 64), we assume that older may any area. The investigation of the Alba syncline and the

Devonian deposits on the Asturian Geanticline also southwesternmost occurrences of the Devonian is still belonged to the Asturo-Leonese facies type. in full swing, and its outcome must be awaited before

The behavior of the Santibanez was Ridge probably reliable conclusions about the southern and western in general analogous to that of the Asturian Geanti- extent of the Asturo-Leonese basin can be reached. cline. The pre-Ermita hiatus here did not reach deeper Remarkable is the occurrence of the Pre-Cambrian than theLower Devonian. The transgressive character (Mora Formation) between Mora and Villablino (de of the part of the Valcovero Formation in the upper Sitter, 1961), not far from the southernmost outcrops northern of the Valsurvio is part area remarkable, of the which does extensive Devonian, not suggest an however. continuation of the Devonian basin in southern

direction. THE LOCATION, NATURE, AND DEVELOPMENT The Asturo-Leonese basin was very shallow. The cross- OF THE DEVONIAN SEDIMENTARY BASINS bedded sandstones, coarse-grained detrital limestones, From distribution of the facies the two patterns we and biostromal limestones point to an epicontinental can distinguish two distinct sedimentary basins: the environment, well aerated, in the photic zone and Asturo-Leonese Basin and the Palentine Basin (Fig. above the wave base. The deposits were formed in 58). the sub-littoral and from zone to a great extent even

the infra-littoral zone (< 50 m deep). Even emersion The Asturo-Leonese Basin has occurred in the Asturo-Leonese Basin. Fossils

The of the Asturo-Leonese facies that soils described Font-Altaba Closas part area are by & (1960, formed the Devonian amended a sedimentary trough during by Koopmans, 1962, pp. 137—138) at the will be called the Asturo-Leonese Basin. This basin of the La Formation Portilla de Luna top Vid near Asturian Geanticline in fringed the the west and south. and by Koopmans (1962, p. 137) at the top of the 174 H. r Adrichem Conodonts Cantabrian A. van Boogaert: of the Mountains

64. for the Famennian. Fig. Facies-pattern map uppermost

65. the Tournaisian. Fig. Facies-pattern map or upper Palaeogeographic synthesis 175

in Valsurvio Desicca- did Palentine Compuerto Formation the area. Famennian, the Basin become a true tion cracks are reported from the Santa Lucia Forma- embayment (Figs. 62 and 63).

Esla The sheltered character of Palentine Basin not tion of the autochthone (Rupke, 1965, p. 20, the was

Fig. 7) and from the Caldas Formation (Smits, 1965). yet clearly expressed in the Silurian, Gedinnian, and

The subsidence was generally small, which meant that Siegenian. Members b and c ofthe Carazo Formation in times of an abundance of reef-building organisms differ considerably from the San Pedro Formation, San but distance bioherms could develop only rarely. Only in the the between the Rio Esla area (the Martin-Ventanilla could bioherms of im- of of the San Pedro Forma- area some easternmost area exposure portance grow against the Santibanez Ridge (Kanis, tion) and the Cardafio-Triolloarea make it impossible

1956, Brouwer, 1964a). to determinethe nature and the place of this change in

Whether calcareous and sedi- which not fundamental. The or sandy argillacious lithology, was, however, the Lebanza Formation differs in litho- ments were deposited depended primarily on also considerably of which is from the lowermost ofthe La Vid supply terrigenous material, mainly con- logy part Formation; trolled by epeirogenetic factors. We do not know to the former, being less argillacious, contains no dolo- what climatic factors role in the mites and consists much of extent played a supply to a greater extent coarse, of terriginous material or reef building. detrital, and fossiliferous limestone. The rich brachiopod fauna does not suggest restricted conditions during the deposition oftheLebanzaForma- The Palentine Basin tion. Binnekamp (1965, pp. 52—53), however, has The sediments of the Palentine facies, with its distinct pointed out the difference between the brachiopod sedimentary character, were deposited in a clearly fauna collected by Comte (1959) in Leon from the limited area that we shall call the Palentine Basin La Vid Formation and the brachiopod fauna from Asturian (Fig. 58). This basin is bounded by the Gean- the Lebanza Formation. It therefore seems permissible ticline in the north and the and the Santi- Palentine Basin west by to distinguish a separate as early as bafiez north- Ridge in the southwest to south. In the from upper Gedinnian to lower Siegenian times. eastern direction a land area is assumed, as explained The sheltered conditions of the Palentine Basin are above (p. 173). emphasized by: in The Palentine Basin may have had a connection a. The limited thickness ofthe deposits, which become the southeast with the main Asturo-Leonese basin, thinner towards the Asturian Geanticline in a because the Barruelo-Muda area appears to have northern well (see Fig. 55) as in western to south- some properties of an area of facies "spill-over". western direction (compare Vidrieros Formation It follows that the Palentine Basin was a sheltered in sections CAL CU and CU III in the almost II, II, area, completely surrounded by positive areas. Gildar-Monto area). The nature of the sediments, being predominantly shales, calcilutites, and nodular limestones, points to b. The occurrence of black shales and bituminous a quiet-water depositional environment. The sea floor limestones in the Gustalapiedra Formation, which below Since was the wave base. the wave action in points to anoxic conditions. The shales of the the sheltered basin would have been Abadi'a Formation darker colour in the not very strong, show a the base wave must have been high and consequently centre of the Palentine Basin than at its southern of great water depth was not essential; a depth more edge. than 50 m does not seem for the necessary deposition The reefs of the Ventanilla lie the c. area appear to on of these quiet-water sediments. The fact that in the southern side of the Santibanez Ridge.

Palentine Basin no erosion resulted from the uplift of d. The occurrence of impoverished coral and cepha- the central Cantabrian area could point, however, to faunas in the Palentine Basin of the basin. it is lopod (Kullmann, a great depth Nevertheless, more 1965, 58—60). probable that this basin did not join the uplift of the pp. adjacent areas and remained a relatively shallow, sub- e. The occurrence of plant remains in the nodular siding trough bordered by marginal faults, in which the limestones in the Palentine Basin. Wagner & Murcia Formation could accumulate. A indicated remains small, deep Wagner-Gentis (1963, p. 151) basin, bordered by positive areas, would show much of drifted land plants in beds of Middle Devonian and which found in the Barruelo-Muda Veen slumping grading features, are rarely age area; van (1965, in the Palentine Basin. p. 58) found a few thin stems of wood in the Cardano Formation The sheltered Palentine Basin was possibly surround- in the Cardano area. ed of shallow in which by a zone very water ocean waves and tidal action were damped out and in which A view ofthe Palentine facies area as a sheltered basin the sediment dead end. is in conflict with the of rich currents transporting came to a not presence a pelagic

We visualize the Santibanez Ridge as having been a microfauna. Conodonts are known to be pronouncedly similar in times of of the The number of conodonts zone. Only general uplift facies-breaking. per sample

Asturian Geanticline and the Santibanez Ridge, as is inversely proportional to the rate of sedimentation occurred from the Frasnian the middle the uppermost to (Lindstrom, 1964, p. 68). Especially slowly- 176 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

66. for the lower Viséan. Fig. Facies-pattern map

67. for the Viséan. Fig. Facies-pattern map upper Palaeogeographic synthesis 177

deposited nodular limestones are rich in conodonts. Next, a slight regression brought the deposition of the

BouCek 57, has that black shale to an end in the Asturo- (1964, pp. 155) reported pelagic Vegamian locally restricted tentaculites (Dacryoconarida) are not to Leonese facies area. The deposition of the Vegamian

one particular facies because of their pelagic mode Formation in the Palentine Basin was not affected.

of life. The scarcity or absence of pelagic elements in Slight erosion of the newly-deposited Vegamian be the Formation in the Asturo-Leonese facies might explained by took place in some regions the Asturo-

small chance of fossilization of their delicate remains Leonese facies area, leaving an irregular distributional in the turbulent environment of that facies and of this with thickness. by pattern formation, varying their "dilution" in the thick sediment With the beyond recovery a new rapid general transgression over very flat surface of facies sequence. the Asturo-Leonese area, the red nodular limestone complex of the Alba Formation This was deposited (Fig. 66). transgression started in THE LOWER CARBONIFEROUS DEVELOPMENT the lower Visean in the or even already uppermost OF THE SEDIMENTARY BASINS Tournaisian (cu Ilp/y). In the Palentine Basin this

As we have seen in the discussion of the Ermita and transgression had no effect. The deposition of black

Vidrieros the of the shales continued into the Visean the Formations, pattern uppermost probably upper (at least in Gildar-Monto Kull- Devonian sedimentation (Fig. 64) continued in the the area: Budinger &

In the Visean, after a lowermost Tournaisian. A general regression soon fol- mann, 1964, p. 421). upper lowed. Only in the centre of the Palentine Basin did short break in the sedimentation, the Alba Formation in the Palentine Basin with the the sedimentation continue with the deposition of the was deposited too,

black shale of the Vegamian Formation. exception of the central and eastern part, e.g. the in which Liebana The San in Later in the Tournaisian, the sea the Vega- area (Fig. 67). Julian region the

mian Formation was deposited spread over the whole Barruelo-Muda area was probably still an an island

Palentine Basin, reaching the Asturo-Leonese facies during this period.

area in the Tournaisian. There, the black shale The character of the Asturian Geanticline as a upper high

complex developed in northeastern and central Astu- disappeared in the Visean. The former positive area northern and in of the Rio formed of the basin in which the Forma- rias, Leon, part Bernesga, part Escapa

Rio Torio, and Rio Curueno areas (Fig. 65). This tion was deposited in the Namurian. Flysch-type Tournaisian sediments in the Palentine Basin in picture continued till the uppermost to were deposited

lowermost Visean. that time.

CHAPTER VI

CONODONT SYSTEMATICS

this In chapter the most important species charac- being photographed, the conodonts were coated

terizing the conodont zones are illustrated and briefly lightly with a sublimate of ammonium chloride.

discussed. For the terminology, we refer to Hass (1962). All the conodont faunas from the samples mentioned We followed the recommendation of in this be found in the have, however, paper are to Rijksmuseum van

Scott el al. (1962) concerning the orientation of the Geologie en Mineralogie, Hooglandse Kerkgracht 17, plate-like conodonts. Leiden, The Netherlands. The illustrated specimens

The conodonts were recovered from limestone samples have been registered separately.

(occasionally dolomitic), weighing about 1 kg, with Genus ANCYRODELLA Ulrich & Bassler, 1926 the exception of the Lower Devonian samples, which Ancyrodella curvata (Branson & Mehl, 1934) were generally twice as heavy. Monochloro-acetic Plate 1, Fig. 1 acid was used to digest the carbonate. The separation of the heavy fraction was carried out in bromo- pure 1934a — Branson & Ancyrognathus curvata n.sp. Mehl, p. 241, form. Voluminous heavy residues were magnetically PI. 19, Figs. 6, 11. separated after Dow (1960). Often found in associa-

1956 & — Ancyrodella curvata (Branson Mehl) Bischoff, p. tion with the conodonts and dermal were teeth, scales, 118, Figs. 9—11.

of For further & plates fish, phosphatic brachiopod shells; and, synonymy, see Glenister Klapper (1966,

ostracods and of tenta- occasionally, posterior tips p. 798). culites.

This has its side The enlargement of the photographs (25 or 40 times) Remarks. Ancyrodella on outer a poste- is indicated in the explanations of the Plates. Before riorly-directed lobe with secondary carina and second- 178 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

keel. This is abundant in Frasnian Gnathodus texanus authors. The latter ary species our by European was faunas. Relations with other Frasnian ancyrodellids not found in the Cantabrian Visean. found There transitional forms Gnathodus can be in Ziegler (1962a, Fig. 2). are of antetexanus tc Gnathodus typicus and slender forms of Gnathodus semi- Ancyrodella rotundiloba (Bryant, 1921) glaber. Ancyrodella rotundiloba rotundiloba (Bryant, 1921) Gnathodus bilineatus (Roundy, 1926) Plate 1, Fig. 3 Plate 2, Figs. 4—6 PI. 1921 rotundilobus — Polygnathus n.sp. Bryant, p. 26, 12, 1926 Polygnathus bilineata Roundy in Roundy et al., Figs. 1—6, Text-fig. 7. n.sp. PI. 10. p. 13, 3, Fig. 1956 Ancyrodella rotuntiloba (Bryant) — Hass, PI. 4, Fig. 21. 1953 Gnathodus bilineatus (Roundy) Hass, p. 78, PI. 14, 1966 Ancyrodella rotundiloba rotundiloba (Bryant) — Glenister Figs. 25—29. & PI. 9—13. Klapper, p. 799, 85, Figs.

1965 Gnathodus — bilineatus (Roundy) Budinger, pp. 53—56,

PI. 2, Figs. 1—8; Text-figs. 15, 16. Remarks. The illustrated specimen is fairly juvenile. 1965 Gnathodus delicatus Branson & with Mehl Budinger, pp. The adult forms are covered many regularly- 56—57, PI. 2, Fig. 12. [non Figs. 9—ll, 13 = G. delicatus placed, rounded nodes. Relations with other Frasnian s.l.]. ancyrodellids can be found in Ziegler (1962 a, Fig. 2). For further synonymy, see Budinger (1965).

Genus ANCYROGNATHUS Branson & Mehl, 1934 Remarks. The long, sharp, transversely-ridged parapet Ancyrognathus asymmetrica (Ulrich & Bassler, 1926) side which the on the inner of the cup, runs alongside Plate 1, Fig. 2 carina, reaches the posterior tip of the conodont. This

— Ulrich & the from Gnathodus delicatus 1926 Palmatolepis asymmetrica n.sp. Bassler, p. distinguishes species 5.1., PI. 18. which has shorter and nodular 50, 7, Fig. a more parapet or

1958 Ancyrognathus asymmetrica (Ulrich & Bassler) — Ziegler, only a row of nodes on the inner side of the cup. The PI. 11. of this does reach the pp. 10, Figs. 10, parapet species not posterior tip. For further Glenister & synonymy, see Klapper (1966, Gnathodus commutatus (Branson & Mehl, 1941) p. 801).

1941 Branson & Spathognathodus commutatus n.sp. Mehl, p. Remarks. Some ofthe broader specimens in our material 98, PI. 19, Figs. I—4. tend towards Ancyrognathus calvini (Miller & Young- 1957 Gnathodus commutatus commutatus (Branson & Mehl) — quist, 1947) but the keel angle, which does not surpass PI. 4, 15. Bischoff, p. 23, Figs. 2—6, 90°, and the posterior lobes, which are not very rounded, distinguish them from that species. Relations Gnathodus commutatus cruciformis Clarke, 1960

with other Frasnian ancyrognathids can be found in Plate 2, Fig. 8

Ziegler (1962a, Fig. 5). & 1958 Gnathodus commutatus nodosus Bischoff—Lys Serre, pp.

Ancyrognathus triangularis Youngquist, 1945 891—892, PI. 9, Fig. 4, [ram Fig. 3 = G. commutatus Plate 1, Fig. 4 commutatus].

1960 Gnathodus 25, PI. cruciformis n.sp. Clarke, p. 4,

1945 Ancyrognathus triangularis — Youngquist, 356, n.sp. p. Figs. 10—12. PI. 54, Fig. 7. 1965 Gnathodus commutatus cruciformis Clarke—Budinger, p. For Glenister & synonymy, see Klapper (1966, p. 802). 49, PI. 4, Figs. 10, 11; Text-fig. 12.

Remarks. The available lie well within the specimens each side of the Remarks. One transverse ridge on cup Relations variability range given by Ziegler (1958). is characteristic for this species. Transitional forms to with other Frasnian ancyrognathids can be found in Gnathodus commutatus nodosus are common. Ziegler (1962a, Fig. 5).

Gnathodus commutatus nodosus Bischoff, 1957 Genus GNATHODUS Pander, 1856 Plate 2, Fig. 7

Gnathodus antetexanus Rexroad & Scott, 1964 Gnathodus commutatus nodosus Plate 2, Fig. 3 1957 n.subsp. Bischoff, p. 23, PI. 4, Figs. 12, 13.

1964 Gnathodus antetexanus — Rexroad & For n.sp. Scott, pp. synonymy, see Budinger (1965). 28—29, PI. 2, Figs. 7—10.

1965 PI. bears node Gnathodus texanus Roundy—Budinger, pp. 60—62, 3, Remarks. The cup normally a prominent on

= = of G.Figs. 7—B only [non Fig. 2 G. typicus, nee Fig. 3 both sides of the carina, but sometimes only one

= G. cf. G. Text- semiglaber, nec 10 cuneiformis]; of the nodes a Figs. 9, these nodes is present. By elongation to fig. 17 [form A only]. ridge, Gnathodus commutatus cruciformis developed from For further see Rexroad & Scott synonymy, (1964). this species.

This reduced than Gnathodus Mehl & Thomas, 1947 Remarks. species has a less cup cuneiformis 10 Gnathodus texanus. It has been repeatedly described as Plate 2, Figs. 9, Conodont systematics 179

Gnathodus — Mehl & 1947 cuneiformis n.sp. Thomas, p. 10, the lower part of the anchoralis Zone and the other in

PI. 2. 1, Fig. the bilineatus-delicatus s.l. Zone). For the present, we Rexroad & Scott For synonymy, see (1964, p. 29). shall the whole delicatus s.l. classify group as Gnathodus

Gnathodus girtyi Hass, 1953 Remarks. This species has along both sides of the carina Plate 16 to 2, Fig. a low row of nodes, only anteriorly fused form a which is towards the carina. The parapet concave 1953 Gnathodus — PI. girtyi n.sp. Hass, p. 80, 14, Figs. 22—24. on the inner side extends slightly further for- parapet 1957 Gnathodus PI. girtyi Hass—Bischoff, pp. 24—25, 4, the other side. The is ward than the one on cup Figs. 16—23. slender and rather symmetric. For comparisons with

1960 Gnathodus clavatus — PI. 4—9. n.sp. Clarke, p. 25, 4, Figs Gnathodus girtyi, see there. 1965 Gnathodus Hass 57 girtyi —Budinger, pp. —58, PI. 2,

Gnathodus cf. G. cuneiformis Mehl & Thomas, 1947 Figs. 14—16 [non PI. 3, Fig. 11 = G. cf. G. girtyi, nee Fig.

12 = G. cf. G. nee 13 = G. Plate 2, Figs. 11, 12 cuneiformis, Fig. typicus?]; Text-fig. 17 [form D only].

Gnathodus PL 1965 girtyi Hass—Budinger, pp. 57—58, 3,

cf. G. PL 14— = G. Fig. 12 \non Fig. 11 girtyi, nee 2, Figs. Remarks. Gnathodus girtyi has a high parapet on either

16 = G. girtyi]. side of the carina. The has broad cup a low, bulge at

1965 Gnathodus PL texanus Roundy—Budinger, pp. 60—62, its outer side; Gnathodus cuneiformis has lower, incom-

10 2 = G. = G. Figs. 9, [non Fig. typicus, Nec Fig. 3 which towards 3, plete parapets are anteriorly concave

semiglaber, nee Figs. 7, 8 = G. antetexanus]; Text-fig. 17 the carina and is more symmetric than the former B only]. [form species.

1957 Gnathodids with of nodes Gnathodus kockeli Bischoff, Remarks. two rows or para- Plate 18 of size the carina here classified 2, Figs. 17, pets unequal along are

as Gnathodus cf. G. cuneiformis. Among others, a number

1957 Gnathodus kockeli — PL n.sp. Bischoff, p. 25, 3, Figs. of had be in this fairly juvenile specimens to placed 27—32.

species. 1959 kockeli PI. Gnathodus Bischoff—Voges, pp. 281—282,

33, Figs. 26, 27. Gnathodus delicatus Branson & Mehl, 1938 s.l. 1964 Gnathodus kockeli BischofF in et PI. Plate 2, Figs. 13—15 —Higgins Higgins al., 5, Fig. 27.

1938b Gnathodus delicatus — Branson & n.sp. Mehl, p. 145, PI. 34, Figs. 25—27. Remarks. For description and comparison with Gnatho- Gnatho- 1964 Gnathodus delicatus Branson & Mehl Higgins in dus commutatus homopunctatus, see Voges (1959).

PI. 24. Higgins et al., p. 226, 5, Fig. dus kockeli is rarely represented in the lower Tournai-

1965 Gnathodus delicatus Branson & Mehl sian conodont faunas of the Cantabrian Mountains. Budinger, pp.

56—57, PI. 2, Figs. 9—ll, 13 [non Fig. 12 = G. bilineatus]. Gnathodus For further Rexroad & Scott punctatus (Cooper, 1939) synonymy, see (1964, pp. Plate 19 29—30). 2, Fig.

1939 — PI. Dryphenotus punctatus n.sp. Cooper, p. 386, 41, Remarks. Gnathodus delicatus has The great variability. Figs. 42, 43, PI. 42, Figs. 10, 11.

older of this have a fairly representatives species 1959 Gnathodus — PI. punctatus (Cooper) Hass, p. 395, 47,

slender 2, whereas the ones cup (PL Fig. 13), younger Figs. 11—18.

tendto have a broad 2, 14, Gnathodus For 58 cup (PL Figs, 15). synonymy, see Budinger (1965, pp. —59). perplexus Branson 145, PI. 34, & Mehl, 1938, p. Fig. included in Rexroad & Scott's for 24, synonymy Remarks. Characteristic for this species is the short

Gnathodus is also a broad variant. For criteria inner side of the delicatus, parapet on the cup curving away this broad from Gnathodus to distinguish type bilineatus, from the carina posteriorly.

see there. Gnathodus semiglaber Bischoff, 1957 Collinson, Scott & Rexroad (1962) found Gnathodus Plate 2, Figs. 20, 23 delicatus in the Mississippi Valley restricted to late Kinderhook and times. & early Valmeyer Budinger 1957 Gnathodus bilineatus semiglaber n.subsp. Bischoff, p. Kullmann and this (1964) Budinger (1965) reported 22, PI. 3, Figs. I—lo, 12—14. Its species as high as the Visean cu Illy. upper 1959 Gnathodus — PI. semiglaber (Bischoff) Voges, p. 284, 33, occurrence in our material is the highest Upper Figs. 38, 39.

bilineatus-delicatus s.l. Zone of approximately cu Ilia— 1965 Gnathodus Bischoff 59 semiglaber —Budinger, pp. —60, cu 111(3 age. PI. 3, Figs. 1, 4—6, Text-fig. 17 [forms D, E]. into It might prove possible to split this species two 1965 Gnathodus texanus Roundy—Budinger, pp. 60—62, PI. subspecies, if enough material from complete sections 3, Fig. 3 \non Fig. 2 = G. typicus, nee Figs. 7, 8 = G.

were studied. An indication for this is the occurrence of = G. cf. antetexanus,Nec Figs. 9, 10 cuneiformis].

maxima in its distribution in section OL in For further Rexroad & Scott two (one synonymy, see (1964, p. 30). 180 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Gnathodus has low The median of nodes Remarks. semiglaber a asymmetrical a compressed appearance. row

whose broad outer side is ornamented. extends further with a ridge of 2 cup scantily (carina) backward, often bears three 3 fused the denticle the Its posterior tip is broad and rows to denticles, posterior being

ofsmall nodes. This species is very variable. There are largest and sometimes cusp-like. The lateral rows of Lateral view: transitions to Gnathodus antetexanus, Gnathodus cuneiformis, nodes consist of 8 to 11 discrete nodes.

and Gnathodus typicus. The posterior edge makes a right angle with the oral

side of the conodont or slants backwards slightly. Gnathodus typicus Cooper, 1939 Aboral view: The expanded pulp cavity is widest Plate 2, Fig. 21 posteriorly, tapering toward the anterior part. A

postero-lateral flange is weakly but distinctly develop-

— PI. 1939 Gnathodus typicus n.sp. Cooper, p. 388, 42, Figs. ed at the outer side. At the innerside a faintanteriorly- 77, 78. directed spur is indicated. 60 PI. 1965 Gnathodus texanus Roundy—Budinger, pp. —62, Distribution. Spain. The part of the La Vid 8 upper 3, Fig. 2 [non Fig. 3 = G. semiglaber, Nec Figs. 7, = G. Lucia Formation and the basal part of the Santa 10 = G. cf. G. antetexanus, Nec Figs. 9, cuneiformus]. Formation in the Rio Esla area. One specimen was PI. 1965 [?] Gnathodus girtyi Hass—Budinger, pp. 57—58, found in the upper part of the Santa Lucia Formation. 11 = G. cf. G. 12 = G. 3, Fig. 13 [turn Fig. girtyi, nec Fig. Belgium. Upper Emsian (Rhenan sence) to Lower cf. G. cuneiformis, nee PI. 2, Figs. 14—16 = G. girtyi]. Couvinian Prof. Dr. O. H. For further Rexroad & Scott (pers. comm. Walliser, synonymy, see (1964, pp.

31—32). Gottingen, Germany).

Icriodus eslaensis n.sp. Remarks. Gnathodus typicus has a short, high on parapet Plate 1, Figs. 9—12 the inner side of the slender cup; the outer side bears

Transitions Gnathodus delicatus 1938 Icriodus Branson & — a few small nodes. to cymbiformis Mehl? Stauffer, p.

and Gnathodus semiglaber exist. 430, PI. 52, Figs. 11, 13.

1940 Icriodus Branson & Mehl cymbiformis Stauffer, p. Gnathodus A Collinson, Scott & Rexroad, 1962 sp. I.425, PI. 60, Figs. 56—58, 68, 37 [?] [nor, Fig. 51 = Plate 2, Fig. 22 cymbiformis].

1940 Icriodus Branson & Mehl expansus Stauffer, p. 425, 1961 Gnathodus cf. G. commutatus (Branson & Mehl) Scott PI. 60, Figs. 70, 71, 59 [?], 64 [?] [mm Figs. 40, 47—48, & PI. Figs. 23—27. Collinson, pp. 123—124, 1, 60—63 = I. expansus]. 1962 Gnathodus A — Collinson, Scott & Rexroad, 8, n.sp., p. 1965 Icriodus cf. I. obliquimarginatus Bischoff & Ziegler —- Chart 3. Adrichem Table Table 9. van Boogaert, p. 166, 2; p. 175,

1964 Gnathodus A in el sp. Higgins Higgins al., p. 227,

PI. 5, Fig. 28. where this Derivatio nominis. After the Rio Esla area,

species was first recognized. think that the smooth from Remarks. We gnathodids Holotype. Specimen AB 9, illustrated in PI. 1, Fig. 9. the lower Tournaisian TR 21 Gnathodus sample are sp. Locus typicus. La Velilla de Valdore. is sub-circular but A. In our material the not The basal of the La Portilla cup Stratum typicum. part

its axis a small with oval, long usually making angle Formation in the Rio Esla area. of conodont. the longitudinal direction the Material. 225 specimens.

Diagnosis. A small slender species of Icriodus with the Genus ICRIODUS Branson & Mehl, 1938 median row of nodes (carina) extended posteriorly Icriodus corniger Wittekindt, 1966 the short lateral of nodes. The extended past rows part Plate 6 1, Figs. 5, consists of 4 to 5 fused denticles, which increase in

height posteriorly.

1966 — Icriodus corniger n.sp. Wittekindt, p. 629, PI. 1, Description. Oral view: The conodont is relatively Figs. 9—12. small, uncurved and slender, with sub-parallel sides.

The median row of nodes consists of 10 to 13 nodes. Remarks. The sharp postero-lateral flange of the ex- the In the posterior part, where it extends beyond panded pulp cavity is characteristic. In some specimens of it consists of fused lateral row nodes, 4 to 5 denticles, this flange bears a low ridge consisting of a few fused which increase in height posteriorly. The last denticle nodes. of is the largest and often cusp-like. The lateral rows do reach the anterior Icriodus cf. I. corniger Wittekindt, 1966 nodes not tip. They usually consists of 4 5 descrete but their number Plate 1, Figs. 7, 8 to nodes, from 3 to 7. The lateral nodes tend alter- can vary to

Material. 4 complete and 5 indentifiable damaged nate slightly with respect to those of the median row.

specimens. Some of them, on one or both sides, may be suppressed,

Description. Oral view: Fairly slender, slightly curved giving an irregular appearance to some specimens. widest the middle. The nodes the Lateral view: The conodont is arched. Its conodont, at on not posterior smaller and is inclined backward. Aboral view: The posterior part are more closely set, giving edge slightly Conodont systematics 181

is In 1938a Icriodus latericrescens — Branson & pulp cavity moderately expanded posteriorly. n.sp. Mehl, pp. 164—165, PI. 26, Figs. 30—37. some specimens a faint antero-lateral spur is indicated.

The 1938 Icriodus latericrescens Branson & Mehl — expanded pulp cavity tapers rapidly anteriorly. Stauffer, p.

PI. 34. Distribution. Spain. The species occurs in the varca 430, 52, Figs. 30,

Zone s.l. (middle to Givetian) in the Canta- upper 1947 Icriodus latericrescens Branson & Mehl — Youngquist,

brian Mountains. U.S.A. Stauffer PI. (1938, 1940) p. 102, 25, Fig. 25. this from the shale in reported species Olentangy 1952 Icriodus latericrescens Branson & Mehl [?] Graves, p. Ohio and Ontario and in clays above the Cedar 612, PI. 81, Fig. 9, [non Figs. 14—16 = I. latericrescens limestone Valley in Minnesota. beckmanni ?].

Remarks. Icriodus angustus Stewart & Sweet, 1965 differs 1952 Icriodus curvatus Branson & Mehl — Graves, PI. 81, from this in that its median of nodes does species row Figs. 10, 11. not reach the anterior tip and its oral ornamentation

1956 Icriodus latericrescens Branson & Mehl — Stewart & forms transverse ridges. PI. 7. Sweet, p. 268, 33, Figs. 2, 6, Icriodus obliquimarginatus seems to be related to the new

1956 Icriodus latericrescens latericrescens Branson & Mehl — species. The former is, however, considerably more PI. 14—17. Ziegler, p. 100, 6, Figs. slender and has a more irregular oral surface in which 1958 Icriodus latericrescens latericrescens Branson & Mehl, the nodes of the lateral rows are sometimes connected Bischoff & PL 8. Sannemann, p. 95, 12, Fig. with the nodes of the median row by small ridges.

1962 Icriodus latericrescens latericrescens Branson & Mehl —

Furthermore, it has a more widely expanded pulp PI. 16. Jentzsch, p. 967, 1, Fig. cavity with an antero-lateral spur. Juvenile specimens 1964 Icriodus latericrescens Branson & Mehl— Orr, pp. 9—10, of Icriodus eslaensis n.sp. conform to the description of PI. 2, Figs. 8—10. Icriodus We with Bischoff & cymbiformis. agree Ziegler 1966 Icriodus latericrescens latericrescens Branson & Mehl — (1957, 62), who that this small icriodid p. suppose Clark & PI. 14. Ethington, p. 679, 83, Figs. 6, 7, 10, 11, represents juvenile forms of various species of Icriodus. 1966 Icriodus latericrescens Branson & Mehl Wittekindt,

629—630, PI. 1, Figs. 6—B. Icriodus latericrescens Branson & Mehl, 1938 pp.

Remarks. This species has been reported from the This subspecies of Icriodus latericrescens has one spur, Lower and Middle Devonian. Its main distribution which points postero-laterally. and abundance in the Lower where occur Devonian, Description. See Ziegler (1956: ".sensu Ziegler") and '" three " " " Ziegler (1956) distinguished subspecies. Philip Wittekindt (1966: sensu Branson & Mehl"). (1965) and Wittekindt the (1966) propose placing The Lower Devonian forms of this subspecies are Lower Devonian forms in a separate, homeomorphous mostly somewhat smaller than representatives of species. The fact that in Icriodus latericrescens Europe Icriodus latericrescens bilatericrescens. There are transi- has been found in the Lower Devonian and in the tional forms between these subspecies.

Givetian (part of the varca but in the Zone), never Characteristic for the Givetian forms (PL 1, Figs. 16,

Eifelian, seems to this view. In the United support 17) are the poorly-developed median row ofnodes and States, however, this species is reported from the often covered with double of the strong spur, a row lower of the part Middle Devonian as well (Branson small nodes. & Mehl, 1938).

The older Lower Devonian forms of Icriodus latericre- Icriodus latericrescens bilatericrescens Ziegler, 1956

scens are generally broad and massive, i.e. ”Icriodus Plate 1, Figs. 18—20

latericrescens (sensu Ziegler, 1956)". The species shows

1956 Icriodus latericrescens bilatericrescens n.subsp. — Ziegler, a tendency to become more slender and to have more 101—102, PI. 6, Figs. 6—13. set of nodes toward the pp. closely rows upper part of the 1958 Icriodus latericrescens bilatericrescens & Lower Devonian (PI. 1, Figs. 13—15, 18, 20). Where Ziegler—Bischoff Sannemann, 96, PI. 12, Fig. 5. this continues p. tendency we may see the slender Middle 1958 Icriodus lalericrescens cf. latericrescens Ziegler—Bischoff & Devonian forms, i.e. ”lcriodus latericrescens (sensu Bran- PL 11. Sannemann, pp. 95—96, 12, Figs. 10, son & Mehl)" as a normal evolution of the same 1962 Icriodus latericrescens bilatericrescens Ziegler—Jentzsch, p. species. There seems to be only an arbitrary boundary 966, PI. 1, Figs. 12, 14, 24. between the "isensu Ziegler" and the "jsensu Branson & Icriodus latericrescens cf. Mehl" forms. 1962 bilatericrescens Ziegler—Jentzsch,

PI. 15. p. 966, 1, Figs. 10, 13, It has not yet been explained why Icriodus latericre- 1962 Icriodus woschmidti Ziegler—Jentzsch, 967, PI. 1, scens is so rarely found in the Middle Devonian in p. Figs. 17, 20—22. Europe. Perhaps the species became more facies- 1965 Icriodus bilatericrescens Ziegler—Philip, PI. 9, dependent in that epoch. Icriodus is known to have its p. 103, Figs. 30—32. own facies preferences, since it is sometimes the only Icriodus latericrescens in faunas. 1966 bilatericrescens Ziegler—Clark & plate-like genus present conodont PI. 15. Ethington, p. 679, 83, Figs. 12, 13, Icriodus latericrescens latericrescens Branson 1938 & — Mehl, 1966 Icriodus pesavis Bischoff & Sannemann Clark &

Plate 13—17 PI. 5. 1, Figs. Ethington, p. 680, 83, Fig. 182 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

— This of Icriodus latericrescens has two 1934a Palmatolepis distorta Branson & Mehl, 237, subspecies spurs, n.sp. p. PI. 14. one 18, Figs. 13, pointing postero-laterally, the other in the oppo- For see Glenistcr & (1966, site direction. synonymy, Klapper p. 809).

Description. See Ziegler (1956). Remarks. Palmatolepis distorta is strongly sigmoidal. It Remarks. The variation of the oral ornamentation, has the side of the close which from of nodes a ridge on outer platform, to ranges widely-set transverse rows the blade. For comparisons with Palmatolepis glabra to closely-set rows, resembles that of Icriodus latericre- peclinata, see Ziegler (1962b). scens latericrescens. A few of the specimens with a well-

developed postero-lateral bear two or even spur Palmatolepis gigas Miller & Youngquist, 1947 three rows of fused, small nodes on the (PI. 1, spur Plate 2, Fig. 27 Fig. 19). Juvenile forms generally a poorlydeveloped 1947 Palmatolepis gigas Miller & Youngquist, antero-lateral spur and may resemble Icriodus la- n.sp. p. PI. 1. tericrescens latericrescens. Icriodus latericrescens bilatericre- 512, 75, Fig.

1956 rhenana PI. scens is not known from the Middle Devonian. Palmatolepis n.sp. Bischoff, p. 129, 8, Figs. 26—28, PI. 10, Fig. 7. Icriodus woschmidti Ziegler, 1960 1963 Palmatolepis gigas Miller & Youngquist—Klapper & Plate 1, Fig. 21; Plate 2, Fig. 1 406—407. Furnish, pp.

For further Glenister & synonymy, see Klapper (1966, Icriodus PI. 1960 a woschmidti n.sp.— Ziegler, p. 185, 15, p. 810). Figs. 16—18, 20—22.

Icriodus woschmidti PI. 1962 [non] Ziegler—Jentzsch, p. 967, Remarks. The slender patform, posteriorly pointed, the 1, Fig, 17—23 [Figs. 17, 20—22 = I. latericrescens bilateri- long inner lobe, and the high free blade, characterize crescens]. this species. Relations with other Frasnian palmato- 1964 Icriodus woschmidti PI. Ziegler—Walliser, pp. 38—39, lepids can be found in Ziegler (1962a, Fig. 8). 9, Fig. 22; PI. 11, Figs. 14—22.

Palmatolepis glabra Ulrich & Bassler, 1926 Ulrich & 1926 Remarks. Characteristic features are the very widely Palmatolepis glabra glabra Bassler,

separated transverse rows of three fused nodes form- Plate 2, Fig. 25

ing ridges and the cusp-like posterior denticle on the

1926 — Ulrich & Palmatolepis glaber n.sp. Bassler, p. 51, oral surface. A postero-lateral spur and an oppositely PI. 9, Figs. 18—20. directed antero-lateral wide spur are present. The very 1934 a Palmatolepis glabra Ulrich & Bassler Branson & of the transverse rows of nodes spacing distinguishes P. PI. 26 22 = Mehl, p. 233, 18, Fig. [non Figs. 9, glabra this species from Icriodus latericrescens bilatericrescens. pectinata].

1960b Palmatolepis glabra glabra Ulrich & Bassler—Ziegler, Icriodus sp. a PI. 1, 11—13. Plate 2, Fig. 2 p. 7, Figs. For Glenister & further synonymy, see Klapper (1966, p.

811). Remarks. This be species appears to related to Icriodus latericrescens bilatericrescens. Instead of the antero- of the Remarks. This species has no inner lobe. Most lateral and broad are material small. postero-lateral spurs, flanges specimens in our were fairly developed in these directions. The expanded pulp is Palmatolepis glabra elongata Holmes, 1928 cavity exceptionally wide. The oral side of the cono- Plate 2, Fig. 26 dont is fusiform, being widest in the centre. The

lateral rows ofnodes as connected are developed ridges 1928 PI. Palmatolepis elongata n.sp. Holmes, p. 33, 11, with the median row. In the lateral view the oral Fig. 33. surface of the conodont is arched. The of the flanges 1960b PI. Palmatolepis glabra elongata Holmes—Ziegler, p. 8, of the reminis- posterior part expanded pulp cavity are 1, Figs. 10, 14.

cent ofIcriodus which a different further Glenister & corniger, has, however, For synonymy, see Klapper (1966, pp. oral ornamentationand its width greatest posteriorly. 811,814).

Only three of Icriodus a were found in specimens sp.

our material, all in 19 from the Rio Esla area. Remarks. This is slender of sample avery subspecies Palmatolepis We shall therefore wait it to present as a new species glabra with a small, lobe-like outer platform. until more material is available for study. Palmatolepis gracilis Branson & Mehl, 1934

Genus PALMATOLEPIS Ulrich & Bassler, 1926 Palmatolepis gracilis gracilis Branson & Mehl, 1934

Plate 2, Figs. 28, 29 The phylomorphogenetic development of Palmatolepis, Branson & 1934 a Palmatolepis gracilis n.sp. Mehl, p. 238, which genus is ofgreat stratigraphic value in the Upper PI. 18, Fig. 8 [non Fig. 2 = P. glabra elongata Holmes, nee Devonian, can be found in Helms (1963). Fig. 5 (= Nothognathella?)].

distorta Branson & 1934 — Palmatolepis Mehl, 1956 Palmatolepis (Deflectolepis) deflectens n.sp. Muller, p. Plate 2, Fig. 24 32, PI. 11, Figs. 28—39. Conodont systematics 183

PI. transitans & 1959 Palmatolepis deflectens Miiller—Helms, p. 648, 6, 1957 Palmatolepis Miiller—Bischoff Ziegler, p. Fig. 20. 81, PI. 16, Figs. 23—27.

1958 transitans 1962b Palmatolepis deflectens deflectens Miiller—Ziegler, p. 56, Palmatolepis Miiller—Ziegler, p. 66, PI. 1,

PI. 3, Figs. 17—22. Figs. 9, 11 —13; PI. 2, Figs. 1—3, 8.

1962 Palmatolepis gracilis gracilis Branson & Mehl — Mehl & Remarks. transitans developed from Ziegler, pp. 204—205. Palmatolepis Poly-

For further Glenister & Relations with synonymy, see Klapper (1966, pp. gnathus asymmetrica asymmetrica. upper 814—815). Givetian and Frasnian polygnathids and palmatole-

pids can be found in Ziegler (1962a, Fig. 8).

Remarks. This Palmatolepis has an extremely reduced Genus pelekysgnathus Thomas, 1949 platform. Some forms transitional to Palmatolepis Pelekysgnathus serrata, Jentzsch, 1962 minuta occur. Specimen AB 42, illustrated in PI. 2, Plate 2, Figs, 34, 35 Fig. 29 has a bifurcated posterior tip. This seems to be

a pathologic anomaly. 1962 Pelekysgnathus serrata n.sp. —Jentzsch, pp. 970—971, PI. 2, Figs. 7, 8; PI. 3, Figs. 6, 9, 15. Palmatolepis quadrantinodosa Branson & Mehl, 1934 1965 49. Pelekysgnathus serrata Jentzsch—Le Fevre, p. Palmatolepis quadrantinodosa marginifera Ziegler, 1960

Plate 2, Fig. 30 have from Remarks. This species appears to developed

Icriodus. It is a small, blade-like conodont with a long, 1960b Palmatolepis quadrantinodosa marginifera, n.subsp. — posteriorly-widening pulp cavity. 11 — PI. PI. 6—8. Ziegler, pp. 12, 1, Fig. 6; 2, Figs.

1962b Palmatolepis quadrantinodosamarginifera Ziegler—Ziegler, Genus polygnathus Hinde, 1879 PI. 6—9. p. 75, 7, Figs. Polygnathus asymmetrica Bischoff & Ziegler, 1957 For further Glenister & synonymy, see Klapper (1966,p.820). Polygnathus asymmetrica asymmetrica Bischoff & Ziegler, 1957 Remarks. This species has a broad, rounded posterior Plate 2, Fig. 36 and the part a prominent ridge on outer platform parallel to the blade. 1957 Polygnathus dubia asymmetrica n.subsp. BischofT & PI. PI. 3. Ziegler, pp. 88—89, 16, Figs 20—22; 21, Fig. Palmatolepis rugosa Branson & Mehl, 1934 1958 Polygnathus dubia asymmetrica Bischoff & Ziegler— Palmatolepis ampla Miiller, 1956 rugosa Ziegler, PI. 1, Figs. 4—6, 8, 10. Plate 2, Fig. 31 1964 Polygnatus asymmetrica asymmetrica BischofF & Ziegler— Ziegler, Klapper & Lindstrom. 423. 1956 p. Palmatolepis (Palmatolepis) ampla n.sp. Muller, p. 28, For further & synonymy, see Glenister Klapper (1966, p. PI. 9, Figs. 35, 36. 828). 1960 c in Palmatolepis rugosa ampla Miiller—Ziegler Kronberg

PI. 13. etal, p. 38, 1, Figs. 3—5; Text-figs. 12, Remarks. Ziegler, Klapper & Lindstrom (1964) have PI. 1962b Palmatolepis rugosa ampla Miiller—Ziegler, p. 78, 8, pointed out that Polygnathus dubia Hinde, 1879 is a Fig. 6. that nomen dubium. They proposed Polygnathus robusti-

costata Bischoff & Ziegler, 1957 be recognized as the Remarks. The ornamentation of this species is less of new types species Polygnathus. Furthermore, they pronounced than that of the other subspecies of for proposed a new subspecific name Polygnathus The inner Palmatolepis rugosa. posterior part of the dubia dubia, i.e. Polygnathus asymmetrica ovalis Ziegler platform is very large. & Klapper. The from Polygnathus Palmatolepis subrecta Miller & Youngquist, 1947 genus Palmatolepis developed : in the lowermost Frasnian. Relations with Plate 2, Fig. 32 asymmetrica palmatolepids can be found in Ziegler (1962a, Fig. 8).

1947 subrecta — Miller & Palmatolepis n.sp. Youngquist, p. communis Branson & 1934 513, PI. 75, Figs. 7—10. Polygnathus Mehl,

For Glenister & communis communis Branson & 1934 synonymy, see Klapper (1966, p. 823). Polygnathus Mehl, Plate 2, Fig. 37

Remarks. The posterior of the platform is arched part 1934b communis Branson & Polygnathus n.sp. Mehl, p. 293, downward. Relations with other Frasnian palmatole- PI. 24, Figs. I^. pids can be found in Ziegler (1962a, Fig. 8). Palma- 1964 Polygnathus communis communis Branson & Mehl —

subrecta is abundant in our Frasnian material. tolepis Rexroad & Scott, p. 33, 34, PL 2, Figs. 17, 18.

For further Rexroad & Scott synonymy, see (1964). Palmatolepis transitans Miiller, 1956

Plate 2, Fig. 33 Remarks. The earliest known occurrence of Polygnathus

1956 transitans —• communis communis in the Cantabrian Mountains is in Palmatolepis (Manticolepis) n.sp. Miiller,

PI. the illustrated p. 18, 1, Figs. 1,2. quadrantinodosa Zone. The specimen

1956 — comes from this zone. It has the charac- Palmatolepis (Manticolepis) cruciformis n.sp. Miiller, already

PI. 9. teristic excavation behind p. 9, 2, Fig. the pulp cavity. 184 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Polygnathus communis carina Hass, 1959 small. For comparisons with Polygnathus longipostica,

Plate 2, Fig. 43 see there. Klapper (1966) has indicated the great

variability range of Polygnathus inornata. 1959 Polygnathus communis Branson & Mehl carina n.var. 391, PI. Figs. 8, 9. Hass, p. 47, Polygnathus linguiformis Hinde, 1879

1964 Polygnathus communis carina Hass—Rexroad & Scott, Polygnathus linguiformis linguiformis Hinde, 1879 34, PI. 2, 25. p. Figs, 24, Plate 2, Fig. 44; Plate 3, Fig. 1

1879 367, PI. ornament Polygnathus linguiformis n.sp. Hinde, p. 17, Remarks. Two or more nodose ridges the Fig. 15. anterior side of the platform of this subspecies of 1966 Polygnathus linguiformis Hinde—Philip, 448—449, Polygnathus communis. pp. PI. 2, Figs. 29—40.

Polygnathus cf. P. communis communis 1966 Polygnathus linguiformis linguiformis Hinde—Wittekindt,

PI. 10—12. Branson & Mehl, 1934 pp. 635—636, 2, Figs.

Plate 38 For further Orr and Witte- 2, Fig. synonymy, see (1964, p. 16, 18) kindt (1966).

1959 Polygnathus communis Branson & Mehl, 1934? — Voges,

290. p. Remarks. The illustrated specimen, AB 57, shown here

in PI. 3, Fig. 1 derives from the lowest fauna (sample Remarks. Conodonts closely resembling Polygnathus com- VIII 12) with Polygnathus linguiformis linguiformis from

munis communis but lacking the characteristic excava- the Rio Esla area. It has an Emsian (most probably tion behind the classified pulp cavity were as Polygna- lower Remarkable is its and Emsian) age. very large thus cf. P. communis communis. wide, Pseudopolygnathus-like pulp cavity. Philip (1966)

also found this characteristic mark in lower Emsian Polygnathus cristata Hinde, 1879 specimens of Polygnathus linguiformis of southeastern Plate 2, Fig. 41 Australia. He pointed out that in the Lower Devo- — 1879 cristatus 366, PI. 17, nian of Prof. H. Walliser observed Polygnathus n.sp. Hinde, p. Europe, Dr. O. Fig. 11. forms with an enlarged pulp cavity occurring together

1957 cristata Hinde—BischofF& 86— Polygnathus Ziegler, pp. with forms with a small pulp cavity. Samples VIII 12 PI. PI. 13. 87, 15, Figs. 13, 16; 17, Figs. 12, and lb 8, both from the Rio Esla area, contain only

cristata PI. 1966 Polygnathus Hinde—Ziegler, pp. 670—671, specimens of Polygnathus linguiformis linguiformis with 4, Figs. 17—23; PI. 5, Figs. 1—5. a large pulp cavity. All higher Emsian and Middle

For further see Orr (1964, 13). synonymy, p. Devonian specimens of this species have a normal, small pulp cavity.

Remarks. This species has a broad, oval platform orna-

linguiformis mucronata Wittekindt, 1966 mented with large unfused nodes. Polygnathus Plate 3, Fig. 2 Polygnathus cf. P. eiflia Bischoff & Ziegler, 1957

1966 Polygnathus linguiformis mucronata n.subsp. — Witte- Plate 2, Fig. 42 PI. 13—15. kindt, p. 636, 2, Figs.

1965 Polygnathus pennata Hinde—van Adrichem Boogaert, Table 5. its p. 169, Remarks. Only one specimen of this subspecies with

pointed posterior tip lacking transverse ribbing was

Remarks. Wittekindt stresses in a found in the Gildar-Monto (1966, p. 633) new area. definition of Polygnathus eiflia the importance of the Polygnathus longipostica Branson & Mehl, 1934 asymmetric platform and the narrow, constricted Plate 3, 4 anterior Because of the slenderness of Figs. 3, part. the pos- terior of the in have part platform our specimens, we 1934b — Branson & Polygnathus longipostica n.sp. Mehl, p. classified them cf. as Polygnathus P. eiflia. 294, PL 24, Figs. 8—11.

For Rexroad & Scott synonymy, see (1964, p. 36—37). Polygnathus inornata E. R. Branson, 1934

Plate 2, Figs. 39, 40 Remarks. This species is related to Polygnathus inornata,

from which it can be by its more slender 1934 inornata — E. R. PI. distinguished Polygnathus n.sp. Branson, p. 309, free 25, Figs. 8, 26. form, longer blade, more pronounced or-

namentation of the and 1938b lobata — Branson & platform, unusually large Polygnathus n.sp. Mehl, p. 146, ends PI. 34, Figs. 44—47. pulp cavity. The carina of mature specimens

For further 19 with denticle. The lowest synonymy, seeKlapper (1966, pp. —20). posteriorly a~ large oc-

currence ofPolygnathus longipostica in our material was Remarks. Devonian Middle Polygnathus inornata has a short blade and a in the uppermost to Upper costatus thick, elongated platform with upturned margins. Zone. These high Devonian specimens were mostly

The ornamentation is restricted to faint transverse fairly small. of ridges on the margins. The pulp cavity is usually Klapper (1966, p. 21) presented a comparison Poly- Conodont systematics 185

gnathus inornata, Polygnathus longipostica and Polygnathus Pseudopolygnathus triangula Vogcs, 1959 symmetrica. He points out that Polygnathus longipostica Pseudopolygnathus triangula pinna taVoges, 1959 less stands more or midway between Polygnathus Plate 3, Figs. 9, 10 inornata and Polygnathus symmetrica. We found Poly- 1959 Pseudopolygnathus triangula pinnata n.subsp. Voges, gnathus symmetrica only rarely in our material. Our pp. 302—303, PI. 34, Figs. 59—66; PI. 35, Figs. I—6. concept of Polygnathus longipostica is probably some- For see synonymy, Budinger (1965, p. 76). what wider because we followed the definition and

of Rexroad & Scott synonymy (1964). Remarks. The triangular, transversely-ribbed platform

of this species has on its inner, anterior tip a typical Polygnathus nodomarginala E. R. Branson, 1934

lobe. The pulp is small for a Plate 3, Fig. 5 cavity relatively pseudo- polygnathid.

1934 — E. R. Polygnathus nodomarginala n.sp. Branson, p. Genus Branson & 1941 310, PI. 25, Fig. 10. SCALIOGNATHUS Mehl,

1956 Polygnathus nodomarginata E. R. Branson — Bischoff & Scaliognathus anchoralis Branson & Mehl, 1941

PI. 6. Ziegler, p. 157, 12, Fig. Plate 3, Fig. 11

1959 E. R. Polygnathus nodomarginata Branson—Helms, pp.

1941 anchoralis — Branson & Scaliognathus n.sp. Mehl, 102, 651—652, PI. 3, Fig. 1. p. PI. 19, Figs. 29—32.

For see synonymy, Budinger (1965, p. 77). Remarks. The unusually large pulp cavity and the long,

thick of ! that platform Polygnathus nodomarginata suggest Remarks. This remarkable species is easily recognized

this species stands more or less midway between by its cross-like to blunt arrow-like shape. Pseudopolygnalhus fusiformis and Polygnathus longipostica. Genus SCAPHINGNATHUS Ziegler, 1960

Polygnathus varca Stauffer, 1940 Scaphignathus velifera Ziegler, 1960 Plate 3, Fig. 6 Plate 3, Fig. 12

1940 — PI. Polygnathus varcus n.sp. Stauffer, 430, 60, p. 1959 PI. Scaphignathus velifera Ziegler—Helms, p. 655, 2, Figs. 49, 53[?], 55. Fig. 19; PI. 5, Figs. 20, 28. 1957 Stauffer—Bischoff& 98— Polygnathus varca Ziegler, pp. 1960b — PI. Scaphignathus velifera n.sp. Ziegler, p. 403, 3, 99, PI. 18, Figs. 32—35; PI. 19, Figs. 7—9. Fig. 1—6. For further Wittekindt 639 synonymy, see (1966, pp. —640). 1962b Scaphignathus velifera Ziegler—Ziegler, pp. 102—103, PI. 11, Figs. 19—24.

Remarks. This has a free blade and species very long 1965 PI. Scaphignathus velifera Ziegler—Budinger, p. 78, 4, often The is small, asymmetric platform. pulp cavity Fig. 17.

situated at the very anterior part of the platform. For with Wittekindt comparisons Polygnathus xyla, see This Remarks. species has a very high, short free blade, (1966). which is offset slightly laterally with respect to the platform in adult specimens. The elongated Genus pseudopolygnathus Branson & Mehl, 1934 narrow,

platform consists of two lateral rows of denticles. In Pseudopolygnathus dentilineata E. R. Branson, 1934 adult forms one of these lateral rows extends some- Plate 3, Fig, 8 what further anteriorly than the other. A median

1934 dentilineata E. R. row of low nodes is in mature Pseudopolygnathus n.sp Branson, p. developed specimens. 317, PI. 26, Fig. 22. Genus For SCHMIDTOGNATHUS 1965 synonymy, see Klapper (1965, pp. 14—15). Ziegler,

Schmidtognathus peracuta (Bryant, 1921) Plate Remarks. The of Pseudo- 3, Fig. 13 narrow, elongated platform

dentilineata is ornamented with of polyignathus a row 1921 — PI. Polygnathus peracutus n.sp. Bryant, p. 25, 10, nodes each side of the carina. or transverse ridges on Fig. 12. The right side of the platform is more fully developed. 1934 PI. Polygnathus peracuta Bryant—Huddle, p. 97, 8, Fig. 8 Pseudopolygnathus fusiformis Branson & Mehl, 1934

1966 Schmidtognathus peracuta (Bryant) — Ziegler, 668, Plate 3, Fig. 7 p. PI. 1, Figs. 1—10.

1934b — Branson & For further and Pseudopolygnathus fusiformis n.sp. Mehl, synonymy description, see Ziegler (1966)

PI. 1—3. p. 298, 33, Figs.

For Rexroad & Scott synonymy, see (1964, pp. 38—39). The is characterized Remarks. genus Schmidtognathus by its usually fairly big, asymmetric pulp cavity. On and Remarks. This species has an elongated very the outer side of this cavity a constriction (fold) is

reduced platform. The margins of the platform are present. The platform is Polygnathus-like. Schmidto- with low with has slender ornamented nodes. For relations Poly- gnathus peracuta a strongly arched, platform ornamentedwith of gnathus nodomarginata, see there. longitudinally arranged rows nodes. 186 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

Genus SIPHONODELLA Branson & Mehl, 1944 is deepest anteriorly, where the pulp cavity closes rather and continues in the keel Siphonodella cooperi Hass, 1959 abruptly as a groove of the blade. Plate 3, Fig. 14 Remarks: is classi- Siphonodella? n.sp. a provisionally PI. 1959 Siphonodella cooperi n.sp. Hass, p. 392, 48, Figs. fied be in the genus Siphonodella because it seems to 35—36. related to Siphonodella sulcata, which also has a widely For synonymy, see Klapper (1966, p. 16) flared pulp cavity. The latter has a regularly orna- mented platform with strong, straight, transverse Remarks. The slender of this elongated platform ridges. Because of the scarcity and bad preservation

bears two or three rostral on its anterior species ridges our we of material, leave the nomenclatureopen.

The outer is ornamented with trans- part. platform Distribution. a was found in Spain. Siphonodella? n.sp. the inner with nodes. verse ridges, the Cantabrian Mountains in the Vegamian Forma- of the anchoralis tion, sample SJ/H6 (lower part Zone, Siphonodella sulcata (Huddle, 1934)? Section MA, Fig. 18), and in the basal part of the Plate 3, Fig. 16 Alba Formation, sample AG 6 (anchoralis Zone, section

1934 sulcata — PI. 8, In both the found in a Polygnathus n.sp. Huddle, p. 101, AG, Fig. 11). cases species was

Figs. 22—23. fauna including numerous reworked elements. It is

et 1962 sulcata Collinson al., 6, that a from the above- Siphonodella (Huddle) p. very probable Siphonodella? n.sp. Chart 2. mentioned localities is also reworked.

Genus SPATHOGNATHODUS Branson & Mehl, 1941 Remarks. One specimen in our material has an affinity Spathognathodus aculeatus (Branson & Mehl, 1934) with Siphonodella: This is a single prominent rostral Plate 3, Fig. 19 ridge on the outer side of the platform. The ornamen-

tation consists of transverse ridges. The aboral side

1934a aculeatus — Branson & Spathodus n.sp. Mehl, p. 186, has a big and long Polygnathus-like or Pseudopolygnathus- PI. 17, Figs. 11, 14. like pulp cavity and a broad keel. We classify this

1934 tridentatus — E. R. Branson, 307, PL Spathodus n.sp. p. specimen as Siphonodella sulcata? 27, Fig. 26.

1959 aculeatus & — Siphonodella obsoleta Hass, 1959 Spathognathodus (Branson Mehl) Helms, 657, PI. 3, Fig. 8. Plate 3, Fig. 15 p. For further synonymy, see Klapper (1966, p. 24).

1959 obsoleta PI. Siphonodella n.sp. Hass, p. 392, 47, Figs. I—2. Remarks. Spathognathodus aculeatus has one to five, but For synonymy, see Klapper (1966, p. 17) mostly three, denticles situated parallel to the blade,

above the pulp cavity. In some specimens the anterior Remarks. obsoleta has but Siphonodella normally two, transition lateral denticle is the largest, thus showing a sometimes up to four, rostral ridges. The single rostral to Spathognathodus anteposicornis Scott.

ridge on the outer side of the platform continues Bischoff & 1957 posteriorly on the outer margin. The inner side of the Spathognathodus bipennatus Ziegler,

platform is ornamented with nodes, the outer side Plate 3, Fig. 21

is smooth to nearly smooth.

1957 — Bischoff & Spathognathodus bipennatus n.sp. Ziegler, 115—116, PI. 31. Siphonodella? n.sp. a pp. 21, Fig.

Plate 18 — 3, Figs. 17, 1966 [non] Spathognathodus bipennatus nevadensis n.subsp.

Clark & PI. 11 Ethington, p. 687, 84, Figs. 1, 6, 8, 10, (= Eognathodus Philip, 1965). Material. 10 more or less damaged specimens.

Diagnosis. Platform conodont with a long, wide pulp

of of Remarks. This has a blade of double cavity over most the length the platform. The species consisting a of denticles in its central and The oral side is ornamented with low transverse ridges. row posterior part.

The upturned margins of the platform tend to form pulp cavity has two laterally flaring lips. rostral ridges. Spathognathodus costatus (E. R. Branson, 1934) Description. Oral view: The thick, slightly curved plat- Spathognathodus costatus costatus (E. R. Branson, 1934) form is constricted anteriorly. The margins are up- Plate 3, Figs. 20, 22 turned and tend to form rostral ridges, particularly

on the side of the outer, more plat- 1934 — E. R. 303 broadly-developed Spathodus costatus n.sp. Branson, pp. —304, form. The is ornamented with somewhat platform low, PI. 27, Fig. 13. irregular, transverse ridges. The carina consists of 1956 Spathognathodus costatus (E. R. Branson) — Bischoff &

round nodes. The blade is unknown. Lateral PI. 3. fused, Ziegler, p. 166, 13, Fig. view:The conodontis arched. Aboral view: The slightly 1962b Spathognathodus costatus costatus (E. R. Branson) — cavity is long and wide and has flaring lips. PI. 1—6, 8—10. pulp very Ziegler, pp. 107—108, 14, Figs.

It most of the of the and For further runs over length platform, synonymy, see Ziegler (1962b) Conodont systematics 187

1964 steinhornensis steinhornensis Remarks. In Spathognathodus costalus costatus a parallel Spathognathodus Ziegler— Walliser, 85. row of denticles accompanies the blade from the p.

1965 steinhornensis Ill — middle of the anterior half to the posterior tip. The Spathognathodus Ziegler—Philip, pp.

of this be short 112, PI. 10, Figs. 1—12. denticles row may developed as 1966 steinhornensis & transverse ridges. [?] Spathognathodus Ziegler—Clark PI. Ethington, p. 686, 84, Fig. 2. Spathognathodus costalus spinulicostatus (E. R. Branson, 1934) Remarks. Walliser (1964, p. 85) has demonstrated that Plate 24 3, Figs. 23, this species is extremely variable. He distinguished three subspecies: 1934 — E. R. 305, Spathodus spinulicostatus n.sp. Branson, p. Spathognathodus steinhornensis eosteinhornensis (uppermost PI. 27, Fig. 19. Silurian), Spathognathodus steinhornensisremscheidensis (lo- 1957 Spathognathodus spinulicostatus spinulicostatus (E. R. wermost Devonian), and Spathognathodus steinhornensis — PI. Branson) Bischoff, p. 57, 4, Fig. 27. steinhornensis(higher in the Lower Devonian) .These sub- 1962b Spathognathodus costatus spinulicostatus (E. R. Branson) in species show so transitional forms that only — PI. 11 — 18. many Ziegler, p. 108, 14, Figs.

For further faunas can an be made synonymy, see Ziegler (1962b). large adequate separation according to the mean of the distribution of their

Remarks. This subspecies of Spathognathodus costatus has, properties. The illustrated specimen derives from a of lower? Emsian The blade is rather besides a row of denticles on one side of the blade, sample age.

second the of the other denticulated and the wide, a shorter row on posterior part irregularly posteriorly side of the blade. placed pulp cavity is asymmetrically cardiform.

A a Spathognathodus costatus ultimus Bischoff, 1957 n.gen. n.sp. Plate 68 in text. Plate 3, Fig. 25 3, Figs. 28, 29; Fig.

1957 ultimus — Spathognathodus spinulicostatus n.subsp. Material. 3 specimens. PI. Bischoff, pp. 57—58, 4, Figs. 24—26. Diagnosis. Simple type of compound conodont with 1962b ultimus Spathognathodus costatus Bischoff—Ziegler, p. denticle behind the main and denticle one cusp. Cusp 109, PI. 14, Figs. 19, 20. are laterally compressed. The long, narrow pulp For further synonymy, see Ziegler (1962b). cavity has two apices, the highest one pointing into

the main cusp, the lower one into the posterior den- Remarks. This subspecies is distinguished from Spatho- ticle. The surface of the conodont is ornamented with the of gnathodus costatus spinulicostatus by development fine longitudinal striations.

transverse ridges on both sides of the blade.

Spathognathodus jugosus (Branson & Mehl, 1934) Plate 3, Fig. 26

1934a — Branson & 190— Spathodusjugosus n.sp. Mehl, pp.

191, PI. 17, Figs. 19,22.

1956 Spathognathodus jugosus (Branson & Mehl) — Bischoff

PI. 8—10. & Ziegler, p. 167, 13, Figs.

For further synonymy, see Klapper (1966, p. 24).

Remarks: Spathognathodus jugosus developed from

Spathognathodus stabilis as judged from its generally slender outline and long, wide pulp cavity. The of the blade consists of double posterior part a row of small denticles which are mostly fused by trans- verse ridges. In some mature specimens a third row of small nodes may be present between the two main rows.

Spathognathodus steinhornensis Ziegler, 1965

Spathognathodus steinhornensis steinhornensis Ziegler, 1956

Plate 3, Fig. 27

1952 Spathognathodus aculeatus Branson & Mehl Graves,

~ PI. 3 p. 612, 81, Fig. [non Figs. 1, 4, s].

1956 steinhornensis — 104— Spathognathodus n.sp. Ziegler, pp. 68. A AB 84 Fig. n.gen. n.sp. a. Specimen (sample pol 1, 105, PI. 7, Figs. 3—10. Arauz-Polentinos area), photographed in trans-

1958 Spathognathodussteinhornensis Ziegler—Bischoff & Sanne- mitted light (200 x ) showing the pulp cavity

9. with and the fine striation. mann, pp. 106—107, PI. 13, Figs. 2, 3, 7, two apices longitudinal 188 H. A. van Adrichem Boogaert: Conodonts of the Cantabrian Mountains

lower and middle Ordovician. Distribution. Spain. The lowermost part of the Multioistodus, however,

Lebanza Formation (sample pol 1) in the Arauz- does not have a pulp cavity with two apices. Our material shows Polentinos area of the Cantabrian Mountains. The scanty of this new genus two dif- of this of the Lebanza Formation is ferent forms. One is age part upper specimen (PI. 3, Fig. 28) some-

Gedinnian to lower Siegenian. what shorter and more flattened than the other two

Remarks. Dr. M. Lindstrdm (pers.comm) pointed out (PI. 3, Fig. 29). Until more material becomes available,

that the new genus has some resemblance to Multiois- we prefer to refrain from making a more detailed is from classification and leave the nomenclature todus Cullison, 1938, which known only the to open.

Samenvatting

tieve eenheden Een kort overzicht van de litteratuur betreffende de gebieden. Als paleogeografische wor-

stratigrafie van het Devoon en het Onder-Karboon den onderscheiden het Asturo-Leonese Bekken, het Geanticlinaal. van het Cantabrisch Gebergte gaat vooraf aan de Palentijnse Bekken en de Asturische stratigrafische en paleontologische waarnemingen van Aan de hand van acht kaartjes van het faciespatroon

de auteur in León: het Río Esla gebied (Gedinnien tot wordt de ontwikkeling van deze eenheden van het

Viséen), het centrale Cantabrische gebied (Famennien Midden-Devoon tot het Onder-Karboon behandeld. het Gildar-Montó De sedimentatie de Asturische Geanticlinaal tot Viseen) en gebied (Eifelien tot op was

Viséen); in Asturië: het kustgebied (Frasnien tot beperkt en waarschijnlijk incompleet. Een epeirogene-

Viséen); in Palencia: het Arauz-Polentinos gebied tische opheffing van dit gebied had plaats in het

(Gedinnien tot Givetien), het Cardano-Triollo gebied bovenste Frasnien tot onderin het Famennien. Deze Martín-Valsurvio de de (Eifelien tot Viséen) en het San opheffing wordt gecorreleerd met afzetting van

gebied (Givetien en Famennien tot Viséen); in San- kwartsitische Murcia-Formatiein het beschutte Palen-

tander: het Liébana gebied (Eifelien tot Viséen). tijnse Bekken. Het opgeheven gebied werd bedekt door

Uit de kalksteenformaties uit deze gebieden zijn cono- de Ermita-transgressie in het bovenste Famennien tot

dontenfaunas gewonnen, waarvan de meeste konden onderste Tournaisien. Na sedimentatie bracht worden ingepast in de opeenvolging van conodonten- een onderbreking van de een

zones vastgesteld in Duitsland. Aldus werden nieuwe lokale transgressie in het hogere Tournaisien de Vega-

gegevens over een aantal formaties uit het Cantabri- mián-Formatie. De Alba-transgressie begon in het sch deel het in het bovenste Tour- Gebergte verkregen. grootste van gebied Er wordt aangetoond, dat de Ermita-Formatieook in naisien ofonderin het Viséen. In het Palentijnse Bek-

Asturie en Palencia voorkomt. Deze eenheid heeft ken ving de afzetting van de Alba-Formatie bovenin

maximaal een bereik van het bovenste Famennien tot het Viséen aan.

het onderste Tournaisien. De Cardaňo-Formatieheeft Het systematische gedeelte behandelt de belangrijkste bereik een van het middelste of bovenste deel van het gidsvormen voor de conodontenzones. Als nieuwe

Givetien tot bovenin het Frasnien. De Vidrieros- elementen worden beschreven Icriodus eslaensis n.sp.

Formatie heefteen bereikvan het bovenste deelvan het uit het middelste tot bovenste deel van het Givetien,

Famennien het onderste Tournaisien. bovenuit het lagere tot Siphonodella? n.sp. a, waarschijnlijk Een de bakent Tournaisien A synthese van stratigrafische gegevens en n.gen. n.sp. a, een eenvoudig het Palentijnse faciesgebied af. Het is duidelijk geschei- samengestelde conodont bovenuit het Gedinnien of

den van het Asturo-Leonese faciesgebied door posi- onderuit het Siegenien.

Sumario

Una reseňa corta de la literatura acerca de la estrati- Viseense) y la región de San Martin-Valsurvio del Devónico Carbonífero Inferior de la Fameniense hasta Santan- grafia y (Givetiense y Viseense); en Cordillera Cantabrica precede las observationes estra- der: la región Lebaniega (Eifeliense hasta Viseense).

tigraficas y paleontologicas del autor en las regiones Se han extraido faunas de conodontes de las formatio- siguientes en León: la región del Río Esla (Gedinense nes de caliza de estas regiones, la mayoría de las cuales

hasta Viseense), la región central cantábrica (Fame- se podían correlacionar con aquellas de la sucesión niense hasta establecida in Alemania. Así obtuvieron Viseense) y la región de Gildar-Montó zonal se

(Eifeliense hasta Viseense); en Asturias: la región de la nuevos datos acerca de unas formationes en la Cordille-

costa (Frasniense hasta Viseense; en Palencia: la ra Cantabrica. Queda demostrado que la Formatión Arauz-Polentinos Give- existe Asturias Palencia. region de (Gedinense hasta de Ermita tambien en y Este

tiense), la region de Cardaňo-Triollo (Eifeliense hasta unidad se extiende maximamente desde la parte más Sumario 189

Murcia la alta del Fameniense hasta la parte más baja del Tour- depositión de la Formatión cuarcitica de en

naisiense. La Formatión de Cardaňo se extiende del Cuenca Palentina protejida. La región elevada estaba

Givetiense medio o superior hasta el Frasniense supe- cubierta por la transgression de Ermita en el Fame-

rior. La Formation de Vidrieros se extiende de la niense superior mas alto hasta el Tournaisiense más

del Fameniense inferior hasta el Tour- parte superior bajo.

naisiense más bajo. Después de una interuptión en la sedimentatión una

Una síntesis de los datos estratigráficos delimita la transgressión local llevó la Formatión de Vegamián en facies Claramente Tournaisiense la de la región de la palentina. está separa- el superior. En mayor parte

da de la de la facies astur-leonesa la de Alba en la región por regiones región transgressión empezó parte alta del Tournaisiense el Viseense inferior. positivas. Como unidades paleogeográficas se distin- más o en la Cuenca la Cuenca Palentina En la Cuenca Palentina la sedimentatión de la Forma- guen Astur-Leonesa,

y el Geanticlinal Asturiano. Por medio de ocho facies- tión de Alba empezó en el Viseense superior. demostrada la evolutión de unidades La sistemática de las formas mapas está estas parte trata especialmente

desde el Devónico Medio hastael Carbonífero Inferior. más importantes de las zonas de conodontes. Como sedimentatión el Geanticlinal Icriodus eslaensis La en Asturiano estaba nuevos elementos están descritos n. sp. limitada Una elevatión del Givetiense medio y problamente incompleta. y superior, Siphonodella? n.sp. a,

epirogenética de este geanticlinal tenía lugar en la probablemente del Tournaisiense superior y n.gen. A alta del Frasniense hasta el Fameniense conodonte del Gedinense parte mas n.sp. a, un simple superior o

inferior. Esta elevatión está correlationada con la el Siegeniense inferior.

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Fig. 1 Ancyrodella curvata (Branson & Mehl) Fig. 11, specimen AB 10a (sample CAL 4, Gildar- Specimen AB 1 (sample cal 15, Gildar-Montó area). Montó area), x 40. Specimen with faintly indicated 25. oral aboral view. oral aboral lateral view. X 1a, view; 1b, spur. 11a, view; 11b, view; 11c,

Fig. 12, specimen AB 10 (sample CAL 4, Gildar-Montó Fig. 2 Ancyrognathus asymmetrica (Ulrich & Bassler) area). X 40. Oral view of broad specimen. Specimen AB 3 (sample CU 4, Gildar-Montó area).

X 25. 2a, oral view; 2b, lateral view; 2c, aboral view. Figs. 13-17 Icriodus latericrescens latericrescens Branson & Fig. 3 Ancyrodella rotundiloba rotundiloba (Bryant) Mehl AB 2 CAR Cardaño-Triollo Specimen (sample 29, area). Fig. 13, specimen AB 11 (sample pol 1, Arauz-Polenti- x 25. Oral view of fairly juvenile specimen. nos area), x 40. 13a, oral view; 13b aboral view.

Fig.Youngquist4 Ancyrognathus triangularis Fig. 14, specimen AB 12 (sample bk 1, Arauz-Polentinos 40. Oral view. Specimen AB 4 (sample CAR 26, Cardaño-Triollo area). area). x X 25. 4a, oral view; 4b, aboral view. Fig. 15, specimen AB 13 (sample VIII 6114, Río Esla 40 Oral view. area). x area. Figs. 5,6 Icriodus corniger Wittekindt Fig. 16, specimen AB 14 (sample Id 30, Río Esla area). Fig. 5, specimen AB 5 (sample Vc 210, Río Esla area). X 40. Oral view. X 40. 5a, oral view; 5b, aboral view. Fig. 17, specimen AB 15 (sample IVc 74, Río Elsa area). Fig. 6, specimen AB 6 (sample 85, Liébana area). X 40. X 40. 17a, oral view; 17b, aboral view. 6a, oral view; 6b, aboral view.

bilatericrescens Figs. 7,8 Icriodus cf. I. corniger Wittekindt Figs. 18-20 Icriodus latericrescens Ziegler AB 16 1, Arauz-Polentinos Fig. 7, specimen AB 7 (sample VIII 20, Río Esla area). Fig. 18, specimen (sample pol 40. Oral view. X 40. 7a, oral view; 7b, aboral view. area). X Arauz-Polentinos Fig. 8, specimen AB 8 (sample VIII 30, Río Esla area). Fig. 19, specimen AB 17 (sample pol 2, 40. Oral view. with well- X 40. 8a, oral view; 8b, aboral view. area). X Damaged specimen

developed spur. 9-12 Icriodus eslaensis Figs. n.sp. Fig. 20, specimen AB 18 (sample POL 8, Arauz-Polen- Fig. 9, holotype, specimen AB 9 (sample IVc 74, Rio tinos area). X 40. 20a, oral view; 20b, aboral view. Esla area). X 40. 9a, oral view; 9b, aboral view; 9c, lateral view. Fig. 21 Icriodus woschmidti Ziegler 19 Arauz-Polentinos Fig. 10, specimen AB 9a (sample Id 30, Río Esla area). Specimen AB (sample ar 1, area).

X 40. Small specimen. 10a, oral view; 10b, lateralview. X 40. 21a, oral view; 21b aboral view.

PLATE 2

Fig. 1 Icriodus woschmidti Ziegler Fig. 24 Palmatolepis distorta Branson & Mehl AB 37 Gildar-Montó Specimen AB 20 (sample ar 1, Arauz-Polentinos area). Specimen (sample CU 15, area). off. oral lateral 25. Oral view. X 40. Anterior tip broken la, view; 1b, X view. Fig. 25 Palmatolepis glabra glabra Ulrich & Bassler 2 Icriodus AB 39 Gildar-Montó Fig. sp. a Specimen (sample CU 15, area). Specimen AB 21 (sample 19, Río Esla area). x 40. 2a, X 25. Oral view. oral view; 2b, aboral view. Fig. 26 Palmatolepis glabraelongata Holmes Fig. 3 Gnathodus antetexanus Rexroad & Scott Specimen AB 40 (sample CU 15, Gildar-Montó area). Specimen AB 22 (sample OL 3, Río Esla area). x 25. X 25. Oral view. 3a, oral view; 3b, aboral view. Fig. 27 Palmatolepis gigas Miller & Youngquist Figs. 4-6 Gnathodus bilineatus (Roundy) Specimen AB 38 (sample CAR 25, Cardaño-Triollo AB 23 POS Gildar-Montó Fig. 4, Specimen (sample 3, 25. Oral view ofsmall area). x specimen showing many area). x 25. Oral view. affinities with Palmatolepis hassi.

AB 23a and AB 23b CAR Figs. 5,6, specimens (sample Figs. 28, 29 Palmatolepis gracilis gracilis Branson & Mehl Cardaño-Triollo 25. Oral view. These 13, area). x spe- Fig. 28, specimen AB 41 (sample CAL 8, Gildar-Montó cimens have been deformed by cleavage. area). x 40. Lateral view. Fig. 7 Gnathodus commutatus nodosus Bischoff Fig. 29, specimen AB 42 (sample CAL 8, Gildar-Montó Specimen AB 25 (sample CU 25, Gildar-Montó area). area). x 40. Oral view. The bifurcated posterior tip is X 25. Oral view. considered to be a pathologic anomaly. Fig. 8 Gnathodus commutatus cruciformis Clarke 30 Specimen AB 24 (sample CU 25, Gildar-Montó area). Fig. Palmatolepis quadrantinodosa marginifera Ziegler AB 43 CU Gildar-Montó X 25. Oral view. Specimen (sample 15, area). X 25. Oral view. Figs. 9, 10 Gnathodus cuneiformis Mehl & Thomas 31 Müller Fig. Palmatolepis rugosa ampla AB 26 OL Río Esla area). Fig. 9, specimen (sample 1, Specimen AB 44 (sample VID 3, Cardaño-Triollo area). X 25. oral view; 9b aboral view. 9a, X 25. Oral view.

Fig. 10, specimen AB 27 sample) OL 1, Río Esla area). Fig. 32 Palmatolepis subrecta Miller & Younguist X 25. Oral view of large specimen. Blade partly bro- Specimen AB 45 (sample cu 4, Gildar-Montó area). x 25. ken off. oral view.

Figs. 11, 12 Gnathodus cf. G. cuneiformis Mehl & Thomas Fig. 33 Palmatolepis transitans Müller AB 28 OL Esla 46 Fig. 11, specimen (sample 2, Río area). Specimen AB (sample cu 6, Gildar-Montó area). 25. Oral view. x X 25. Oral view.

Fig. 12, specimen AB 28a (sample OL 2, Río Esla area). Figs. 34, 35 Pelekysgnathus serrata Jentzsch X 25. oral view. Fig. 34, specimen AB 47 (sample pol 1, Arauz-Polentinos Figs. 13-15 Gnathodus delicatus Branson & Mehl s.l. area). X 40. 34a, lateral view; 34b, aboral view.

Fig. 13, specimen AB 29 (sample OL 1, Río Esla area). Fig. 35, specimen AB 48 (sample VIII 1, Río Esla area).

X 25. Oral view. x 40. Lateral view.

Fig. 14, specimen AB 30 (sample OL 5, Río Esla area). Fig. 36 Polygnathus asymmetrica asymmetrica Bischoff & Ziegler 25. Oral view. X Specimen AB 49 (sample cu 6, Gildar-Montóarea). x 25.

Oral view. The anterior of this is Fig. 15, specimen AB 30a (sample TR 4’, Cardaño-Tri- part specimen missing. ollo 25. Oral view. area). x Fig. 37 Polygnathus communis communis Branson & Mehl

Fig. 16 Gnathodus girtyi Hass Specimen AB 50 (sample CAL 7, Gildar-Montó area) 40. Small Anterior of blade broken off. Specimen AB 31 (sample POS 3, Gildar-Montó area). X specimen. part oral aboral view. X 25. Oral view. Blade broken off. 37a, view; 37b,

Fig. 38 Polygnathus cf. P. communis communis Branson & Mehl Figs. 17, 18 Gnathodus kockeli Bischoff Specimen AB 52 (sample OL 1, Río Esla area). x 40. AB 32 TR Fig. 17, specimen (sample 21, Cardaño-Triollo Anterior of blade part broken off. 38a, oral view; 38 b, area). x 25. Oral view. aboral view.

Fig. 18, specimen AB 32a (sample TR 21, Cardaño-Triollo Figs. 39, 40 Polygnathus inornata E. R. Branson area). x 25. Oral view. Anterior part of blade broken 39, AB 55 CAL Gildar-Montó off. Fig. specimen (sample 14, area). X 25. 39a, oral view; 39b aboral view. Fig. 19 Gnathodus punctatus (Cooper) Fig. 40, specimen AB 56a (sample PE 4, Asturias, coastal Specimen AB 33 (sample OL 1, Río Esla area). X 25. area). X 25. Oral view of reworked, attrited specimen. Oral view. Anterior part of blade broken off. Fig. 41 Polygnathus cristata Hinde Figs. 20, 23 Gnathodus semiglaber Bischoff Specimen AB 53 (sample CAR 30, Cardaño-Triollo area). 20. AB 34a FE central Canta- Fig. specimen (sample 3, X 25. Oral view. Anterior part ofthis specimen is missing. brian area). X 25. Oral view. Blade broken off. Fig. 42 Polygnathus cf. P. eiflia Bisschoff & Ziegler Fig. 23, specimen AB 24 OL Río Esla (sample 2, area). Specimen AB54 (sample CAR 34, Cardaño-Triolloarea). X 25. Oral view. X 40. Oral view. Blade broken off.

Fig. 21 Gnathodus typicus Cooper Fig. 43 Polygnathus communis carina Hass Specimen AB 35 (sample OL 4, Río Esla X 25. area). Specimen AB 51 (sample OL 1, Rio Esla area). X 25. Oral view. Blade broken off. Blade broken off. 43a, oral view; 43b, aboral view.

22 Gnathodus A Scott & Rexroad 44 Fig. sp. Collinson, Fig. Polygnathus linguiformis linguiformis Hinde Specimen AB 36 (sample TR 21, Cardaño-Triollo area). Specimen AB 57a (sample CU 3, Gildar-Montó area).

X 25. Oral view. X 25. Oral view.

PLATE 3

Fig. 1 Polygnathus linguiformis linguiformis Hinde Fig. 15 Siphonodella obsoleta Hass Specimen AB 73 IIf Río Esla 25. Specimen AB 57 (sample VIII 12, Río Esla area). (sample 224, area). x Oral view. X 25. This specimen has an exceptionally large pulp cavity. la, aboral view; 1b, oral view. Fig. 16 Siphonodella sulcata (Huddle)? AB 71 IIf Río Esla 25. Fig. 2 Polygnathus linguiformis mucronata Wittekindt Specimen (sample 223, area). x 16a, oral view; 16b, latero-oral view; 16c, aboral view. Specimen AB 58 (sample CAL 4, Gildar-Montó area). 25. view. Blade broken off. X Oral 18 Figs. 17, Siphonodella? n.sp. a

17, AB 75 central Canta- Figs. 3, 4 Polygnathus longipostica Branson & Mehl Fig. specimen (sample SJ/H6, brian area). X 35. 17a, oral view; 17b, aboral view. Fig. 3, specimen AB 59 (sample CU 0, Gildar-Montó Fig. 18, specimen AB 76 (sample SJ/H6, central Can- area). x 25. 3a, oral view; 3b, aboral view. tabrian area). X 25. 18a, oral view; 18b, aboral view. Fig. 4, specimen AB 60 (sample CAL 14, Gildar-Montó

area). X 25. 4a latero-oral view; 4b, aboral view. Fig. 19 Spathognathodus aculeatus (Branson & Mehl) Specimen AB 77 (sample FE 1, central Cantabrian area). Fig. 5 Polygnathus nodomarginala E. R. Branson X 25. 19a, oral view; 19b, lateral view. Specimen AB 61 (sample CU 0, Gildar-Montó area). Figs. 20, 22 Spathognathodus costatus costatus (E. R. Branson) x 25. 5a, oral view; 5b, aboral view.

Fig. 20, specimen AB 79 (sample CAL 13, Gildar-Montó Fig. 6 Polygnathus varca Stauffer area). x 25. 20a, lateral view; 20b, oral view. Specimen AB 62 (sample CU 16, Gildar-Montó area). x 25. Fig 22, specimen AB 79a (sample CU 0, Gildar-Mon- Fig. 7 Pseudopolygnathus fusiformis Branson & Mehl tó area). x 25. Oral view.

Specimen AB 64 (sample CU 13, Gildar-Montóarea). Fig. 21 Spathognathodus bipennatus Bischoff & Ziegler small affinities X 25. Oral view of specimen showing Specimen AB 78 (sample IVc 74, Río Esla area). x 25. with nodomarginala. Polygnathus Oral view.

Fig. 8 Pseudopolygnathus dentilineata E. R. Branson Figs. 23, 24 Spathognathodus costatus spinulicostatus (E. R. AB 63 AG 4, Río Esla x 25. Specimen (sample area). Branson) Oral view of large, slightly damaged specimen. Fig. 23, specimen AB 80 (sample CU 0, Gildar-Montó 25. Oral view. Figs. 9, 10 Pseudopolygnathus triangula pinnata Voges area). x

Fig. 9, specimen AB 65 (sample FE 2,central Cantabrian Fig. 24, specimen AB 80a (sample CU 13, Gildar-Montó area). x 25. Oral view. area). X 25. Oral view.

AB 66 OL Río Esla Fig. 10, specimen (sample 1, area). Fig. 25 Spathognathodus costatus ultimus Bischoff 25. 10 oral view; aboral view. X a, 10b, Specimen AB 81 (sample CU 13, Gildar-Montó area).

X 25. Oral view. Fig. 11 Scaliognathus anchoralis Branson & Mehl

Specimen AB 67 (sample OL 2, Río Esla area). X 25. Fig. 26 Spathognathodus jugosus (Branson & Mehl) Oral view. Specimen AB 82 (sample kutt 77, Gildar-Montó area). X 25. 26a, oral view; 26b, lateral view. Fig. 12 Scaphignathus velifera Ziegler

Specimen AB 68 (sample LV 435, Liébana area). x 25. Fig. 27 Spathognathodus steinhornensis steinhornensis Ziegler lateral oral view. 12a, view; 12b, Specimen AB 83 (sample bk 6, Arauz-Polentinos area).

X 25. 27a, lateral view; 27b, oral view. Fig. 13 Schmidlognaihus peracuta (Bryant)

AB 69 CAR Cardaño-Triollo Specimen (sample 30, A Figs. 28, 29 n.gen. n.sp. a area). X 25. 13a, Latero-oral view; 13b, aboral view. Fig. 28, specimen AB 84 (sample pol 1, Arauz-Polen- Fig. 14 Siphonodella cooperi Hass tinos areal. X 40. Lateral view.

Specimen AB 70 (sample SJ/H6, central Cantabrian Fig. 29, specimen AB 85 (sample pol 1, Arauz-Polen-

area). X 25. 14a, oral view; 14b, aboral view. tinos area). X 40. Lateral view.