Plantilla de artículo 2013
Andean Geology 42 (2): 213-236, May, 2015
Andean Geology
doi: 10.5027/andgeoV42n2-a04

The Valanginian Olcostephaninae Haug, 1910 (Ammonoidea) from the Andean Lower Cretaceous Chañarcillo Basin, Northern Chile
Francisco Amaro Mourgues1, 2, Luc G. Bulot3, Camille Frau4

1 IRD-LMTG Observatoire Midi-Pyrénées, 14 Avenue Edouard Belin, 31400 Toulouse, France.

2 Servicio Nacional de Geología y Mineraía, Sección Paleontología y Estratigrafía, Tiltil 1993, Ñuñoa, Santiago, Chile.

3 UMR CNRS 7730 CEREGE, Aix-Marseille Université, Case 67, 3 Place V. Hugo, 13331 Marseille Cédex 03, France.
bulot@cerege.fr

4 9bis Chemin des Poissoniers, 13600 La Ciotat, France.
camille_frau@hotmail.fr

* Permanent adress: TERRA IGNOTA. Heritage & Geosciences Consulting, Dr. Cádiz 726, Ñuñoa, Santiago, Chile.
amourgues@terraignota.cl

Ammonites of the genus Santafecites Etayo-Serna and subgenus Olcostephanus (Viluceras) Aguirre-Urreta and Rawson are described for the first time from Chile. The succession of Olcostephaninae from the Chañarcillo Basin of northern Chile is described in the light of new collections and revision of historical material. The occurrence of mixed Andean and Mediterranean faunas supports the correlations proposed with the ammonite scales of the Neuquén Basin and the Mediterranean Province.

Keywords: Olcostephaninae, Lower Cretaceous, Northern Chile, Systematic Paleontology, Biostratigraphy.        

 

ABSTRACT

1. Introduction

Olcostephanids are an important component of the Valanginian ammonite faunas of the Chañarcillo Basin (northern Chile), where they occur at two discrete intervals in the lower part of succession. The first one is characterized by the conspicuous occurrence of the cosmopolitan Olcostephanus (Olcostephanus) atherstoni (Sharpe, 1856) whereas the other is a level with Olcostephanus (Viluceras) permolestus (Leanza, 1957).

The first Olcostephaninae species mentioned in the Chilean literature is Olcostephanus (O.) curacoensis (Weaver, 1931) but the ammonites were not described and/or illustrated (Tavera, 1956; Corvalán, 1974; Corvalán in Segerstrom et al., 1963). These authors assigned the Argentinean species to different genera, all of which are now regarded as junior objective synonyms of Olcostephanus Neumayr, 1875. It was reported from several localities of the Punta del Cobre and Abundancia formations south and north of Copiapó. O. (O.) curacoensis is herein considered, in agreement with Riccardi et al. (1971) and Aguirre-Urreta and Rawson (1997), a synonym of O. (O.) atherstoni.

The second level is characterized by a very distinctive fauna of evolute olcostephanids. This was recorded by Corvalán in Segerstrom et al. (1963) and Corvalán (1974) with a new species, ‘Holcostephanus copiapoensis nov. sp.’, but neither a description nor an illustration was given; this is therefore a nomen nudum according to the International Code of Zoological Nomenclature. Examination of the specimen labeled ‘Holcostephanus copiapoensis nov. sp.’ in the Corvalán collection by one of us (FAM), allows us to regard this taxon as an invalid synonym of Olcostephanus (Viluceras) permolestus.

The insights issued from this paleontological study permits to discuss and precise the former ammonite biostratigraphic long distance correlations between the European and South Central Andean regions. In this way, the stratigraphic record of the Andean Chañarcillo Basin, represent a link between the Neuquén Basin and Tethys ammonite biostratigraphic record.

2. Stratigraphy and fossil localities

The oldest sequences recorded until now from the marine Chañarcillo Basin are the Punta del Cobre and Abundancia formations, which are respectively assigned to an Early and Late Valanginian age (Mourgues, 2004; Aguirre-Urreta et al., 2007). These outcrops are located southeast of the city of Copiapó, in the northernmost part of the Chañarcillo Basin (Fig. 1). Nevertheless, (?) Middle to Upper Berriasian ammonites (Malbosiceras gr. malbosi in Aguirre-Urreta et al., 2007, fig. 6A) were collected 80 km north of Copiapó and are currently under study.

The Abundancia Formation (‘Capas de Abundancia’, Biese in Hoffstetter et al., 1957) is the oldest lithological unit of the Chañarcillo Group (Segerstrom and Parker, 1959). It is characterized by well bedded grey mudstones interbedded with arkoses, which pass vertically and laterally into the Nantoco Formation limestones, and lies on the Punta del Cobre Formation (Segerstrom and Ruiz, 1962; Marschik and Fontboté, 2001, p. 413). This latter unit is composed of a pile of andesitic to basaltic andesitic lava flows, conglomerates, sandstones, tuffs and dacitic dome complexes, including sedimentary layers in its upper levels. The contact between this unit and the overlying Abundancia Formation is transitional, and has been defined by the first occurrence of a continuous bed of massive limestone or its metamorphosed equivalent. The following localities have been investigated.

 

fig.1

 

FIG. 1. Geographic distribution of the olcostephanidfaunas of the Chañarcillo Basin (simplified after SERNAGEOMIN, 2002). 

 

 

2.1. Cerro La Vinchuca

Discontinuous outcrops of the Chañarcillo Group were observed southwest of Inca del Oro, which were previously investigated by Moraga (1977). Two different levels have yielded ammonites, where two distinctive faunas were identified. From bottom to top these are: Lissonia Gerth, 1925 (to which Raimondiceras Spath, 1924 is likely to be a synonym) and O. (O.) atherstoni. The historical collection of SERNAGEOMIN contains a single specimen herein identified as Santafecites santafecinus (d’Orbigny, 1842) from the same locality (Chong collection, SNGM 07) and described below.

2.2. Puquios area

The Puquios area is characterized by complex tectonics that affects the whole Mesozoic sequence. It is known in the literature as the ‘Caos de Puquios’ and was studied by Sepúlveda and Naranjo (1982) and Mpodozis and Allmendinger (1992). These authors describe a series of 200 m of bioclastic and oolithic limestones that yielded ammonites of Late Valanginian to Barremian age. The material is kept in the collection of SERNAGEOMIN and contains a single specimen of Santafecites santafecinus (Sepúlveda collection, SNGM 1536) described below.

2.3. Quebrada Meléndez

The Quebrada Meléndezis one of the reference sections for the Lower Cretaceous of the Chañarcillo Basin. The succession crops out in a tributary creek, east of the Copiapó valley (Fig. 2). It shows the most complete series of the Upper Valanginian to Lower Aptian strata. From this locality O. (O.) curacoensis and Lissonia riveroi (Lissón, 1907) were reported (Corvalán, 1974). We have observed that near the top of the Abundancia Formation, an ‘Ammonitico rosso’ bed contains abundant O. (Viluceras) permolestus associated with numerous neocomitids, such as juvenile N. (Neocomites) peregrinus (Rawson and Kemper, 1978), Neocomites (Sabbaiceras) beaumugnensis (Sayn, 1907) and Rodighieroites cardulus Company, 1987. These neocomitids are under study with associated material from others localities.

2.4. Quebradas Nantoco-Los Algarrobos

A volcanoclastic succession in the upper part of the Punta del Cobre Formation crops out at Quebrada Los Algarrobos. Two fossiliferous siltstone levels were recognized; a lower one with Lissonia sp., and an upper one containing O. (O.) atherstoni (Fig. 2). From the same locality, previous investigators have mentioned L. riveroi and O. (O.) curacoensis and other neocomitids such as ‘Acanthodiscus’, ‘Thurmanniceras’ by Corvalán in Segerstrom et al. (1963) and ‘Cuyaniceras’ by Marschik and Fontboté (2001).

The upper fossiliferous level with O. (O.) atherstoni was also identified at Quebrada Nantoco. We collected three specimens from that locality (SNGM 1021 [7-9]), which are associated with a specimen of Bochianites sensu lato, several meters above a bed with Lissonia.

fig.2

 

FIG. 2. Lithologic sections of the upper part of the Abundancia Formation (Quebrada Meléndez) and upper part of the Punta del Cobre Formation (Quebradas Nantoco-Los Algarrobos).

 

3. Systematic Paleontology

The material described herein is stored in the Paleontological Collections of the Servicio Nacioal de Geología y Minería (SERNAGEOMIN) Museum, Santiago, with the prefix SNGM. Most of the material belongs to the collection of one of us (F.A.M.). Specimens from the Corvalán, Chong and Sepúlveda collections have also been included in this study.

Dimensions of specimens are indicated as follow: d: diameter; wh: whorl height; wt: whorl thickness; wu: width of umbilicus. M: macroconch; m: microconch. In the synonymy list, v indicates that we have seen the specimen(s).

Superfamily Perisphinctaceae Steinmann, 1890
Family Olcostephanidae Haug, 1910
Subfamily Olcostephaninae Haug, 1910

(=Taraisitinae Cantú-Chapa, 1966; Capeloitinae Cantú-Chapa, 2009; =Garcitinae Cantú-Chapa, 2012; =Saynoceratinae Cantú-Chapa, 2012)

The subfamily Taraisitinae is herein as a junior subjective synonym of Olcostephaninae since its type genus Taraisites Cantú-Chapa, 1966 is a junior subjective synonym of Olcostephanus (Olcostephanus) (see discussion in González Arreola et al., 2014, p. 59). The type genus of Garcitinae is Garcites Cantú-Chapa, 2001, a taxon based on pyritized internal moulds of juveniles olcostephanids. In our opinion the diagnostic features retained by Cantú-Chapa (2001) are diagenetic artifacts due to the alteration of pyrite on the ventral area of the moulds. As a consequence Garcitinae is a nomen dubium based on olcostephanids that cannot be identified at the genus level. Bulot et al. (1990a) showed that the Valanginites-Saynoceras lineage directly derives from the subgenus Olcostephanus by complex heterochronies. Placing Valanginites Kilian, 1910 and Saynoceras Munier-Chalmas, 1894 in different subfamilies is artificial and do not reflects the iterative evolution of the Olcostephaninae. We therefore regard Saynoceratinae as an artificial grouping of micromorphic genera that derivates from Olcostephanus sensu stricto at different moments (Bulot et al., 1990a). The same is true for the Capeloitinae, whose type genus Capeloites Lissón, 1937a, derivates from the Olcostephanus rootstock in the late Lower Hauterivian (Kemper et al., 1981; Bulot, 1990a; Rawson, 2007).

Genus Santafecites Etayo-Serna, 1985

Type-species: Ammonites santafecinus d’Orbigny, 1842, by original designation of Etayo-Serna (1985, p. xxiv-22).

Remarks: In Colombia, Santafecites was reported from the base of the Upper Valanginian together with Saynoceras verrucosum (d’Orbigny, 1841), above the Valanginites, Raimondiceras, Lissonia and Acantholissonia Leanza, 1972 assemblage that characterize the Lower Valanginian (Etayo-Serna, 1985 and personal communication, 1999). It should be noted that there is striking similarities between S. santafecinus and Sphaeroceras broggianus Lissón, 1937b from the Valanginian of Peru. This later species that has been differently interpreted in the literature (Benavides-Cáceres, 1956; Nikolov, 1965; Riccardi and Westermann, 1970; Klein, 2005) and recently designated as the type species of Peruvites by Cantú-Chapa (2012). Unfortunately the exact stratigraphic position of the Peruvian taxon is poorly constrained even if it seems closely allied to Valanginites argentinicus Leanza and Wiedmann, 1980 from the early Lower Valanginian of the Neuquén Basin. Santafecites is therefore likely to derivate from an endemic Valanginites-like form (=Peruvites), but pending a thoroughful description of the Colombian representative of the latter this hypothesis remains speculative.
Occurrence: Early Upper Valanginian of Colombia (Etayo-Serna, 1985). The genus is herein reported from Chile for the first time (?Upper Valanginian). Ammonites santafecinus was also mentioned from Perú by Steinmann (1930) and Lissón and Boit (1942) but this occurrence was never confirmed.

Santafecites santafecinus (d’Orbigny, 1842)
Figs. 3, 4A-J

v 1842 Ammonites santafecinus d’Orbigny, p. 34, Pl. 1, Figs. 3, 4.
1985 Santafecites santafecinus (d’Orbigny), Etayo-Serna, p. xxxiv-22.

Holotype: By monotypy, the specimen figured by d’Orbigny, 1842, p. 34, Pl. 1, Figs. 3, 4, from Santa Fe (de Bogotá), Colombia, kept in the Museum National d’Histoire Naturelle de Paris (d’Orbigny collection, R-3148).

Material: Two specimens (table 1). One large and crushed adult from the Puquios area (SNGM 1536, Sepúlveda collection) and a fragmentary specimen from Cerro La Vinchuca (SNGM 07, Chong collection).

Description: Umbilicus small with poorly defined umbilical region. Ribs organized in bundles with branching at, and above, the umbilical shoulder. The ornamentation seems to be smoother on the outer part of the whorl (? body chamber). No clear bullae or tubercles are visible. The ribs are interrupted on the venter and form a weak groove. They are alternate on both sides of the groove. Suture line is too poorly preserved for description.

Remarks: Since the original figure and description by d’Orbigny (1842), the species was never figured nor described again. The holotype is illustrated here as well as some Colombian specimens (Figs. 4A, B, E, F, H, and I) from the Breistroffer collection (Grenoble University). The larger crushed specimen matches well the holotype (Fig. 3) and the reference material from these collection (compare Figs. 4A, B, E, F, H, and I). It shows all typical features of the genus and species, e.g., ribs originating from periumbilical bullae interrupted on the ventral area and not opposite. The smaller specimen (Dmax ca. 40 mm), represented by half a whorl still septate at D. ca. 36 mm, is slightly different. It is also a typical sphaeroconewith rounded venter and whorl section broader than wide, with the maximum width on the umbilical wall. The smaller specimen (SNGM 07) differs from the Colombian material in its smaller size, lack of bullae and weaker ornamentation. Nevertheless, there is no doubt that it belongs to the genus Santafecites (general morphology, ventral groove and alternate ribs that are interrupted on the venter). It may represent the microconch form of S. santafecinus.

Occurrence: As for genus.

 

fig.3

 

FIG. 3. Santafecites santafecinus (d’Orbigny, 1842). Holotype, MHNP R-3148, d’Orbigny (specimen figured by d’Orbigny, 1842, p. 34, Pl. 1, Figs. 3, 4, from Santa Fe (de Bogotá).

 

 

fig.4

 

FIG. 4. Santafecites santafecinus (d’Orbigny, 1842). A-B, E-F, H-I. side and ventral view of BF-28d to f from Colombia (Breistroffer collection, University of Grenoble); C-D. side and ventral view of SNGM 07 (Chong collection, Cerro La Vinchuca,Chile); G, J. side and ventral view of SNGM 1536(Sepúlveda collection, Puquios, Chile).

 

Genus Olcostephanus Neumayr, 1875

Remarks: As herein understood, the genus Olcostephanus is subdivided into three subgenera: Olcostephanus, Jeannoticeras Thieuloy, 1965 and Viluceras Aguirre-Urreta and Rawson, 1999. Parastieria Spath, 1923a and Mexicanoceras Imlay, 1938 are considered as separate genera. Holcostephanus Sayn, 1889 and Astieria Pavlow, 1892 are junior objective synonyms of OlcostephanusSubastieria Spath, 1923b; Rogersites Spath, 1924; Maderia Imlay, 1938; Taraisites, Satoites Cantú-Chapa, 1966 and Lemurostephanus Thieuloy, 1977 are junior subjective synonyms of Olcostephanus.

Subgenus Olcostephanus Neumayr, 1875

Type-species: Ammonites asterianus d’Orbigny, 1840, p. 115, Pl. 28, Figs. 1, 2 (only), by subsequent designation of Baumberger (1910, p. 3). As already pointed out by Bulot (1990a, p. 5), the specimen from the d’Orbigny collection figured and considered as lectotype by Baumberger (1910, Pl. 32, Figs. 1, 2) does not match the original description or figure of d’Orbigny. The revision of the type material (Bulot, 1990a) suggests that the original drawing is reconstructed from different specimens, some of which belongs to O. (O.) guebhardi (Kilian, 1902), a taxon closely allied to O. (O.) atherstoni (see below); the others to the O. (O.) densicostatus (Wegner, 1909)- sayni (Kilian, 1895) lineage. This view was supported by Busnardo et al. (2003, p. 48-49) and Gauthier et al. (2006, p. 29-30).

Diagnosis: Compressed to strongly inflated cadicones, with strongly arched to well rounded venter. Primary ribs usually present on umbilical wall, commonly terminating in tubercles at umbilical shoulder, from which arise straight or slightly curved secondary ribs, usually in fasciculate bundles. Commonly 3-4 secondaries per bundle, although there may be as many as 6-9 or as few as 2. Secondary ribs may bifurcate on the flanks, while intercalated ribs between bundles are the rule. Ribbing always passes uninterrupted across venter; also it may weaken in some species. Parabolae or constrictions may or may not be present, but are absent on the outer whorls of macroconchs. The genus is dimorphic; microconchs are small and bear lappets on the aperture; macroconchs are larger, with a simple peristome (modified after Cooper, 1981, p. 161).

Occurrence: Olcostephanus (Olcostephanus) is widely distributed in the Mediterran-Caucasian and Indo-Pacific subrealms of the Tethyan Realm and adjacent areas of the Boreal Realm sensu Westermann (2000). In the western Tethys, the subgenus is known to range from the highest Upper Berriasian (F. boissieri Zone, T. otopeta Subzone) to the high Lower Hauterivian,    L. nodosoplicatum Zone, O. (O.) variegatus biohorizon (Bulot, 1990a; Bulot, 1992; Bulot et al., 1993; Bulot and Thieuloy, 1995). In South America, Olcostephanus (Olcostephanus) is known from Colombia, Perú, Chile and Argentina. The oldest occurrence was reported from the early Valanginian of the Neuquén Basin (Aguirre-Urreta and Rawson, 1999) and possibly Colombia (Etayo Serna, written communication 1999). ‘Middle’ Valanginian Olcostephanus of the atherstoni group occur in all countries listed above except Perú (Riedel, 1938; Haas, 1960; Riccardi et al., 1971; this paper). Late early Hauterivian Olcostephanus of the sayni-variegatus (Paquier, 1900) plexus are known from all those countries except Chile (Riedel, 1938; Haas, 1960; Robert et al., 1998; Aguirre-Urreta and Rawson, 2001).

Olcostephanus (Olcostephanus) atherstoni (Sharpe, 1856)
Figs. 5, 6

v   1856    Ammonites Atherstoni Sharpe, p. 196, Pl. 23, Figs. 1a, b.
v   1856    Ammonites baini Sharpe, p. 197, Pl. 23, Figs. 2a-b.
v   1860    Ammonites asterianus (d’Orbigny), Pictet and Campiche, p. 298, Pl. 43, Figs. 1, 2.
1863    Ammonites Schenki Oppel, p. 286, Pl. 81, Figs. 4a-c.
v   1878    Ammonites asterianus (d’Orbigny), Bayle, Pl. 55, Fig. 1 (only).
v   1902    Holcostephanus (Astieria) guebhardi Kilian, p. 866, Pl. 57, Fig. 2.
1902    Astieria cf. atherstoni (Sharpe), Karakasch, p. 11, Pl. 1, Fig. 3.
1903    Holcostephanus (Astieria) schencki (Oppel), Uhlig, p. 130, Pl. 18, Figs. 2a-c.
v   1907    Astieria atherstoni (Sharpe), Baumberger, p. 39-47, Pl. 21, Fig. 3, Pl. 23, Fig. 1, Pl. 24, Figs. 2, 5, Text-figs. 114-117.
v   1907    Astieria cf. atherstoni (Sharpe), Baumberger, Pl. 25, Fig. 4.
v   1908    Astieria leptoplana, Baumberger, p. 9-12, Pl. 26, Fig. 4, Pl. 28, Fig. 2.
v   1908    Astieria imbricata Baumberger, p. 14-18, Pl. 26, Figs. 2-3, Text-figs. 123-126.
v   1908    Astieria actinota Baumberger, p. 18-20, Pl. 26, Fig. 1.
v   1908    Holcostephanus wilmanae Kitchin, p. 195, Pl. 9, Figs. 1,1a.
v   1908    Holcostephanus rogersi Kitchin, p. 201, Pl. 9, Fig. 3, Pl.10, Fig. 2.
v   1908    Holcostephanus modderensis Kitchin, p. 202, Pl.10, Fig. 3.
v   1919    Astieria catulloi Rodighiero, p. 83, Pl. 9, Fig. 9.
v   1930    Rogersites sphaeroidalis Spath, p. 144, Pl. 13, Fig. 5, Pl. 15, Fig. 1.
v   1930    Rogersites otoitoides Spath, p. 149, Pl. 14, Fig. 1.
v   1930    Astieria psilostoma var. lateumibilicata Roch, p. 314, Pl. 16, Figs. 3a, b
1931    Astieria curacoensis Weaver, p. 427, Pl. 49, Figs. 326-327, Pl. 50, Fig. 328.
1931    Astieria sudandina Windhausen, Pl. 33, Fig. 1 (nomen nudum).
v   1932    Rogersites douvillei Besairie, p. 44, Pl. 5, Figs. 9, 9a, Text-fig. 2.
1933    Astieria atherstoni (Sharpe), Roman, p. 21-22, Pl. 4, Fig. 1 (only)
v   1936    Rogersites curvicostatus Besairie, p. 141, Pl. 12, Figs. 7, 10, Pl. 13, Fig. 8.
v   1936    Rogersites douvillei Besairie, p. 138, Text-fig. 9, n° 2.
v   1936    Rogersites baini var. ambikyi Besairie, p. 138, Pl. 13, Fig. 5, Text-fig. 9, n° 3.
1939   Olcostephanus (Rogersites) schencki (Oppel), Spath, p. 30, Pl. 2, Fig. 6, Pl. 18, Figs. 9-10.
1944    Holcostephanus midas Leanza, 1944, p. 16, Pl. 1, Figs. 1a-c.
1944    Holcostephanus auritus Leanza, 1944, p. 18, Pl. 2, Figs. 1a-c.
1958    Olcostephanus curacoensis (Weaver), Corvalán and Pérez, 1958, p. 38, Pl. 12, Fig. 27.
1970 Olcostephanus cf. O. atherstoni Baumberger (non Sharpe), Imlay and Jones, p. B38-B39, Pl. 8, Fig. 15, Pl. 9, Figs. 1-3, 6-10.
v   1962    Holcostephanus schencki Uhlig (non Oppel), Collignon, p. 36, Fig. 859.
v   1962    Holcostephanus atherstoni (Sharpe), Collignon, p. 38, Fig. 860.
v   1962    Holcostephanus douvillei (Besairie), Collignon, p. 43, Fig. 869.
v   1971 Olcostephanus atherstoni (Sharpe), Riccardi et al., p. 91, Pl. 12, Figs. 3-4, Pl. 13, Figs. 1-5, Text-figs. 3-10.
1980    Olcostephanus (Olcostephanus) atherstoni (Sharpe), Leanza and Wiedmann, p. 945, Pl. 1, Fig. 1.
1980    Olcostephanus (Olcostephanus) sakalavensis (Besairie), Leanza and Wiedmann, p. 946, Pl. 1, Fig. 3.
v   1977    Olcostephanus (Olcostephanus) salinarius Spath, Fatmi, Pl. 3, Figs. 1a, b (only).
v   1981    Olcostephanus (Olcostephanus) sp. (= ‘Proastieria’ Stolley), Kemper et al., p. 268-269, Pl. 35, Fig. 2.
v   1981    Olcostephanus (Olcostephanus) atherstoni (Sharpe). Cooper, 1981, p. 182, Figs. 19, 24-26, 27A-D, 28-33, 38, 40-43, 55, 118, 143C-D, 151A.
v   1981    Olcostephanus (Olcostephanus) rogersi (Sharpe), Cooper, 1981, p. 221, Figs. 70-73, 74A-B, 75A-B, 76, 131A-B, 150E-G.
v   1981    Olcostephanus (Olcostephanus) baini baini (Sharpe), Cooper, 1981, p. 263, Figs. 114-129, 130A-B, 131C-J, 132-135, 143A-B, 144A-D, 150C-D, 151B-D.
v   1981    Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath), Cooper, 1981, p. 263, Figs. 144E-G, 145-146, 149, 150A-B, F.
v   1981    Olcostephanus (Olcostephanus) ventricosus (von Koenen), Cooper, p. 300, Fig. 154 (only).
v   1981    Olcostephanus atherstoni (Sharpe), Leanza, p. 169-170, Pl. 15, Figs. 1-8, Pl. 19, Figs. 16-17.
v   1985    Olcostephanus astierianus (d’Orbigny), Cecca, p. 157, Pl. 6, Figs. 1a, b.
v   1985    Olcostephanus densicostatus (Wegner), Company, p. 118, Pl. 1, Figs. 3-7.
v 1986  Olcostephanus psilostomus psilostomus (Neumayr and Uhlig), Kvantaliani and Sakharov, p. 56, Pl. 1, Figs. 1, 2.
v   1986    Olcostephanus convolutus (von Koenen), Kvantaliani and Sakharov, p. 57, Pl. 1, Figs. 3.
1986         Olcostephanus cf. guebhardi (Kilian), Howlett, p. 71, Figs. 2a, b.
v   1987   Olcostephanus densicostatus (Wegner), Company, p. 594, Pl. 6, Figs. 5-6.
v   1988    Olcostephanus atherstoni (Sharpe), Riccardi, Pl. 5, Figs. 1-2.
v   1990a Olcostephanus (Olcostephanus) guebhardi (Kilian), Bulot, p. 87, Pl. 6, Figs. 1-10, Pl. 7, Figs. 1-3, Pl. 8, Figs. 1-7, Pl. 11, Figs. 1-2.
v   1990b Olcostephanus (Olcostephanus) guebhardi (Kilian), Bulot, p. 8, Pl. 1, Figs. 4, 5.
v   1991    Olcostephanus (Olcostephanus) guebhardi morphotype querolensis (Kilian), Ettachfini, p. 113, Pl. 11, Figs. 1, 3, 7, 8.
v 1991  Olcostephanus (Olcostephanus) guebhardi aff. morphotype querolensis (Kilian), Ettachfini, p. 113, Pl. 11, Fig. 2.
v   1991    Olcostephanus (Olcostephanus) guebhardi lateumbilicatus (Roch), Ettachfini, Pl. 11, Figs. 4, 6.
v   1991    Olcostephanus (Olcostephanus) cf. psilostomus lateumbilicatus (Roch), Ettachfini, Pl. 11, Figs. 10, 11.
v   1992    Olcostephanus (Olcostephanus) guebhardi morphotype querolensis Bulot, Pl. 1, Fig. 1.
v   1992    Olcostephanus (Olcostephanus) guebhardi (Kilian), p. 151, Bulot, Pl. 1, Figs. 2a, b.
1992    Olcostephanus aff. guebhardi (Kilian), Kemper, Pl. 35, Figs. 1a, b.
v   1993    Olcostephanus (Olcostephanus) atherstoni (Sharpe), Aguirre-Urreta, Pl. 2, Figs. 6, 7.
v   1995    Olcostephanus atherstoni (Sharpe), Aguirre-Urreta and Rawson, p. 453, Figs. 6h-i.
v   1997    Olcostephanus atherstoni (Sharpe), Aguirre-Urreta and Rawson, Pl. 1, Figs. c-e.
1997     Olcostephanus (O.) catulloi (Rodighiero), Faraoni et al., Pl. 5, Fig. 17.
2001    Olcostephanus (Olcostephanus) guebhardi (Kilian), Wippich, Pl. 10, Figs. 2-11, Pl. 11, Figs. 1-2.
2003    Olcostephanus (Olcostephanus) guebhardi (Kilian), Wippich, Fig. 8d, Fig. 9e.
2003    Olcostephanus (Olcostephanus) guebhardi (Kilian), Lukeneder and Harzhauser, Fig. 3a.
2004    Olcostephanus (Olcostephanus) guebhardi (Kilian), Lukeneder, Fig. 6.
2004    Olcostephanus (Olcostephanus) atherstoni (Sharpe), Mourgues, Fig. 5a, g.
2004    Olcostephanus (Olcostephanus) guebhardi lateumbilicatus (Roch), Ettachfini, Pl. 23, Fig. 2-6.
2004    Olcostephanus (Olcostephanus) guebhardi morphotype querolensis Bulot, Ettachfini, Pl. 23,    Fig. 7-10.
2005    Olcostephanus (Olcostephanus) atherstoni (Sharpe), Aguirre-Urreta et al., Fig. 5a.
2007   Olcostephanus (Olcostephanus) guebhardi morph. type querolensis Bulot, Lukeneder, Fig.4F-I.
2007    Olcostephanus (Olcostephanus) atherstoni (Sharpe), Aguirre-Urreta et al., Fig. 6C, G, Fig. 10F.
2010    Olcostephanus guebhardi (Kilian), Vašíček, p. 399, Pl. 1, Fig. 2, Pl. 2, Fig. 2.
?    2010    Olcostephanus guebhardi (Kilian), Fözy et al., Fig. 3R.

 

fig.5

 

FIG. 5. Olcostephanus (O.) atherstoni (Sharpe, 1856). A-B. SNGM 1021-17; C-D. SNGM 1021-14; E-F. SNGM 1021-8; G-H. SNGM 1021-26; I. SNGM 1021-23; J-K. SNGM 1021-19; L-M. SNGM 1021-18; N. SNGM 1021-15; O. SNGM 1021-20; P, T. SNGM 1022-3; Q, U. SNGM 1021-10/11; R. SNGM 1021-29; S. SNGM 1021-16; V. SNGM 1022-15; W. SNGM 1022-4. All specimens from bed PC2 at Quebrada Los Algarrobos (Mourgues collection) except E and F from Quebrada Nantoco.

 

 

fig.6

 

FIG. 6. Olcostephanus (O.) atherstoni (Sharpe, 1856). A. SNGM 1022-7; B. SNGM 1022-15; C-D. SNGM 1022-13; E-F. SNGM 1022-14; G. SNGM 1022-16; H. SNGM 1022-4; I. SNGM 1022-3; J-K. SNGM 1022-2; L-M. SNGM 1022-10; N-O. SNGM 1022-5 from bed PC2 at Quebrada Los Algarrobos (Mourgues collection). All specimens natural size except H (x 0.7).

 

Holotype: By monotypy, the specimen figured by Sharpe (1856, p. 196, Pl. 23, figs. 1a, b), from the Sundays River, South Africa (British Museum, BM-C32202), refigured by Cooper (1981, p. 183-184, figs. 24, 25).

Material: 41 specimens (tables 2 and 3): 3 from Quebrada Nantoco (SNGM 1021 [7-9]), 36 from Quebrada Los Algarrobos (SNGM 1021 [10-28], SNGM 1022 [2-18]) and 2 from 3 km to northwestern of Cerro La Vinchuca (SNGM 1537 [1-2]).

Description: Microconch [m]: the microconch consists of immature examples of 25-30 mm diameter, with at least some part of the body chamber preserved. Septation ceases at diameters of about 20-25 mm. The shell is slightly involute, cadicone with moderately depressed to semicircular whorls. The umbilical slope is moderately steep with a rounded margin. The umbilicus is moderately deep and narrow where its diameter (wu) is approximately 28% of the shell diameter (d). The venter and flanks are rounded. Maximum width of whorl at umbilical bullae. Rursiradiate primary ribs arise near the umbilical seam, progressively becoming prominent on the umbilical slope, and forming bullae-like tubercles on the upper part of the umbilical margin. There are 9-12 primary ribs per half-whorl. Bundles of three to four (exceptionally 5) secondary ribs associated with each bulla, but an additional one or two may be intercalated between two bundles. There are 40-45 secondary ribs per half-whorl. No constrictions were observed our examples.

Macroconch [M]: the shell of the macroconch is moderately small with diameters between 70-85 mm. Some specimens are septate to at least 40-50 mm. The body chamber occupies at least ¾ of the last whorl. These forms are more involute than the microconchs, with the umbilicus (wu) reaching approximately a fifth of the shell diameter (d). The umbilical slope is steep with a rounded margin. Ribbing is finer and denser than in microconchs, and slightly rursiradiate. There are 13-15 primary ribs per half-whorl. Bundles of four to six secondary ribs are associated with each bulla, but an additional rib may be intercalated between two bundles. There are 55-70 secondary ribs per half-whorl. Neither constrictions nor parabolae was observed in our material.

Discussion. As already mentioned by Riccardi et al. (1971), Cooper (1981), Company (1987) and Bulot (1990a, 1992), the Olcostephanus of the atherstoni plexus are characterized by a marked dimorphism. The Chilean material also illustrates this feature (Fig. 7). The dimorphism clearly affects size, coiling and ribbing. The microconchs are more evolute, smaller and with coarser ribs than the macroconchs.

 

fig.7

 

FIG. 7. Plot of umbilical width (wu) against shell diameter (d) for Olcostephanus (O.) atherstoni (Sharpe, 1856) and O. (Viluceras) permolestus (Leanza, 1957).

 

It should also be noted that the material from Atacama is closer to the population from the Mediterranean Tethys than to the one from South Africa and Argentina (Neuquén Basin). As a whole, our population is very similar to O. (O.) guebhardi and its morphotypes as defined by Bulot (1990a, 1992). This includes not only O. (O.) guebhardi s. str. but also O. (O.) guebhardi morphotype querolensis Bulot, the O. (O.) densicostatus (Wegner) in Company (1987, Pl. 15, Figs. 1-8, Pl. 19, Figs. 16, 17).

On the other hand, some of the South African microconchs figured by Cooper (1981, p. 265, Fig. 115, Fig. 129B) under O. (O.) baini baini, considered herein as a junior subjective synonym of O. (O.) atherstoni, hardly differ from the microconchs of O. (O.) guebhardi. Similarly, the larger macroconchs of O. (O.) guebhardi (morphotypes querolensis and hollwedensis) fall within the adult size of the smaller South African and Argentinean O. (O.) atherstoni. As a consequence, we cannot see any reason to keep O. (O.) guebhardi separate from O. (O.) atherstoni. Because of priority rules, O. (O.) guebhardi is a junior subjective synonym of O. (O.) atherstoni.

Nevertheless, it should be noted that in South-eastern France the oldest representatives of O. (O.) atherstoni, O. (O.) guebhardi ‘forme primitive’ in Bulot (1990a, 1992), first occur in the middle part of the B. campylotoxus Subzone (B. subcampylotoxus biohorizon sensu Bulot and Thieuloy, 1995). The acme of the species ranges from the upper part of the B. campylotoxus Subzone to the uppermost part of the K. biassalense Subzone (B. campylotoxus biohorizon to K. inostranzewi Zone sensu Bulot and Thieuloy, 1995). These forms, always cadicones, lose at the adult stage the sphaerocone morphology of the precursors. It is among those populations that the morphology is closer to the typical O. (O.) atherstoni from South Africa. They also include large numbers of typical O. (O.) guebhardi.

At the top of the K. biassalense Subzone, the populations are affected by a new morphological change that affects mainly the density of ribbing, while the general form of the shell remains unchanged. Bulot (1990a, 1992) introduced the name querolensis for this morphotype that shows its acme in the S. verrucosum Subzone and disappears at the top of the K. pronecostatum Subzone. Most of the Chilean macroconchs described herein clearly belong to this morphotype.
Similarly, Ettachfini (1991) showed that the morphotype succession identified in south-east of France is also represented in the Valanginian of the Essaouira basin (Morocco). In South-eastern Spain, O. (O.) atherstoni is represented only by the morphotype querolensis, which shows the same distribution as in France. The German morphotype hollwedensis Bulot (=‘Proastieriasensu Stolley, 1937) is known only from the lower part of the P. hollwedensis Zone of the Lower Saxony and Yorkshire basins of Northern Europe (Kemper et al., 1981). Similar forms also occur in the ‘Marnes à Astieria’ of the Jura platform in Switzerland (Bulot, 1992). This morphology is by far the closest to the typical O. (O.) atherstoni from South Africa. The time span of the hollwedensis morphotypes merely represents an equivalent of the uppermost B. campylotoxus and lower K. biassalense Subzones of the Mediterranean standard ammonite scale.

From the Neuquén basin, the species described by Leanza (1944), O. (O.) auritus and O. (O.) midas, correspond respectively to the macroconchs and microconchs of our O. (O.) atherstoni.

Subgenus Viluceras Aguirre-Urreta and Rawson, 1999

Type-species: Simbirskites permolestus Leanza, 1957, by original designation of Aguirre-Urreta and Rawson (1999, p. 351).

Diagnosis: An evolute to serpenticone subgenus of Olcostephanus characteristically with well-developed constrictions and flares, particularly in the adult growth stage.

Discussion: The relationships between Viluceras, Simbirskites, Lemurostephanus and Olcostephanus have been discussed at length by Aguirre-Urreta and Rawson (1999, p. 343 and 351-352) and we fully agree with those authors on the systematic position of Viluceras as a subgenus of Olcostephanus. As already mentioned above and for the reasons given by Aguirre-Urreta and Rawson (1999, p. 351-352), we also regard Lemurostephanus as a junior subjective synonym of Olcostephanus s. str.

Occurrence: Until the recognition of its occurrence in Chile (Mourgues, 2004), the subgenus was known only from the middle part of the Upper Valanginian of the Neuquén Basin in Argentina (Aguirre-Urreta and Rawson, 1999). The reported occurrence of O. (V.) permolestus from south-east France (Autran, 1993) is erroneous and will be discussed below.

Olcostephanus (Viluceras) permolestus (Leanza, 1957)
Fig. 8

v      1957   Simbirskites permolestus Leanza, p. 16, Pl. 3, Fig. 1.
v      1963   ‘Holcostephanus copiapoensis nov. sp.’: Corvalán in Segerstrom et al., p. 10.
v      1974   ‘Holcostephanus copiapoensis nov. sp.’: Corvalán, p. 19.
v     1980   Olcostephanus (Lemurostephanus) permolestus (Leanza): Leanza and Wiedmann, p. 949, Pl. 1, Fig. 2, Pl. 2, Fig. 1.
v       1993   Olcostephanus (Lemurostephanus) permolestus (Leanza): Aguirre-Urreta, Pl. 2, Fig. 4 (holotype refigured from opposite flank).
non 1993    Olcostephanus (Lemurostephanus) permolestus (Leanza): Autran, Pl. 4, Fig. 2.
v     1995   Olcostephanus (Lemurostephanus) permolestus (Leanza): Aguirre-Urreta and Rawson, Pl. 1g (re-illustration of Aguirre-Urreta, 1993, Pl. 2, Fig. 4).
v     1997   Olcostephanus (Lemurostephanus) permolestus (Leanza): Aguirre-Urreta and Rawson, Figs. 6a, b.
v     1999   Olcostephanus (Lemurostephanus) permolestus (Leanza): Aguirre-Urreta and Rawson, p. 352-353, Figs. 4e, f, I, j; Figs. 5a-n; Fig. 6; Figs. 7a-f, i, j.
v       2002    Olcostephanus (V.) permolestus (Leanza): Mourgues, p. 75.
v       2004   Olcostephanus (Viluceras) permolestus (Leanza): Mourgues, Fig. 5f.
2005   Olcostephanus (Lemurostephanus) permolestus (Leanza): Aguirre-Urreta et al., Fig. 5b.
2007  Olcostephanus (Viluceras) permolestus (Leanza): Aguirre-Urreta et al., Figs. 6D-E, Fig. 10I.

Holotype: By original designation, the specimen figured by Leanza (1957, p. 16, Pl. 3, Fig. 1), from El Durazno hill, Central West Argentina. Universidad de Buenos Aires, CPBA 7018. Refigured by Aguirre-Urreta (1993, Pl. 2, Fig. 4) and Aguirre-Urreta and Rawson (1995, Pl. 1, Fig. g; 1999, p. 350, Fig. 5a).

Material: 19 specimens (table 4 and 5), from Quebrada de Meléndez (SNGM 1023 [1-17]; SNGM 1538 [1, 2], Corvalán collection).

Description: The examples are dimorphic in size. Small microconch of 35-45 mm diametre, serpenticone with a large umbilicus (wu=35-48% of d). The subcircular whorl section overlaps approximately a third of the preceding whorl. The umbilical slope is quite shallow and rounded and becomes moderately steep in the adult macroconch. The rectiradiate primary ribs are coarse, slightly concave, and form bullae-like tubercles on the upper part of the umbilical  margin. From each bulla arise associated bundles of two to three (frequently two) secondary ribs, gently rursiradiate on the upper third of the whorl. An additional one may be intercalated between two bundles. Constrictions are frequent.

Remarks: Both our macroconchs and microconchs fall in the range of intraspecific variation accepted by Aguirre-Urreta and Rawson (1999) for this species (compare our Figs. 8B, E and K with their Figs. 5h, i and n). As in the Argentinean material, microconchs show a significant variation in rib density, degree of inflation, as well as in strength and frequency of constrictions and flares.

Outside southern South America, a single specimen was reported from south-east France by Autran (1993) as O. (Lemurotephanus) permolestus. As already noted by Bulot (1990a) and Aguirre-Urreta and Rawson (1999), this record is based on the misidentification of macroconch forms of O. (O.>) nicklesi.

 

fig.8

 

FIG. 8. O. (Viluceras) permolestus (Leanza, 1957). A, G. SNGM 1538-1; B. SNGM 1023-2; C. SNGM 1023-12; D, J. SNGM 1538-2; E. SNGM 1023-9; F. SNGM 1023-5; H-I. SNGM 1023-8; K-L. SNGM 1023-3. All specimens from bed G3-3 at Quebrada de Meléndez (Mourgues collection), except A, G, D and J (Corvalán collection).

 

4. Biostratigraphic implications

As in the Neuquén Basin, the genus Olcostephanus occurs at two main distinct levels. The lower one is characterized by a monospecific assemblage of O. (O.) atherstoni, while the upper one is marked by the co-occurrence of O. (V.) permolestus with various Neocomitidae such as N. (Neocomites) peregrinus, Neocomites (Sabbaiceras) beaumugnensis and Rodighieroites cardulus.

Both O. (O.) atherstoni and O. (V.) permolestus were retained by Aguirre-Urreta and Rawson (1997, 1999) as index species for zones and subzones of the Valanginian of the Neuquén Basin (Fig. 9). As already pointed out by those authors, the lower and middle part of the O. (O.) atherstoni Zone, O. (O.) atherstoni and Karakaschiceras attenuatum subzones, span the Lower/Upper Valanginian boundary on evidence of ammonites common to Argentina and the Mediterranean Tethys.

Until now, O. (O.) atherstoni was considered by one of us (L.G.B.) as the southern hemisphere counterpart of the North Tethyan species O. (O.) guebhardi. As discussed above in the light of the Chilean material, we are now convinced that the two taxa should be considered as synonyms. For the reasons given in our discussion of the intraspecific variability of O. (O.) atherstoni, we therefore consider that the base of the Argentinean O. (O.) atherstoni Zone correlates with the middle part of the B. campylotoxus Zone of the Mediterranean zonal scheme (Fig. 9).

 

fig.9

 

FIG. 9. Ammonite zonal scheme for the Chañarcillo Basin and proposed correlation with the Neuquén Basin (Aguirre-Urreta and Rawson, 1999; Rawson, 1999) and the West Mediterranean Province (Reboulet et al., 2011).

 

Even if the First Apparition Data (FAD) of Karakaschiceras sensu Bulot and Thieuloy (1995) and Neohoploceras Spath, 1939 occur at the base of the K. biassalense Subzone of the Mediterranean zonal scheme, we herein consider that the Argentinean K. attenuatum Subzone better correlates with the K. pronecostatum Subzone of the Mediterranean zonal scheme. This view is supported by the fact that Neohoploceras arnoldi sensu Aguirre-Urreta (1998) is closely allied, if not identical, to Neohoploceras gr. depereti/schardti sensu Bulot and Thieuloy (1995). Similarly, the general morphology of K. attenuatum (Behrensen, 1892) and K. neumayri (Behrensen, 1892) is closer to that of the group of K. pronecostatum Felix, 1891, including K. quadristrangulatum (Sayn, 1907), than to the late Lower Valanginian forms of the K. biasselense/inostranzewi plexus. As a consequence, the correlation proposed herein is slightly different from that formerly published by Rawson (1999, 2007) and Aguirre-Urreta et al. (2007).

So far, the Argentinean O. (V.) permolestus Subzone has been correlated with the O. (O.) nicklesi Subzone of the Mediterranean ammonite scale (Aguirre-Urreta and Rawson, 1999, p. 356; Aguirre-Urreta et al., 2007; Rawson, 2007). As Viluceras is unknown outside southern South America, this view was mainly based on the co-occurrence O. (O.) mingrammi (Leanza), an evolute Olcostephanus, that shows some morphological affinities with the plexusof O. (O.) nicklesi Wiedmann and Dieni, 1968. Nevertheless, those authors also noted, that in south western Europe, the range of evolute Olcostephanus s. str. started in the N. peregrinus Subzone and carried on in the C. furcillata Subzone with such species as O. (O.) detonii (Rodighiero, 1919) and O. (O.) mittreanus (d’Orbigny, 1850). Our new material shows clearly that O. (V.) permolestus is associated with Neocomites (N.) peregrinus in Chile. We therefore suggest that the O. (O.) permolestus Subzone correlates with the N. peregrinus Subzone of the Mediterranean standard scale of Reboulet et al. (2011) (Fig. 9).

The stratigraphic record of the Chañarcillo Basin represents a bridge between the Neuquén Basin and the Tethys region, where the typical Andean faunas were documented in association with Tethysian ammonites which allows us to precise the former biostratigraphic long distance correlations, for the Valanginian ammonite biozones.

 

Acknowledgements
This research was supported by the following projects: ‘Levantamiento Hoja Vallenar’ (SERNAGEOMIN) and ‘Bilans et processus des systèmes carbonatés’ (Fre CNRS 2761, University of Marseille I). Additional financial support was provided by UMR CNRS 5563- IRD (University of Toulouse III).

Access to collections in Chile was made possible by courtesy of Prof. E. Pérez d’A. and Dr. A. Rubilar (SERNAGEOMIN). Prof. M.B. Aguirre-Urreta (Universidad de Buenos Aires) and Prof. P.F. Rawson (Scarborough) are gratefully acknowledged for access to collections in their care and fruitful discussions on the systematics and biostratigraphy of the olcostephanids of the Neuquén Basin. Personal communications by Prof. Fernando Etayo-Serna (Bogotá) and Dr. C. Arévalo (SERNAGEOMIN) have improved an original draft of this manuscript.

The photographic work was carried out by Mrs. A. Fontana (Santiago) and A. Arnoux (University of Marseille). For their help in the field, we are also very grateful to Mr. P. Herrera (SERNAGEOMIN, Copiapó) and Prof. E. Jaillard (University of Grenoble).

 

References

Aguirre-Urreta, M.B. 1993. Neocomian Ammonite Bio-stratigraphy of the Andean Basin of Argentina and Chile. Revista Española de Paleontología 8: 57-74.

Aguirre-Urreta, M.B. 1998. The ammonites Karakaschiceras and Neohoploceras (Valanginian Neocomitidae) from the Neuquén Basin, West-Central Argentina. Journal of Paleontology 72: 39-59.

Aguirre-Urreta, M.B.; Rawson, P.F. 1995. Estratigrafía y ammonites del Valanginiano superior de la región de Chos Malal, Cuenca Neuquina. In Congreso Argentino de Paleontología y Bioestratigrafía, No. 6, Actas: 7-14. Trelew.

Aguirre-Urreta, M.B.; Rawson, P.F. 1997. The ammonite sequence in the Agrio Formation (Lower Cretaceous), Neuquén Basin, Argentina. Geological Magazine 134: 449-458.

Aguirre-Urreta, M.B.; Rawson, P.F. 1999. Lower Cretaceous ammonites from the Neuquén Basin, Argentina: Viluceras, a new Valanginian subgenus of Olcostephanus. Cretaceous Research 20: 343-357.

Aguirre-Urreta, M.B.; Rawson, P.F. 2001. Lower Cretaceous ammonites from the Neuquén Basin, Argentina: a Hauterivian Olcostephanus fauna. Cretaceous Research 22: 763-778.

Aguirre-Urreta, M.B.; Rawson, P.F.; Concheyro, G.A.; Bown, P.R.; Ottone, E.G. 2005. Lower Cretaceous (Berriasian-Aptian) biostratigraphy of the Neuquén Basin. In The Neuquén Basin, Argentina: A case Study in Sequence Stratigraphy and Basin Dynamics     (Veiga, G.D.; Spalletti, L.A.; Howell, J.A.; Schwarz, E.; editors). Geological Society, Special Publications 252: 57-81. London.

Aguirre-Urreta, M.B.; Mourgues, F.A.; Rawson, P.F.; Bulot, L.G.; Jaillard, E. 2007. The Lower Cretaceous Chañarcillo and Neuquén Andean basins: ammonoid biostratigraphy and correlations. Geological Journal 42: 143-173.

Autran, G. 1993. L’évolution de la marge nord-est Provençale (Arc de Castellane) du Valanginien moyen à l’Hauterivien à travers de l’analyse biostratigraphique des séries de la région de Peyroules: séries condensées, discontinuités et indices d’une tectonogenèse distensive. Annales du Muséum d’Histoire Naturelle de Nice, Paléobiologie 10: 1-240.

Baumberger, E. 1907. Die ammonitiden der unteren Kreide im der westschweizerischen Jura. Mémoire de la Société Paléontologique Suisse 34: 1-47.

Baumberger, E. 1908. Die ammonitiden der unteren Kreide im der westschweizerischen Jura. Mémoire de la Société Paléontologique Suisse 35: 1-40.

Baumberger, E. 1910. Die ammonitiden der unteren Kreide im der westschweizerischen Jura. Mémoire de la Société Paléontologique Suisse 36: 1-57.

Bayle, E. 1878. Fossiles Principaux des Terrains. Service de la Carte Géologique détaillée de la France. Explication de la Carte géologique de France 4 (1), Atlas: 158 pls. Paris.

Behrendsen, O. 1892. Zur Geologie des Ostabhanges der argentinischen Cordillere. Zeitschrift Deutsche Geologische Gesellschaft 44: 1-42

Benavides-Cáceres, V.E. 1956. Cretaceous System in Northern Perú. Bulletin of the American Museum of Natural History 108: 353-494.

Besairie, H. 1932. Fossiles caractéristiques du nord et du nord-ouest de Madagascar. Annales géologiques du Service des Mines de Madagascar 2: 37-53.

Besairie, H. 1936. Recherches géologiques à Madagascar, Première suite: la Géologie du Nord-Ouest. Mémoires de l’Académie Malgache 21 : 1-259.

Bulot, L.G. 1990a. L’evolution des Olcostephanidae (Ammonitina, Cephalopoda) dans le context paléobiogéographique du Crétacé Inférieur (Valanginien-Hauterivien) du Sud-Est de la France. Diplôme supérieur d’Etude et de la Recherche (Unpublished), Université de Bourgogne: 178 p.

Bulot, L.G. 1990b. Révision des types et figurés de la collection Matheron 2. Olcostephanus (Olcostephanus) perinflatus (Matheron, 1878) et Olcostephanus (Olcostephanus)? mittreanus (Matheron non d’Orbigny, 1850). Mésogée 50: 3-8.

Bulot, L.G. 1992. Les Olcostephaninae valanginiens et hauteriviens (Ammonitina, Cephalopoda) du Jura franco-suisse: Systématique et intérêt biostratigraphique. Revue de Paléobiologie 11: 149-166.

Bulot, L.G.; Thieuloy, J.-P. 1995. Les biohorizons du Valanginien du Sud-Est de la France: un outil fondamental pour les corrélations au sein de la Téthys occidentale. In Lower Cretaceous Cephalopod Biostratigraphy of the Western Tethys, Géologie Alpine, Mémoire Hors Série 20: 15-41.

Bulot, L.G.; Company, M.; Thieuloy, J-P. 1990. Origine, évolution et systématique du genre Saynoceras. Géobios 23: 399-413.

Bulot, L.G.; Thieuloy, J.-P.; Blanc, E.; Klein, J. 1993. Le cadre stratigraphique du Valanginien supérieur et de l’Hauterivien du Sud-Est de la France: définition des biochronozones et caractérisation de nouveaux horizons. Géologie Alpine 68: 13-56.

Busnardo, R.; Charollais, J.; Wiedmann, M.; Clavel, B. 2003. Le Crétacé inférieur de la Veveyse de Châtel (Ultrahelvétique des Préalpes externes; canton de Fribourg, Suisse). Revue de Paléobiologie 22: 1-174.

Cantú-Chapa, A. 1966. Se propone una nueva subdivisión de la Familia Olcostephanidae (Ammonoidea) del Cretácico Inferior (Taraisitinae Subfam. nov. y Taraisites gen. nov.). Ingeniería Petrolera 6: 15-17.

Cantú-Chapa, A. 2001. The Taraises Formation (Berriasian-Valanginian) in northeastern Mexico: Subsurface and outcrop studies. In The western Gulf of Mexico Basin: Tectonics, sedimentary basins, and petroleum systems (Bartolini, C.; Buffler, R.T.; Cantú-Chapa,  A.; editors). American Association of Petroleum Geologists, Memoir 75: 353-370.

Cantú-Chapa, A. 2009. Ammonites of the Cretaceous Taraises and Lower Tamaulipas formations in Eastern Mexico. In Petroleum systems in the southern Gulf of Mexico (Bartolini, C.; Román Ramos, J.R.; editors). American Association of Petroleum Geologists, Memoir 90: 191-216.

Cantú-Chapa, A. 2012. The suture line patterns in Olcostephanidae and Spiticeratidae, ammonites of the Upper Jurassic-Lower Cretaceous: systematic considerations. Paleontología Mexicana 62: 58-72.

Cecca, F. 1985. Alcune ammoniti provenienti della ‘Maiolica’ dell’Appennino Centrale (Umbria, Marche e Sabina). Bollettino del Servizio Geologico d’Italia 103: 133-162.

Collignon, M. 1962. Atlas des fossiles caractéristiques de Madagascar (Ammonites). VIII. Berriasien, Valanginian, Hauterivien, Barremien. Service Géologique de Madagascar: 96 p. Tananarive.

Company, M. 1985. La Subzona de Verrucosum (Valanginianse Superior) en el Barranco de La Querola (SE de España). Mediterranea 4: 113-127.

Company, M. 1987. Los Ammonites del Valanginiense del Sector Oriental de las Cordilleras Béticas (SE de España). Ph.D. Thesis (Unpublished), Universidad de Granada: 294 p.

Cooper, M.R. 1981. Revision of the Late Valanginian Cephalopoda from the Sunday River Formation of South Africa, with special reference to the genus Olcostephanus. Annals of the South African Museum 83: 147-366.

Corvalán, J. 1974. Estratigrafía del Neocomiano marino de la región al sur de Copiapó, Provincia de Atacama. Revista Geológica de Chile 1: 13-36. doi: 10.5027/andgeoV1n1-a02.

Corvalán, J.; Pérez, E. 1958. Fósiles guías chilenos, Titoniano-Neocomiano. Instituto de Investigaciones Geológicas, Chile, Manual 1: 48 p.

Etayo-Serna F. 1985. Paleontología estratigráfica del Sistema Cretácico en la Sierra Nevada del Cocuy. In Proyecto Cretácico (Etayo-Serna, F.; Laverde, F.; editors). Publicaciones Geológicas Especiales del Ingeominas 16 (24): 1-47.

Ettachfini, M. 1991. Le Valanginien de l’Atlas atlantique (Maroc): stratigraphie et ammonitofaune. Strata, série 2, Mémoires 15: 1-177.

Ettachfini, M. 2004. Les ammonites néocomiennes dans l’Atlas atlantique (Maroc). Biostratigraohie, Paléontologie, Paléobiogéographie et Paléoécologie. Strata, série 2, Mémoire 43: 1-223.

Faraoni, P.; Flore, D.; Marini, A.; Pallini, G.; Pezzoni, N. 1997. Valanginian and early Hauterivian ammonite successions in the Mt. Catria group (Central Apennines) and in the Lessini Mts. (Southern Alps), Italy. Paleopelagos 7: 59-100.

Fatmi, A.N. 1977. Neocomian Ammonites from northern areas of Pakistan. Bulletin of the British Museum of Natural History, Geology 28: 255-296.

Felix, J. 1891. Versteinerungen aus der Mexikanischen Jura und Kreide Formation. Paleontographica 37: 140-199.

Fözy , I.; Janssen, N.M.M.; Price, G.D.; Knauer, J.; Pálfy , J. 2010. Integrated isotope and biostratigraphy of a Lower Cretaceous section from the Bakony Mountains (Transdanubian Range, Hungary): A new Tethyan record of the Weissert event. Cretaceous Research 31: 525-545.

Gauthier, H.; Busnardo, R.; Combémorel, R.; Delanoy, G.; Fischer, J.C.; Guérin-Franiatte, S.; Joly, B.; Kennedy, W.J.; Sornay, J.; Tintant, H. 2006. Volume IV. Céphalopodes crétacés. In Révision critique de la Paléontologie Française d’Alcide d’Orbigny (Fischer, J.C.; editor). Backhuys Publishers: 1-292. Leiden.

Gerth, E. 1925. La fauna Neocomiana de la Cordillera Argentina en la parte meridional de la provincia de Mendoza. Actas de la Academia Nacional de Ciencias9: 57-132.

González-Arreola, C.; Barragán, R.; Moreno-Bedmar, J.A. 2014. Olcostephanidae (Ammonoidea) from the upper Valanginian Taraises Formation (Durango State, Mexico). Cretaceous Research 49: 55-62.

Haas, O. 1960. Lower Cretaceous Ammonites from   Colombia, South America. American Museum Novitates 2005: 1-62.

Haug, E. 1910. Période Crétacée. In Traité de géologie (Haug, E, editor), Les Périodes Géologiques 2: 1153-1396. Paris.

Hoffstetter, R.; Fuenzalida, H.; Cecioni, G. 1957. Lexique Stratigraphique International, Amérique Latine, Chile 5: 444 p.

Howlett, P.J. 1986. Olcostephanus (Ammonitina) from the Fossil Bluff Formation, Alexander Island, and its stratigraphical significance. British Antarctic Survey, Bulletin 70: 71-77.

Imlay, R.W. 1938. Ammonites of the Taraises Formation of Northern Mexico. Geological Society of America, Bulletin 49: 539-602.

Imlay, R.W.; Jones, D.L. 1970. Ammonites from the Buchia zones in northwestern California and southwestern Oregon. Geological Survey, Professional Paper 647(B): B1-B59.

Karakasch, N. 1902. Note sur le Crétacé inférieur de Biassala (Crimée). Description de quelques espèces de Céphalopodes du Crétacé inférieur de la Crimée. Annales de l’Université de Grenoble 14: 1-15.

Kemper, E. 1992. Die tiefe Unterkreide im Vechte-Dinkel-Gebiet. Dobler-Druck GmbH & Co KG.: 95 p. Alfeld (Leine).

Kemper, E.; Rawson, P.F.; Thieuloy, J.-P. 1981. Ammonites of Tethyan ancestry in the Early Lower Cretaceous of north west Europe. Palaeontology 24: 251-311.

Kilian, W. 1895. Note stratigraphique sur les environs de Sisteron et contributions à la connaissance des terrains secondaires du Sud-Est de la France. In Réunion extraordinaire dans les Basses-Alpes. Bulletin de la Société Géologique de France, Série 3 (23): 659-803.

Kilian, W. 1902. Sur quelques fossiles remarquables de l’Hauterivien de la région d’Ecragnolles. Bulletin de la Societé Géologique de France 4 (2): 864-867.

Kilian, W. 1910. Erste Abteilung: Unterkreide (Palaeocretacicum), Leiferung 2. Das bathyale Palaeo-cretacicum im südöstlichen Frankreich. In Lethaea geognostica. II. Das Mesozoicum, Band 3 (Kreide) (Frech, F.; editor). Schweizerbart: 169-288. Stuttgart.

Kitchin, F.L. 1908. The invertebrate fauna and paleontological relationships of the Uitenhege Series. Annals of the South African Museum 7: 21-225.

Klein, J. 2005. Lower Cretaceous Ammonites I. Perisphinctaceae 1. In Fossilium Catalogus I: Animalia Pars 139 (Riegraf, W.; editor). Backhuys Publishers: 1-484. Leiden.

Kvantaliani, I.V.; Sakharov, A.S. 1986. Valanginian ammonites from Northern Caucasus. Geologica Balcanica 16: 55-68.

Leanza, A.F. 1944. Las Apófisis yugales de Holcostephanus. Museo de La Plata, Serie Paleontología 9: 13-22.

Leanza, A.F. 1957. Acerca de la existencia de Simbirskites en el Neocomiano argentino. Revista de la Asociación Geológica Argentina 12: 5-17.

Leanza, H.A. 1972. Acantholissonia, nuevo género del Valanginiano de Neuquén, Rep. Argentina y su posición estratigráfica. Revista de la Asociación Geológica Argentina 27: 248-264.

Leanza, H.A. 1981. Faunas de Ammonites del Jurásico superior y del Cretácico inferior de América del Sur, con especial consideración de la Argentina. In Cuencas sedimentarias del Jurásico y Cretácico de América del Sur (Volkheimer, W.; Musacchio, E.; editors), Comité Sudamericano del Jurásico y Cretácico 2: 559-597. Buenos Aires.

Leanza, H.A.; Wiedmann, J. 1980. Ammoniten des Valangin und Hauterive (Unterkreide) von Neuquén und Mendoza, Argentinien. Eclogae Geologicae Helvetiae 73: 941-981.

Lissón, C.I. 1907. Contribución a la Geología de Lima y sus alrededores. Librería é Imprenta Gil: 124 p. Lima.

Lissón, C.I. 1937a. Dos Amonites del Perú. Boletín de Minas 3: 3-9.

Lissón, C.I. 1937b. Género Sphaeroceras Bayle, Sphaeroceras Broggianus nov. sp. Revista de Ciencas 422: 153-155.

Lissón, C.J.; Boit, B. 1942. Edad de los fósiles peruanos y distribución de sus depósitos en la república con un mapa paleontológico del Perú. Sociedad Geológica del Perú, Cuarta edición: 320 p. Lima.

Lukeneder, A. 2004. The Olcostephanus Level: an Upper Valanginian ammonoid mass-occurrence (Lower Cretaceous, Northerrn Calcareous Alps, Austria). Acta Geologica Polonica 54: 23-33.

Lukeneder, A. 2007. Cephalopod evolution: A new perspective-Implications from two Early Cretaceous ammonoid suborders (Northern Calcareous Alps, Upper Austria). Denisia 20: 395-404.

Lukeneder, A.; Harzhauser, M. 2003. Olcostephanus guebhardi as a cryptic habitat for an Early Cretaceous coelobite community (Valanginian, Northern Calcareous Alps, Austria). Cretaceous Research 24: 477-485.

Marschik, R.; Fontboté, L. 2001. The Punta del Cobre Formation, Punta del Cobre-Candelaria area, northern Chile. Journal of South American Earth Sciences 14: 401-433.

Moraga, A. 1977. Geología del Cuadrángulo Quebrada Desierto (Quebrada Salitrosa), III Región, Atacama, Chile. Servicio Nacional de Geología y Minería, Carta Geológica de Chile, Serie Geología Básica 25: 12 p. 1 mapa escala 1:50.000.

Mourgues, F.A. 2002. Biostratigraphy of the marine Chañarcillo Basin (Lower Cretaceous), Atacama region, Northern Chile, and its relationship with the Neuquén-Mendoza Basin (Tithonian-Lower Cretaceous), Argentina. In European Meeting on the Paleontology and Stratigraphy of Latin America, No. 3, Abstract Volume: 73-76. Toulouse.

Mourgues, F.A. 2004. Advances in Ammonite Biostratigraphy of the Marine Atacama Basin (Lower Cretaceous), Northern Chile, and its Relationship with the Neuquén Basin, Argentina. Journal of South American Earth Sciences 17: 3-10.

Mpodozis, C.; Allmendinger, R. 1992. Extensión cretácica a gran escala en el Norte de Chile (Puquios-Sierra de Fraga 27°S): significado para la evolución tectónica de los Andes. Revista Geológica de Chile 19 (2): 167-197. doi: 10.5027/andgeoV19n2-a03.

Munier-Chalmas, E.C.P.A.; de Lapparent, A.A. 1894. Note sur la nomenclature des terrains sédimentaires. Bulletin de la Societé Géologique de France, Série 3, 21 (1893): 438-488.

Neumayr, M. 1875. Die Ammoniten der Kreide und die Systematik der Ammonitiden. Zeitschrift der deutschen Geologischen Gesellschaft 27: 854-942.

Nikolov, T. 1965. A new Valanginian Ammonite (Dobrodgeiceras benavidesi sp. n.) from Perú. Comptes Rendus de l’Académie bulgare des Sciences 18: 1153-1155.

Oppel, A. 1863. Über ostindische Fossilreste aus den sekundären Ablagerungen von Spiti und Gnari Khorsum in Tibet. Palaeontologische Mitteilungen: 267-304.

d’Orbigny, A. 1840-1842. Paléontologie française. Terrains crétacés. I. Céphalopodes: 662 p. Paris.

d’Orbigny, A. 1842. Coquilles et échinodermes fossiles de Colombie (Nouvelle-Grenade), recueillis de 1821 à 1833 par M. Boussaingault. Masson Editors: 64 p. Paris.

d’Orbigny, A. 1850. Prodome de Paléontologie stratigraphique universelle des animaux mollusques et rayonnés. Masson: 289 p. Paris.

Paquier, V. 1900. Recherches géologiques dans le Diois et les Baronnies orientales. Annales de l’Université de Grenoble 12: 1-806.

Pavlow, A. 1892. Ammonites de Speeton et leurs rapports avec les ammonites des autres pays. In Argiles de Speeton et leurs équivalents (Pavlow, A.; Lamplugh, G.W.; editors). Bulletin de la Société Impériale des Naturalistes de Moscou, Nouvelle Série 5 (1891): 97-155.

Pictet, F.J.; Campiche, G. 1858-1860. Description des fossiles du terrain crétacé des environs de Sainte-Croix. 1ére Partie. Matériels pour la Paléontologie Suisse 2 (2): 1-380.

Rawson, P.F. 1999. Long-distance correlations in the Valanginian-Hauterivian: Argentina-Western Mediterranean-NW Europe. Scripta Geologica, Special Issue 3: 151-158.

Rawson, P.F. 2007. Global relationships of Argentine (Neuquén Basin) Early Cretaceous ammonite faunas. Geological Journal 42: 175-183.

Rawson, P.F.; Kemper, E. 1978. Varlheideites, n. gen. (Ammonoidea, Neocomitinae) aus dem Obervalangin Nordwest-Deutschlands. Geologisches Jahrbuch, Reihe A 45: 163-181.

Reboulet, S.; Rawson, P.F.; Moreno-Bedmar, J.A.; Aguirre-Urreta, M.B.; Barragán, R.; Bogomolov, Y.; Company, M.; González-Arreola, C.; Idakieva Stoyanova, V.; Lukeneder, A.; Matrion, B.; Mitta, V.; Randrianaly, H.; Vašiček, Z.; Baraboshkin, E.J.; Bert, D.; Bersac, S.; Bogdanova, T.N.; Bulot, L.G.; Latil, J.-L.; Mikhailova, I.A.; Ropolo, P.; Szives, O. 2011. Report on the 4th international meeting of the IUGS lower cretaceous ammonite working group, the ‘Kilian Group’. Cretaceous Research 32: 786-793. Dijon.

Riccardi, A.C. 1988. The Cretaceous System of Southern South America. Geological Society of America, Memoir 68: 1-161.

Riccardi, A.J.; Westermann, G.E.G. 1970. The Valanginian Dobrodgeiceras Nikolov (Ammonitina) from Perú Journal of Paleontology 44: 888-892.

Riccardi, A.C.; Westermann, G.E.G.; Levy, R. 1971. The Lower Cretaceous Ammonitinae Olcostephanus, Leopoldia and Favrella from west-central Argentina. Palaeontographica A 136: 83-121.

Riedel, L. 1938. Amonitas del cretácico inferior de la Cordillera Oriental. Estudios geológicos y paleontológicos sobre la Cordillera Oriental de Colombia, Parte 2. Departamento de Minas y Petróleo, Ministerio de Industrias y Trabajo: 7-78. Colombia.

Robert, E.; Bulot, L.G.; Dhondt, A.; Jaillard, E.; Villa-gómez, R.; Rivadeneira, M.; Paz, M. 1998. La transgresión del Cretáceo inferior en el margen andino (Perú y Ecuador): datos preliminares. Boletín de la Sociedad Geológica del Perú 88: 73-86.

Roch, E. 1930. Etudes géologiques dans la région méridionale du Maroc occidental. Notes et Mémoires du Service de la Carte Géologique du Maroc 9: 1-542.

Rodighiero, A. 1919. Il sistema Cretaceo del Veneto Occidentale compreso fra l’Adige e il Piave con speciale riguardo al Neocomiano dei Sette Comuni. Paleontographia Italica 25: 39-125.

Roman, F. 1933. Sur quelques formes de Céphalopodes de l’Yonne et des régions voisines. Travaux du Laboratoire de Géologie de la Faculté des Sciences de Lyon 22: 6-22.

Sayn, G. 1889. Notes sur quelques Ammonites nouvelles ou peu connues du Néocomien inférieur. Bulletin Société Géologique de France 17: 679-688.

Sayn, G. 1907. Les ammonites pyriteuses des marnes valanginiennes du sud-est de la France. Mémoire de la Société Géologique de France, Paléontologie 23: 29-68.

Segerstrom, K.; Parker, R.L. 1959. Cuadrángulo Cerrillos, Provincia de Atacama. Instituto de Investigaciones Geológicas, Carta Geológica de Chile 1: 33 p.

Segerstrom, K.; Ruiz, C. 1962. Cuadrángulo Copiapó, Provincia de Atacama. Instituto de Investigaciones Geológicas, Carta Geológica de Chile 3: 115 p.

Segerstrom, K.; Thomas, H.; Tilling, R.I. 1963. Cuadrángulo Pintadas, Provincia de Atacama. Instituto de Investigaciones Geológicas, Carta Geológica de Chile 12: 53 p.

Sepúlveda, P.; Naranjo, J.A. 1982. Hoja Carrera Pinto, Región de Atacama. Servicio Nacional de Geología y Minería, Carta Geológica de Chile, Segie Geología Básica 53: 60 p. 1 mapa escala 1:100.000.

SERNAGEOMIN. 2002. Mapa Geológico de Chile en 3 hojas. Servicio Nacional de Geología y Minería, Chile. Carta Geológica de Chile, Serie Geología Básica 75, 1 mapa 1:1.000.000.

Sharpe, D. 1856. Description of Fossils from the Secondary Rocks of Sundays River and Zwartkop River, South Africa, collected by Dr. Atherstone and A.G. Bain, Esq. Geological Society of London, Transactions 7: 193-203.

Spath, L.F. 1923a. On the ammonite horizons of the Gault and contiguous deposits. Summary of Progress of the Geological Survey of Great Britain and the Museum of Practical Geologyfor 1922. Memoirs of the Geological Survey of Great Britain 1923: 139-149.

Spath, L.F. 1923b. Ammonoidea of the Gault. Part I. Paleontographical Society 75 (1921): 1-72. London.

Spath, L.F. 1924. On the ammonites of the Speeton Clay and the subdivisions of the Neocomian. Geological Magazine 61: 73-89.

Spath, L.F. 1930. On the Cephalopoda of the Uitenhage Beds. Annals of the South African Museum 28: 131-157.

Spath, L.F. 1939. The Cephalopoda of the Neocomian Belemnite Beds of the Salt Range. Geological Survey of India, Memoires, Palaeontologica Indica, N. S. 25: 1-154.

Steinmann, G. 1890. Cephalopoda. In Elemente der Plaläontologie (Steinmann, G.; Döderlein, L.; editors). W. Engelmann: 344-475. Leipzig.

Steinmann, G. 1930. Geología del Perú. 458 p. Heidelberg.

Stolley, E. 1937. Die Gliederung des norddeutschen marinen Unterneokoms. Zentralblatt für Mineralogie Geologie und Paläontologie B: 434-456, 497-506.

Tavera, J. 1956. Fauna del Cretáceo Inferior de Copiapó. Facultad de Ciencias Físicas y Matemáticas, Universidad de Chile, Anales 13: 205-216.

Thieuloy, J.-P. 1965. Un Céphalopode remarquable de l’Hauterivien basal de la Drôme. Himantoceras nov. gen. Bulletin de la Societé géologique de France, Série 7 (6): 205-213.

Thieuloy, J.-P. 1977. Les ammonites boréales des formations néocomiennes du Sud-Est Francais (Province Subméditerranéenne). Géobios 10: 395-461.

Uhlig, V. 1903. Himalayan fossils. The fauna of the Spiti Shales. Geological Survey of India, Memoires, Palaeontologica Indica 15: 1-511.

Vašiček, Z. 2010. Early Cretaceous ammonites from the Butkov Quarry (Manín Unit, Central Western Carpathians, Slovakia). Acta Geologica Polonica, 60: 393-415.

Weaver, C. 1931. Paleontology of the Jurassic and Cretaceous of Western Central Argentina. University of Washington, Memoir 1: 1-469.

Wegner, R. N. 1909. Übersicht über die bisher bekannten Astieria-Formen der Ammonitenngattung Holcostephanus nebst Beschreibung zweier neuer Arten. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie 1: 77-92.

Westermann, G.E.G. 2000. Marine faunal realms of the Mesozoic: review and revision under the new guidelines for biogeographic classification and nomenclature. Palaeogeography, Palaeoclimatology, Palaeoecology 163: 49-68.

Wiedmann, J.; Dieni, I. 1968. Die Kreide Sardiniens und Ihre Cephalopoden. Palaeontographica Italica 64: 1-171.

Windhausen, A. 1931. Geología Argentina 2: 645 p. Buenos Aires.

Wippich, M.G.E. 2001. Die tiefe Unter-Kreide (Berrias bis Unter-Hauterive) im Südwestmarokkanischen Becken: Ammonitenfauna, Bio- und Sequenzstratigraphie. Ph.D. Thesis (Unpublished). University Bochum: 142 p.

Wippich, M.G.E. 2003. Valanginian (Early Cretaceous) ammonite faunas from the western High Atlas, Morocco, and the recognition of western Mediterranean ‘standard’ zones. Cretaceous Research 24: 357-374.