Caribbean Lower Tithonian Ammonites from Central-East

Geobios 42 (2009) 117–132

Original article Caribbean Lower Tithonian ammonites from central-east Mexico§ Ammonites des Caraı¨bes dans le Tithonien infe´rieur du centre-est du Mexique Ana Bertha Villaseñor a, Federico Olóriz b,* a Departamento de Paleontología, Universidad Nacional Autónoma de México, Instituto de Geología, Ciudad Universitaria, 04510 México, D.F., Mexico b Departamento de Estratigrafía y Paleontología, Facultad de Ciencias, Universidad de Granada, Av. Fuentenueva s/n, 18002 Granada, Spain Received 6 March 2008; accepted 20 June 2008 Available online 25 November 2008

Abstract Imlay’s ammonite species Paradontoceras butti and Paradontoceras antilleanum are ﬁrst reported from central-east Mexico under precise stratigraphic control. Mexican and Cuban ammonites referred to these species have been studied and proved to have been gathered from a relatively limited area within the southern paleomargin of the north American Plate, which embraces eastern Mexico and north-western Cuba. The palaeontological examination, the multivariate analysis conducted on Mexican and Cuban data, the precise biostratigraphic and biogeographic control support the reinterpretation of Imlay’s species P. antilleanum and P. butti. The latter has been designated the single, valid nominal species- level taxon which comprises the two species proposed by Imlay. The new genus Housaites has been created since the taxon Butticeras Housˇa is not available for taxonomic nomenclature. Housaites butti (Imlay) is upper Lower Tithonian (Semiforme/Verruciferum Zone of the Tethyan standard) in Mexico. The stratigraphic range in Cuba is not conclusively known. # 2008 Elsevier Masson SAS. All rights reserved.

Résumé Les espèces d’ammonites décrites par Imlay sous les noms de Paradontoceras butti et Paradontoceras antilleanum sont décrites pour la première fois du centre-est du Mexique, dans un contexte stratigraphique bien contrôlé. Les ammonites mexicaines et cubaines rapportées à cette espèce sont ici étudiées et regroupées ; elles proviennent toutes d’une zone relativement restreinte correspondant à la limite sud de la plaque nord- américaine, laquelle comprend l’est du Mexique et le nord-ouest de Cuba. L’étude paléontologique et l’analyse multivariée, biostratigraphi- quement et biogéographiquement contrôlées, menées sur les spécimens mexicains et cubains, soutiennent la réinterprétation des deux espèces. L’espèce butti est ici désignée comme seul taxon spéciﬁque valide, réunissant les deux espèces initialement décrites par Imlay. Le nouveau genre Housaites est créé, le nom Butticeras Housˇa étant non disponible. Housaites butti (Imlay) caractérise la partie supérieure du Tithonien inférieur au Mexique (Zone à Semiforme/Verruciferum de l’échelle téthysienne standard). L’intervalle stratigraphique couvert à Cuba n’est pas connu de façon concluante. # 2008 Elsevier Masson SAS. Tous droits réservés.

Keywords: Ammonites; Housaites; Upper Jurassic; Lower Tithonian; Mexico; Caribbean

Mots clés : Ammonites ; Housaites ; Jurassique supérieur ; Tithonien inférieur ; Mexique ; Caraïbes

1. Introduction

Imlay (1942: pp. 1454–1456) described two new ammonites § Corresponding editor: Pierre Hantzpergue. species, Paradontoceras butti n. sp. Imlay, and P.antilleanum n. * Corresponding author. sp. Imlay, from north-western and north-central Cuba (Pinar del E-mail address: [email protected] (F. Olóriz). Río, and Pinar del Río and Santa Clara provinces, respectively).

This author studied 19 specimens of these taxa, collected by himalayitins in Loma Rinconada section (Sierra de Cruillas, employers of the Atlantic Refining Company, most probably Tamaulipas, Mexico): Lytohoplites carribeanus Imlay together R.H. Palmer and R.E. Dickerson, and interpreted their age as with Corongoceras filicostatum Imlay, in horizons containing being Late Portlandian (Imlay, 1942: p. 1434), which preceded Kossmatia victoris Burckhardt, as well as Durangites, his Tithonian deposits without ammonites (Imlay, 1942: Table Paradontoceras?, and Simoceras? from horizons yielding 4). Later on references to these Cuban taxa were made by Physodoceras and Metahaploceras? Judoley and Furrazola (1968), Housˇa and De la Nuez (1973, Myczyn´ski (1989) gave a synthetic, valuable revision of the 1975), Imlay (1980), Myczyn´ski (1989, 1994, 1996), Myc- Tithonian in Cuba. This author identified Imlay’s species zyn´ski and Psczółskowski (1994), Fernández-Carmona (1995), P. antilleanum and P. butti from the uppermost Lower Tithonian Schweigert and Zeiss (1998), and Pszczółkowski and Myc- and throughout the Upper Tithonian in Sierra de los Órganos zyn´ski (2003). With the exception of Schweigert and Zeiss and Sierra del Rosario in western Cuba (Myczyn´ski, 1989: Figs. (1998), these authors discussed the stratigraphic location and 14, 15). taxonomy of Cuban ammonites first described by Imlay (1942), From Sierra de los Órganos, this author reported the but there was no improvement of the precise, morphological occurrence of Paradontoceras butti Imlay associated to description made by Imlay (1942). In contrast, biostratigraphic Hildoglochiceras (Salinites) cf. gallardoi (Judoley and interpretation varied between references to horizons that belong Furrazola) and H. (S.) grossicostatum Imlay from the Tithonian to both the Lower and Upper Tithonian, less frequently to a El Americano Member? in Sierra de Cabezas. It is worth Berriasian age. mentioning that Salinites was considered a mainly Upper Judoley and Furrazola (1968: pp. 118–119) interpreted a Tithonian ammonite by Myczyn´ski (1989, 1990). In addition, Middle Tithonian age for Paradontoceras antilleanum Imlay and Myczyn´ski (1989) cited Paradontoceras antilleanum Imlay and P. butti Imlay in the Artemisa Fm. (Cuba), associated to P.butti Imlay from the El Americano Member in sections B-HA Haploceras sp., Pseudoanahamulina sp. (Pseudoanahamulina (Tithonian horizon above Lytohoplites carribeanus Imlay) and rosariensis in pp. 54–55), Pseudolissoceras cf. P. zitteli C-HA (Tithonian horizon above Lytohoplites sp. together (Burckhardt), and Virgatosphinctes spp. with Aulacosphinctoides sp., Pseudoinvoluticeras sp. cf. Housˇa and De la Nuez (1973: p. 18) were pioneers in P. mozambicum Collignon, ‘‘Virgatosphinctes’’ pinarensis questioning the appertaining of Imlay’s species P. antilleanum Judoley and Furrazola, and others). and P. bu tt i to genus Paradontoceras, and proposed the new In Sierra del Rosario, records of Paradontoceras butti Imlay genus Butticeras instead, but without clear reference to precise were cited by Myczyn´ski (1989) from La Catalina section stratigraphy since they only mentioned the occurrence of a new [Tithonian horizon also yielding Vinalesites rosariensis (Imlay) perisphinctid (assumedly Butticeras) with simple ribs in the below poorly preserved specimens of Dickersonia]. At the upper Lower Tithonian from western Cuba. In the English Cinco Pesos section, P. antilleanum Imlay was reported from a version of the same abstract, Housˇa and De la Nuez (1975: p. 57) Tithonian horizon above Virgatosphinctes sp. and below mentioned ‘‘a new genus of the family Perisphinctidae with Vinalesites rosariensis (Imlay), as well as from horizons simple ribs’’ (literal translation) from their horizon above the between those yielding Neochetoceras below and Lytohoplites middle part of the Lower Tithonian in western Cuba and below an above (Myczyn´ski, 1989: Fig. 11). Upper Tithonian horizon characterized by Kossmatia, Blanfor- Myczyn´ski (1989) identified Paradontoceras antilleanum diceras, Raymondiceras?, Riasanites?, Rapidoceras n. g. (‘‘Hap- Imlay and P. butti Imlay as endemic ammonites from the loceras’’ Judoley and Furrazola, 1968) and others. However, Caribbean province (i.e., Mexico, Cuba and southern USA), these authors cited Butticeras n. g. (genotype Paradontoceras and even he foresaw their occurrence in southern USA. This butti Imlay, 1942), together with Vinalesites, Leptoceras, etc., of author followed Imlay (1980) when interpreting the inclusion the probable Upper Hauterivian. of species butti and antilleanum in Paradontoceras, even In Mexico, Verma and Westermann (1973) interpreted recognizing the doubts cast by Verma and Westermann (1973) Substeueroceras (syn. Paradontoceras?) among ammonites on the validity of this genus and the new taxon Butticeras collected from S Catorce (San Luis Potosí), but they did not created by Housˇa and De la Nuez (1973, 1975). mention Cuban records. Imlay (1980: p. 35) mentioned Myczyn´ski and Psczółskowski (1994) provided data Paradontoceras cf. P. butti Imlay gathered from the Pimienta from Sierra del Rosario. In the El Toro section Fm. in PEMEX sample 20851, located 3 km south from Jonotla ‘‘Paradontoceras’’ antilleanum Imlay and ‘‘P.’’ butti Imlay (Puebla, Mexico), and interpreted an earliest Late Tithonian age were gathered, together with Lytohoplites sp., Protancyloceras for this ammonite. This citation has been for a long time the hondense (Imlay), Pseudolissoceras cf. zitteli (Burckhardt), single known record of species P. butti from Mexico. When and Vinalesites rosariensis (Imlay), among others, below discussing on his Kossmatia-Durangites Ammonite assem- Lower Tithonian horizons with ‘‘P.’’ antilleanum Imlay and blage, Imlay (1980: pp. 34–35) regarded the same age for the Vinalesites rosariensis (Imlay). Presence of ‘‘Paradontoceras’’ ammonite assemblage composed of Burckhardt’ species antilleanum Imlay and ‘‘P.’’ butti Imlay in the Cinco Pesos Pseudolissoceras zitteli, Kossmatia victoris and Grayiceras? section in Lower Tithonian horizons with Protancyloceras mexicanum, among others, reported from Mazatepec (Puebla) hondense (Imlay) was also mentioned, whereas in La Villa–La by Cantú-Chapa (1967: p. 21). In addition, Imlay (1980: p. 35) Revuelta section, they are thought to belong to the uppermost cited ex-situ specimens within the stratigraphic range of Lower Tithonian (Myczyn´ski and Psczółskowski, 1994: Fig. 6). A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 119

From La Catalina section ‘‘Paradontoceras’’ butti Imlay, movements (Salvador, 1987: p. 445), show SW-NE folding together with Hildoglochiceras (Salinites) sp. and Vinalesites resulting in NNW–SSE anticlines and synclines interrupted rosariensis Imlay were reported in horizons above ammonites southward by rocks of the Transmexican Volcanic Axis (eje interpreted as belonging to the late Early Tithonian. neovolcánico in López Ramos, 1979a: 291). Nowadays, near ‘‘Paradontoceras’’ antilleanum Imlay and ‘‘P.’’ butti Imlay Mazatepec, dense forest coverage makes it difficult to access were registered from La Piedra de Genaro section, in Lower- Upper Jurassic outcrops off the Apulco River banks. In the area, Upper Tithonian horizons and the lower part of the Upper the Upper Jurassic section shows a regional deepening of 108 to Tithonian, respectively (Myczyn´ski and Psczółskowski, 1994: 308 NE, and is composed of coarse bedded lower and thinner Fig. 4). For these species of ‘‘Paradontoceras’’, Myczyn´ski and upper parts, the latter being locally affected by faulting that Psczółskowski (1994: Fig. 8) concluded a biostratigraphic disturbs stratigraphical continuity. Upper Jurassic carbonates, distribution ranging from the Lower Tithonian Pseudolisso- from mid-shelf to lower-ramp deposits of the San Andres Fm. ceras spp. to the Upper Tithonian Himalayites and Corongo- (Hernández De La Fuente, 1996) underlie mainly upper Lower ceras zones (ammonite assemblages). Kimmeridgian and Tithonian, brownish to black siltstones of Fernández-Carmona (1995) referred to genus Butticeras in the Tamán Fm. (see Cantú-Chapa, 1971 for regional the Artemisa Fm. from western Cuba with no mention of biostratigraphy and correlation). Parallel rather than low-angle, specimens at the species level, but he did not provide fine cross lamination is common in the latter, where silty information about precise biostratigraphical horizons within interbeds are typical together with occasional intercalation of the Tithonian. more calcareous horizons. The lower Tithonian succession Myczyn´ski (1996) clearly stated the morphological differ- shows local evidence of synsedimentary sliding (slumps) and ence between Butticeras and Paradontoceras in the typical includes horizons containing more calcareous concretions. At occurrence of simple ribs in the former, and accepted the present, the Tithonian, and especially the Lower to lower Upper taxonomic status proposed by Housˇa and De la Nuez (1973, Tithonian succession at the Apulco River section is widely 1975) as a valid one. In contrast, Myczyn´ski (1996) considered covered by alluvium related to severe flooding during 1999, and invalid the interpretation of Lytohoplites carribeanus Imlay, therefore outcrop limitations are common in the area. Over- which is associated to Paradontoceras antilleanum and P. butti lying the Tamán Fm., cherty horizons intercalated in grayish to in the Lower Tithonian part of their ranges, and reclassified it as darkish, more calcareous clayey limestones indicate the Paralytohoplites gen. nov. with carribeanus Imlay as the type Pimienta Fm. (López Ramos, 1979b: p. 315; Fig. 1(B) for species. Formation-level stratigraphy from the east-central Mexico According to the above, the taxonomic interpretation of series). Imlay’s species P. antilleanum and P. butti was changing from Paradontoceras to ‘‘Paradontoceras’’ and then Butticeras, 3. Material and methods whereas their biostratigraphic ranges have been most com- monly interpreted as upper Lower Tithonian, although In our revision of classical Upper Jurassic sections of references to the Upper Tithonian and even Berriasian have Mexico, field missions during the past seven years provided also been admitted (see below). new data from eastern Sierra Madre in the Huasteca region. The present research reports the new finding of Imlay’s Among the outcrops visited, Tithonian deposits in the Apulco species from central-east Mexico, in the Apulco River section River section near Mazatepec (Puebla) were studied by Cantú- near Mazatepec (Puebla), where they were registered together Chapa (1967, 1971) and Hernández De La Fuente (1996), while with Tethyan genera Simocosmoceras and Simoceras, two taxa Tithonian deposits in the Mazatepec area were revisited by previously reported by Villaseñor et al. (2000, 2003) and Stinnesbeck et al. (1993). In the Apulco River section, new Villaseñor and Olóriz (2001). The taxonomic interpretation of insights into the occurrence of Tethyan influences were species antilleanum and butti is discussed and the new genus reported by Villaseñor et al. (2000, 2003) and Villaseñor and Housaites (Lithacoceratinae) is proposed to include them. Olóriz (2001). Recent research in the Apulco River Valley by Nowadays, the area of occurrence of Housaites is limited to the authors verified the occurrence of Imlay’s species P. butti central-eastern Mexico and north-western and north-central and P. antilleanum associated to Lower Tithonian ammonites in Cuba. the Apulco River section MT-2, which is 12.5 m thick and composed of brownish and more or less calcareous siltstones 2. Geological setting and silty limestones. The ammonites studied were collected from calcareous siltstones of the Tamán Formation, on the right The Apulco River section MT-2 studied in the Apulco bank of the Apulco River, close to the hydroelectric station Valley, near Mazatepec (Puebla), is located in the Sierra Norte (Fig. 1(C, D)). The Tithonian stratigraphic interval identified de Puebla, which belongs to the geologic subprovince of for Imlay’s species P. butti and P. antilleanum is 1.8 m thick Eastern Sierra Madre (López Ramos, 1979a: p. 291, Figs. 6-1, within the lowermost part of the MT-2 section. Fossiliferous 6-8; grey colours in Fig. 1(A)) and to the Mexican Belt of horizons in siltstones show fossil preservation as imprints and/ Thrusts and Folds (Suter, 1990), which is considered as a or leaf preservation. The most abundant macrofossils are geologic province by Ortega-Gutiérrez et al. (1992; punctuated ammonite remains (1175 specimens and fragments of line in Fig. 1(A)). Regional structural trends, due to Laramid haploceratids, perisphinctids, and rare simoceratids and 120 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132

Fig. 1. Regional geology, location, and studied section. A. Geological North-Central Provinces of Mexico according to López-Ramos (1979), with the ‘‘Miogeoclinal del Golfo de Mexico’’ province (19), the ‘‘Cinturón Mexicano de Pliegues y Fallas’’ province (20), and the ‘‘Plataforma Valle San Luis’’ province (23) as interpreted A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 121

Fig. 2. Stratigraphic range for large and small types of the studied ammonites at the Apulco River section MT-2 (black circles for large specimens, empty circles for small specimens), and pie diagram showing ammonites versus bivalves throughout the same stratigraphic interval. Détail des distributions stratigraphiques des spécimens de grande et de petite taille collectés dans la section Apulco MT-2 (cercles noirs pour les grands spécimens, cercles blancs pour les petits spécimens) et diagramme en camembert représentant les pourcentages des ammonites versus bivalves pour le même intervalle stratigraphique.

Simocosmoceras); bivalves are also common (670; study in in Mexican versus Cuban specimens, and then within the new progress). genus Housaites. The 66 specimens described were gathered from four beds, 15 In precise references to the specimens described, labeling to 20 cm thick each (Fig. 2), and show phenotypes corresponding indicates IGM (Institute of Geology of the UNAM, Mexico), to Imlay’s species P. bu tt i and P. antilleanum. Collected in situ, followed by catalogue-number indicating specimen’s identiﬁ- with precise stratigraphic control, they represent 5.6% of the total cation within the National Paleontological Collection housed in number of ammonite remains sampled in these beds (64% in the Institute of Geology (UNAM). When more than one Fig. 2). These 66 ammonites – preserved as imprints – provide no specimen is included within the same hand-sample, a last information related to shell volume. Shell size of 48 specimens number is added which identiﬁes the precise individual. was approached showing a size range between 12.1 and 70 mm, In description of shell morphology, the following abbrevia- and the mean shell size close to 26.5 mm. Two ammonite shell- tions are used: DM, shell diameter; U, width of the umbilicus; size classes have been distinguished: H, whorl height; ER2, number of external ribs per half a whorl; UR2, number of umbilical ribs per half a whorl. less than 30 mm (35 specimens); more than 30 mm (13 specimens). 4. Systematic palaeontology

Body chamber preservation must be interpreted from Order AMMONOIDEA Zittel, 1884 sculpture since no suture lines are preserved. Given the scarce Suborder AMMONITINA Hyatt, 1889 illustration published of Imlay’s species P. butti and Superfamily PERISPHINCTOIDEA Steinman in Stein- P. antilleanum (Imlay, 1942: Pl. 7, Figs. 10–12, Pl. 8, Figs. mann and Doderlein, 1890 4–9; Judoley and Furrazola, 1968: Pl. 80, Figs. 1–4; Myczyn´ski Family ATAXIOCERATIDAE Buckman, 1921 and Psczółskowski, 1994: Pl. 1, Fig. 1a), we performed a Subfamily LITHACOCERATINAE Zeiss, 1968 comparative analysis with inclusion of the 13 individuals Genus Housaites nov. gen illustrated from the Pinar del Rio province in western Cuba by Derivatio nominis: The name refers to our recently the authors mentioned (four specimens interpreted as P. butti, deceased colleague Václav Housˇa, who was the first and nine specimens interpreted as P. antilleanum). On this ammonitologist proposing separation of Imlay’s species basis, we evaluated the interpretation of phenotype variability P. antilleanum and P. butti from genus Paradontoceras. by Ortega-Gutiérrez et al. (1992). B. Regional stratigraphy (modified from Morán-Zenteno, 1994). C. Location. D. Studied section showing the stratigraphical range of Housaites. Géologie régionale, localisation géographique et section étudiée. A. Provinces géologiques du centre-nord du Mexique d’après López-Ramos (1979), avec la province du « Miogeoclinal del Golfo de Mexico » (19), la province du « Cinturón Mexicano de Pliegues y Fallas » (20) et la province de « Plataforma Valle San Luis » (23) d’après Ortega-Gutiérrez et al. (1992). B. Stratigraphie régionale (modifié de Morán-Zenteno, 1994). C. Localisation géographique de la zone étudiée. D. Section étudiée montrant la distribution stratigraphique de Housaites. 122 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132

Fig. 3. 1–10. Housaites butti (Imlay). 1, IGM 6146-1; 2, IGM 6149; 3, IGM 6149-4; 4, IGM 6150 (from bed 21); 5, IGM 6141-4; 6, IGM 6153; 7, IGM 6157; 8, IGM 6159-2; 9, IGM 6171; 10, IGM 6085-1 (from bed 21b). Apulco River section MT-2. Scale = 1 cm. 1–10. Housaites butti (Imlay). 1–4 (horizon 21); 5–10 (horizon 21b) Section Apulco MT-2. Échelle = 1cm.

Type species: Paradontoceras butti Imlay, 1942, from the Remarks: Specimens included in Housaites have been Viñales limestone, western Cuba, Pinar del Rio province. classically interpreted as Paradontoceras (Imlay, 1942, 1980; Diagnosis: Housaites includes moderately coiled ammo- Judoley and Furrazola, 1968; Myczyn´ski, 1989) and then nites, with subovate whorl section, flattened flanks, and less ‘‘Paradontoceras’’ (Myczyn´ski and Psczółskowski, 1994). than 130 mm in shell size. Dense ribbing in preadults shows However, Housˇa and De la Nuez (1973, 1975) proposed the bifurcations below the middle flank. Variably crowded, simple new genus Butticeras for them, which was used by Fernández- ribs are typical of adult stages. Throughout ontogeny, rib- Carmona (1995) when he referred to significant Tithonian division points rise to close the mid-flank and ribs cross the ammonites from the Guaniguanico Mountains in western Cuba. venter without modification. No constrictions. Peristomal This proposal was accepted by Myczyn´ski (1996) who made structures and suture lines unknown. remarks on a more or less similar ribbing in other Late Jurassic Holotype: U.S. National Museum 103417. ammonites such as Acuticostites Semenov, 1898, Pavlovia A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 123

Fig. 4. 1–13. Housaites butti (Imlay). 1, IGM 6147-1; 2, IGM 6149-2; 3, IGM 6152; 4, IGM 6151-1; 5, IGM 6175 (from bed 21); 6, IGM 6116-1 (from bed 21b); 7, IGM 6154-2; 8, IGM 6154-3; 9, IGM 6154-4; 10, IGM 6161; 11, IGM 6164; 12, IGM 6169; 13, IGM 6172 (from bed 22b). Apulco River section (MT-2). Scale = 1 cm. 1–13. Housaites butti (Imlay). 1–5 (horizon 21); 6 (horizon 21b); 7–13 (horizon 22b). Section Apulco MT-2. Échelle = 1cm.

Ilovaiski, 1917, Anavirgatites Spath, 1925, and Lytohoplites Butticeras was earlier occupied by Anderson (1958: p. 272) Spath, 1925. Myczyn´ski (1999), Pszczółkowski and Myczyn´ski to identify a small group of mortoniceratid ammonites (2003) and López-Caballero (2006) used Butticeras butti and (Campanian). According to Wright et al. (1996: p. 197), the B. antilleanum. taxon created by Anderson (1958) is considered a junior However, the use of the genus level taxon-name Butticeras synonymous of Submortoniceras Spath, 1926. According to all implies some problems according to the Code. Butticeras the aforementioned, Butticeras Housˇa, in Housˇa and De la Nuez HousˇainHousˇa and De la Nuez, 1973 is an invalid name due to (1973, 1975), is not available for taxonomic nomenclature the fact that it was unavailable when proposed (ICZN, 2000 because of inadequate description and publication. Hence the arts. 11; 13.1.1; 9; 8). Hence Housˇa’s Butticeras (Housˇain new name Housaites is proposed here. Housˇa and De la Nuez, 1973, 1975) is considered nomen nudum Biostratigraphical range: The precise range of genus and therefore it is not available for taxonomic nomenclature Housaites seems to be inconclusively known. The new (ICZN art. 13). Moreover, the genus level taxon-name described specimens of Housaites span a short stratigraphical 124 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 125 interval within the mid Lower Tithonian (two-fold division) in shows a rather unclear low bifurcation at 38.8 mm diameter. Rib Mexico. The stratigraphical range in Cuba seems to be larger, interspaces are comparatively wide on the last whorl preserved. including the lower Upper Tithonian, according to Myczyn´ski Although postjuvenile, mature stages are assumed to be related to and Psczółskowski (1994: Fig. 8), but it could be more similar the development of simple and more widely spaced ribs, the data to that registered in Mexico on the basis of synthetic data on body chamber length, suture line and peristomal structures are provided by Myczyn´ski (1994: p. 98; 1999, pp. 106–107). not precisely known. Reference to potential records within the Berriasian was given by Pszczółkowski and Myczyn´ski (2003), who ‘‘strongly 5. Multivariate analysis suggest’’ (literal translation) that Butticeras butti disappeared early in Berriasian times. Housaites has been reported from Since typical specimens previously thought to belong to western and north-central Cuba and central-east Mexico. P. antilleanum show the combination of smaller shells and persistent bifurcate ribbing throughout their preserved shells, Housaites butti (Imlay, 1942) while simple rib development in greater-size H. butti specimens Figs. 3(1–10), 4(1–13) starts between 25 and 34 mm, morphospecies separation could 1942. Paradontoceras butti - Imlay, p. 1454, Pl. 7 Fig. 10, be artificial, due to rather indistinguishable inner whorls (see 11, 12. Fig. 5(A, B)). 1942. Paradontoceras antilleanum - Imlay, p. 1455, Pl. 8, Thus, we conducted a statistical analysis on a combined Figs. 4, 5–9. database including previous reports of antilleanum and butti 1968. Paradontoceras antilleanum - Judoley and Furrazola, measurements (Judoley an Furrazola, 1968: p. 118, 119), Pl. 80, Figs. 1–4. complemented with measurements we obtained from both 1994. ‘‘Paradontoceras’’ butti - Myczyn´ski and illustrations (from Imlay, 1942: Pl. 7, Fig. 11; Pl. 8, Fig. 4; Psczółskowski, Pl. 1, Fig. 1a. Judoley and Furrazola, 1968: Pl. 80, Fig. 1; Myczyn´ski and ? 1999. Butticeras butti - Myczyn´ski, p. 106, Fig. 7[1]. Psczółskowski, 1994: Pl. 1, Fig. 1a) and the new collected ? 1999. Butticeras antilleanum - Myczyn´ski, p. 106, Fig. 7[2]. material (Appendix A). The null hypothesis we considered was that only one paleobiospecies could comprise the morphologic Material: 66 specimens from siltstones within the Tamán variability observed on the basis of quantitative and qualitative Formation at the Apulco River section MT-2, Apulco River characters. In such a context, and taking into account how Valley, Mazatepec, Puebla, Mexico (Figs. 1 and 2). Besides, 13 limited observation is, due to preservation, smaller individuals Cuban specimens were studied and labeled USNM, J&F, and (i.e., typical antilleanum phenotypes) could be preadults, M&P (plus catalogue number) corresponding to Imlay (1942), incomplete remains corresponding to inner whorls, and/or Judoley and Furrazola (1968), and Myczyn´ski and microconchiates. However, stratigraphic separation seems to be Psczółskowski (1994) collections, respectively. in beds 22b and 23 (Fig. 2) where only small specimens were Diagnosis: As stated for genus Housaites, with adult stage collected–12 specimens which represent 18% of the new showing less dense, simple ribbing. collected specimens, 10 of which were excluded from the Measurements: see Appendix A. analysis because of the lack of data on ribbing (see below). Description: Moderately evolute to evolute shells (U/D For multivariate analysis we used PAST v. 1.72 (Hammer between 0.29 and 0.50) that show a trend for decreasing et al., 2001) with log-transformation of data due to both their umbilical width during ontogeny. Shell diameter varies from 12 belonging to different units and non-normal distribution. to 29.9 mm in small specimens and from 30 to 70 mm in large Cluster analysis was conducted following Ward’s method, shells (Appendix A). In specimens preserved with a subtle shell which has been specially recommended for morphometrics volume, the umbilical wall is small and gently oblique, showing a (Hammer and Harper, 2006: p. 75). rounded umbilical edge. Very dense, fine, rursiradiate simple and bifurcate ribs dominate throughout the ontogeny, showing 5.1. Cluster analysis furcations on the middle flank. Internal and median whorls have dense, thin, slightly prorsiradiate simple and bifurcate ribs; Fig. 6(A) shows clustering of specimens including those that bifurcations occur on the middle flank, but lower division points provide information on at least one of the ribbing variables. are occasionally present. In large specimens the outer, preserved Three groups have been identified: (i) a small group (dark grey whorl is dominated by simple ribs, which stand out of shell box) which comprises specimens without information about diameters between 29 to 34 mm. One specimen (IGM 6153-1) ER/2 and includes specimens with small and large size shells;

Fig. 5. Scatter diagrams for coiling and external ribs. A. Umbilicus versus shell diameter. B. External ribs per half a whorl (ER/2) versus shell diameter in Mexican and Cuban specimens (note difﬁculty for ER/2 counting under whorl overlapping at lower shell size in large specimens). C. External ribs per half a whorl versus shell diameter for Cuban specimens only. Graphes de dispersion pour l’enroulement et les côtes externes. A. Diamètre de l’ombilic versus diamètre de la coquille. B. Côtes externes par demi-tour (ER/2) versus diamètre de la coquille (noter la difﬁculté de mesurer ER/2 dans les tours internes des grands spécimens du fait de l’enroulement). C. Côtes externes par demi- tour versus diamètre de la coquille des spécimens cubains. 126 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132

Fig. 6. Cluster diagrams. A. Three main groups considering specimens with quasi-complete data for all variables. B. Two main groups considering only specimens with records for the complete set of variables. Bolds for Cuban specimens (Italics for butti); underlined for small Mexican specimens; normal case for large Mexican specimens. Dark grey box for specimens in which the complete dataset of variables was unavailable. Diagrammes degroupement. A. Trois groupes principaux pour les spécimens ayant des données presque complètes pour toutes les variables. B. Deux groupes principaux pour les spécimens ayant des données pour l’ensemble des variables sélectionnées. Spécimens cubains en caractères gras (butti en italiques) ; petits spécimens mexicains en caractères soulignés, grands spécimens mexicains en caractères normaux. Le cadre en gris foncé montre les spécimens à données incomplètes. A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 127

(ii) the group in the middle (light grey box), which includes the to only 0.17 with respect to UR/2. Therefore, PC1 informs majority of large size shells and shows Cuban butti specimens about the covariation of variables under the severe influence of (bold italics in Fig. 6(A)) and antilleanum specimens (normal shell size which forces sculpture and determines: bold in Fig. 6(A))–notice the inclusion in this group of IGM 6085-1 which has a small diameter but shows simple ribs; and a reduction in peripheral ribbing rather than in coiling degree; (iii) the group which includes the majority of small size shells a very limited influence over development of primary ribs (non-shaded box). Except for group (i), the separation of groups (see Component 1 in Fig. 7(A)). (ii) and (iii) has been made within high values of similarity (< 0.4), and the influence of preserved shell size is clear for PC1 seems to be responsible for the clear separation of intragroup clustering. typical butti phenotypes. Fig. 6(B) shows a more secure information when specimens PC two (PC2) shows a significant and positive correlation that lack some of the variables are removed (i.e., group [i] with variables U/D and UR/2 (0.59 and 0.64, respectively), and above). Two groups have been differentiated in this figure a slightly lower correlation with ER/2 (0.44). Correlation with according to shell size, which show smaller specimens other variables, whether positive or negative, is lower than 0.10 (underlined in the non-shaded box) separated from greater and therefore, not worth mentioning. Thus, PC2 informs about ones (light grey box). In these groups, almost all Cuban shell type in terms of coiling degree relating sculpture through specimens allocated to P. antilleanum (i.e., Imlay’s [1942] direct covariation with the number of primary ribs and the USNM103421a paratype; Judoley and Furrazola’s [1968] slightly lower, direct correlation with the proliferation of specimens J-F-126 and J-F-127) are included, together with peripheral ribs. PC2 seems to help reinforce morphologic Imlay’s holotype of P. butti USNM103417 (bold italics). Thus, separation of typical antilleanum vs. butti phenotypes since the clustering in Fig. 6(B) confirms the above-mentioned major latter are larger, more evolute shells showing a lower relative influence of shell diameter for discrimination, but it does not number of bifurcate ribs. help to refute the null hypothesis. PC three (PC3) shows a low proportion of explained variance (13.76%), and hence its interpretation is confusing in a context of 5.2. Principal Component Analysis (PCA) strong influence of shell size. The values for H/D (0.87) and H (0.34) together with U/D (–0.24) and UR/2 (0.23) indicate a well This type of analysis is especially appropriate when known but here significantly secondary relationship between variability and the number of involved codependent variables whorl-high and coiling degree, which is interpreted as a result of (Table 1) are high. taphonomic noise (crushing). In addition, PC3 informs about the Even so, the PCA of log-transformed data was conducted on low incidence of shell type (coiling degree and whorl-high) on the complete database since values from different units should sculpture in terms of the number of primary ribs. be performed in this way (Hammer and Harper, 2006: p. 86). PC four (PC4) shows even a lower proportion of the Taking into account the complete data set, the PCA shows that explained variance (8.62%), but together with the previous PC1 up to four Principal Components (PC) are needed to explain to PC3 94.3% of the total variance is explained. High 92.2% of the total variance. We removed the specimens that covariation of coiling and sculpture seems to be evident showed no data for some of the variables and then performed through the significant, inverse correlation between U/D (0.68) the PCA analysis again. Thus, PC one and two explain 71.9% of and UR/2 (–0.59). Thus, PC4 indicates covariation between variance; together with component three, the explanation periumbilical sculpture and coiling, irrespective of shell size. In potential of PCA reaches 85.7%; and including component this way, this component connects with phenotype traits four, 94.3% of variance is explained (see eigenvalues and relatively ‘‘free’’ from the inertial influence of shell size during percentage of variances in Fig. 7). growth. This interpretation points to a combination of variables PC one (PC1) shows positive correlation close to 0.5 among in response to most probably genetic than environmentally Dm, U, H variables, though a lower one with U/D (0.21), and an forced phenotype expression (reaction norm), the former intermediate but negative correlation with ER/2 (– 0.35). connecting shell type and sculpture–a paradox for PCA when Correlation with other variables is not significant, going down focused on low variable phenotype traits?

Table 1 Pearson correlation matrix between variables. Matrice de corre´lation de Pearson entre variables. DM UHU/DH/D ER/2 UR/2 DM – 1.4678E-20 4.1239E-20 0.17566 0.016569 4.0692E-05 0.15868 U 0.96102 – 7.5372E-15 0.0023852 0.014271 0.00032418 0.057391 H 0.95853 0.91366 – 0.25623 0.48935 0.00010311 0.13194 U/D 0.23079 0.49056 0.19427 – 0.30214 0.97635 0.024088 H/D À0.39681 À0.40502 À0.11901 À0.17686 – 0.14206 0.94821 ER/2 À0.62837 À0.56565 À0.60187 À0.0051205 0.24961 – 0.59351 UR/2 0.23994 0.31964 0.25591 0.37534 À0.01122 0.092018 – 128 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132

Fig. 7. Principal Component Analysis conducted on specimens providing the complete data for all the studied variables; biplot of scores and loadings on the four ﬁrst principal components. A. PC1 vs PC2. B. PC1 vs PC3. Analyses en Composantes Principales pour les spécimens ayant des données complètes pour toutes les variables sélectionnées ; graphiques bivariés des scores et charges sur les quatre premières composantes principales. A. PC1 vs PC2. B. PC1 vs PC3. A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 129

5.3. Remarks range for which Late Jurassic biogeographical units of variable hierarchy have been both proposed (e.g., Uhlig, 1911 in Imlay (1942: p. 1455) separated his species P. butti from Pozˇariska and Brochwicz-Lewin´ski, 1975 but see Arkell, 1956; P. antilleanum when he recognized its coarser and less dense Imlay, 1965; Ziegler, 1971; Enay, 1972, 1980; Cariou, 1973; ribbing throughout the ontogeny. He considered that the Westermann, 1984; Callomon, 1992a) and/or related to absence of intermediate forms reinforced his interpretation of intervals of enhanced endemism (e.g., Cecca, 1999). distinct taxa at the species level. Later authors that identified the Considered to be derived from Lithacoceratinae stocks species created by Imlay did not add new precise comments. during the Early Tithonian, Housaites have not received a The detailed analysis of the ammonites sampled from the Rio precise and conclusive palaeobiogeographic interpretation, as Apulco River section together with measurements obtained the information available at present does not favor it. It is from illustrations of P. butti given by Imlay (1942: Pl. 7, Figs. assumed that the palaeobiogeographic scenario and the 10–12) and Myczyn´ski (1996: Pl. 1, Fig. 1a; 1999, Fig. 7[1]) underlying biogeographic dynamics must be in accordance reveals that clear intraspecies variability is recognized in the with that assumed for ammonites showing Tethyan affinity and final diameter where both furcations disappear and ribbing collected under precise biostratigraphic control in the area (e.g., density changes. In addition, the probable constriction that Olóriz, 1987, 1992; Olóriz et al., 1990; Villaseñor et al., 2000, could be envisaged in Imlay’s holotype (Imlay, 1942: Pl. 7, Fig. 2003; Villaseñor and Olóriz, 2001). 11) at ca. 60 mm of shell diameter is here interpreted as resulting from a rather delayed beginning of simple ribbing. 7. Stratigraphic range After Imlay (1942), citations with illustration of Imlay’s species P. antilleanum were rare. Judoley and Furrazola (1968: Housaites butti (Imlay) has been mainly reported in records Pl. 80, Figs. 1–4) illustrated this ‘‘species’’, but their from both Sierra de los Órganos and Sierra del Rosario, in the description was the literal translation of the original text in Pinar del Rio province. Precise citation of associated species Imlay (1942: pp. 1455–1456). Myczyn´ski (1999: pp. 106, Fig. was given in the introductory chapter. The first record was 7[2]) also illustrated a single specimen interpreted as interpreted as Upper Portlandian (Imlay, 1942: p. 1434). P. antilleanum Imlay. Summarizing, and taking into account updated biostratigra- The close analysis of shell morphology and the multivariate phical correlation provided by Callomon (1992b), Olóriz et al. analysis of measurements of illustrations of P. butti and (1999) and Villaseñor et al. (2000), the citation made by P. antilleanum given by Imlay (1942: Pl. 8, Figs. 4–9) and Judoley and Furrazola (1968: p. 119) clearly indicates the Judoley and Furrazola (1968: Pl. 80, Figs. 1–4) reveal that middle part of the Lower Tithonian through association with taxonomic interpretations made by these authors are in need of Pseudolissoceras and so-called Virgatosphinctes. Reports by revision. Obtaining precise measurements from the specimen Housˇa and De la Nuez (1973: p. 19; 1975: p. 57) refer to Lower illustrated by the above-mentioned Polish author was hindered Tithonian horizons, but more biostratigraphical precision is by fossil preservation. On the basis of rib curves, the paratype unavailable, especially for unclear allusion to the Hauterivian. U.S. Nat. Mus. 103421a illustrated by Imlay (1942: Pl. 8, Data from Myczyn´ski (1989, 1990), Myczyn´ski and Fig. 4), as well as specimens J-F-126 and J-F-127 illustrated by Psczółskowski (1994), and Myczyn´ski (1996) open the Judoley and Furrazola (1968: Pl. 80, Figs. 1, 4) belong to possibility for a more extended range including Upper Housaites butti (Imlay) [Fig. 5(C), see also clusters in Fig. 6(A, Tithonian horizons. The association of Housaites butti (Imlay) B)]. The multivariate analysis presented supports the inclusion with Lytohoplites in Sierra de los Órganos (Myczyn´ski, 1989, of all the new specimens and the previously reported ones (with 1990) indicates uppermost Lower Tithonian. In Sierra del illustration) within the new genus Housaites and the single Rosario, its combined record with Vinalesites below himalayi- species H. butti. tins (doubtful Dickersonia in Myczyn´ski, 1989: p. 59), as well as with Lytohoplites, Protancyloceras, Pseudolissoceras, and 6. Palaeobiogeographic range Vinalesites, and with Virgathosphinctes and ammonites interpreted as Hildoglochiceras (Salinites)(Myczyn´ski and As stated for genus Housaites. In Mexico, reported records Psczółskowski, 1994) also indicates mid-to-uppermost Lower of this species refer to the Sierra Norte de Puebla, eastern Sierra Tithonian (two-fold division) to lower Upper Tithonian. In Madre (Imlay, 1980: p. 35; this paper). The specimens addition, lower Upper Tithonian horizons in Sierra de los described were collected from the Apulco River section MT- Órganos could apply to records of Housaites butti (Imlay) from 2 near Mazatepec, Puebla, Mexico. horizons above those containing Lytohoplites with so-called The known, proved records of Housaites indicate a relatively Aulacosphinctoides, Pseudoinvoluticeras, and ‘‘Virgato- limited area within the southern palaeomargin of the north- sphinctes’’, as well as for horizons where this species only American Plate, which embraces eastern Mexico and north- associates to Hildoglochiceras (Salinites)(Myczyn´ski, 1989). western Cuba. Although the occurrence that was foreseen in In Sierra del Rosario, lower Upper Tithonian horizons, most southern USA (Myczyn´ski, 1989) must be demonstrated, the probably from the Microcanthum Zone (Olóriz et al., 1999: Fig. endemic character of Housaites within Mexico-Caribbean 8), could be the most reasonable interpretation of records in the areas is real, as Myczyn´ski (1989) proposed for ‘‘species’’ butti Himalayites-Corongoceras assemblage identified by Myc- and antilleanum. These areas belonged to a palaeogeographical zyn´ski and Psczółskowski (1994), which contains the upper- 130 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 most occurrences of Chitinoidella below an assemblage of specially relevant, (iv) no difference exists between inner calpionellids of a rather inconclusive age. The interpretation of whorls of individuals thought to belong to nominal species precise horizons for Housaites butti (Imlay) within the P. antilleanum and P. butti. In such a context, we interpret that Microcanthum Zone depends on the precise tintinnoids it the null hypothesis of antilleanum and butti phenotypes being associates with (Chitinoidella in the lowermost and calpio- extreme types in a palaeobiospecies can apply. More research nellids for envisaged records in upper horizons within the on newly collected and well-preserved material is necessary to Microcanthum Zone). Later, Myczyn´ski (1999: p. 106) consider if these extreme types are an expression of sexual assigned an Early Tithonian age without making any comments dimorphism or just morphotypes of intraspecies diversity. The that supported his interpretation, but Psczółskowski and possibility of incomplete preservation being the single fact Myczyn´ski (2003: p. 555, 562) reported P. butti as well as responsible for separation of these extreme types seems to be P. antilleanum in the lower Tithonian of Sierra de Camaján and less probable. mentioned that their observations suggest the disappearance of In accordance, we have selected the species H. butti (Imlay) these species early into the Berriasian. as the single, valid species-level taxon to refer the ammonites In Mexico, only Housaites butti (Imlay) was reported by that have been analyzed. The use of genus-level taxon Imlay (1980: p. 35) based on an isolated record from Jonotla Butticeras (HousˇainHousˇa and De la Nuez, 1973, 1975) (Puebla) and, therefore, the earliest Late Tithonian age has been proven to be invalid, and the new genus Housaites has interpreted by this author is not reliable. Doubts also apply been proposed instead. to the same age interpretation made by Imlay (1980: p. 35) of On the basis of the material collected under precise the association of Burckhardt’ species Pseudolissoceras zitteli, biostratigraphic control, the stratigraphic range for Kossmatia victoris and Grayiceras? mexicanum in the Housaites butti (Imlay) in Mexico seems to be upper Lower Mazatepec area (Puebla), on the assumption of the real Tithonian (Semiforme/Verruciferum Zone of the Tethyan occurrence of Pseudolissoceras zitteli Burckhardt in this standard). The stratigraphic range of this ammonite taxon in assemblage. The Middle Tithonian age interpreted by Cantú- Cuba is not conclusively known. Chapa (1967: p. 21) for this assemblage, which he defined, better accords with the widely accepted range for Acknowledgements Pseudolissoceras zitteli (Burckhardt) according to Callomon (1992b), Olóriz et al. (1999), and Villaseñor et al. (2000). The authors thank the field facilities provided by the Imlay’s (1980: p. 35) citation of Paradontoceras? and authorities of the Hydroelectric station in Mazatepec. Mr. Simoceras? within the stratigraphical range of himalayitins Antonio Altamira helped with the photography of specimens. in Tamaulipas (see above) cannot be used for supporting the Fabrizio Cecca (Université Pierre-et-Marie-Curie, Paris) and real occurrence of Housaites butti Imlay in the Upper Tithonian Günter Schweigert (Staatliches Museum für Naturkunde of Mexico. Stuttgart) made interesting suggestions regarding an early The specimens of Housaites butti (Imlay) that have been draft of this paper. This research was supported by CONACYT, described were collected east from Jonotla, but in the same Project 27555T, PAPIIT Project IN 103702, the Institut of region (Sierra Norte de Puebla) as the specimen mentioned by Geology UNAM (Mexico), Project CGL2005-01316 (Spain) Imlay (1980). They are associated to Simocosmoceras and and the EMMI Group of the University of Granada (RNM 178, Simoceras (Villaseñor and Olóriz, 2001; Villaseñor et al., 2000, Junta de Andalucia, Spain). 2003), and therefore indicate the mid Lower Tithonian (Semiforme/Verruciferum Zone in Olóriz et al., 1999). Appendix A. Supplementary data

8. Conclusions Supplementary data associated with this article can be found, in the online version, at doi:10.1016/j.geobios. Mexican and Cuban ammonites referred to Imlay’s species 2008.06.003. P. antilleanum and P.butti have been studied and proved to have been gathered from a relatively limited area within the southern References paleomargin of the north-American Plate, which embraces central-eastern Mexico and north-western Cuba. Anderson, F.M., 1958. Upper Cretaceous of the Pacific Coast. Memoir Geo- The multivariate analysis (PCA) conducted on Mexican and logical Society of America 71, 1–378. Cuban data offers limited resolution since the sample seems to Arkell, W.J., 1956. Jurassic Geology of the World. Oliver and Boyd Ltd, Edinburgh, London. show strong dependence of shell size. However, the relevance Buckman, S., 1909–1930. Yorkshire type ammonites. Wheldon and Wesley of the information gathered from the PCs that have been (Eds), London. analyzed increases in combination with the following facts: (i) Callomon, J.H., 1992a. Upper Jurassic. In: Westermann, G.E.G. (Ed.), The no significative segregation exists in terms of biostratigraphy Jurassic of the Circum-Pacific, 24. Ammonites of the circum-Pacific region within the sample retrieved from the Apulco River section; (ii) (Hillebrandt, von A., Westermann, G. E. G., Callomon, J. H., Detterman, L. M.) Cambridge University Press, New York, pp. 342–359. no segregation exists between Apulco River’ specimens and Callomon, J.H., 1992b. Upper Jurassic, especially of Mexico. In: Westermann, those gathered from other areas in Mexico; (iii) no significant G.E.G. (Ed.), The Jurassic of the Circum-Pacific, Part IV. Biochronology, difference exists between Mexican and Cuban specimens; and, 12. Ammonites zones of the circum-Pacific region (Hillebrandt, von A., A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132 131

Smith, P., Westermann, G. E. G., Callomon, J. H.) Cambridge University Myczyn´ski, R., 1990. Tithonian stratigraphy in the Sierra de los Organos, Press, New York, pp. 261–273. western Cuba: correlation of the ammonite and microfossil zones. In: Cantú-Chapa, A., 1967. Estratigrafía del Jurásico de Mazatepec, Puebla Pallini, G., Cecca, F., Cresta, S., Santantonio, M. (Eds.), Fossili, Evoluzione, (México). Instituto Mexicano del Petróleo, Tecnología de la Exploración, Ambiente, Atti II Convengo Internazionale di Pergola, 1987. pp. 405–415. Sección Geología. Monografía 1, 3–24. Myczyn´ski, R., 1994. Caribbean ammonite assemblages from Upper Jurassic– Cantú-Chapa, A., 1971. La Serie Huasteca (Jurásico Medio-Superior) del Lower Cretaceous sequences of Cuba. Studia Geologica Polonica 105, 91– Centro Este de México. Revista del Instituto Mexicano del Petróleo 3, 108. 17–40. Myczyn´ski, R., 1996. Discusión sobre la presencia de los géneros Paradonto- Cariou, E., 1973. Ammonites of the Callovian and Oxfordian. In: Hallam, A. ceras Spath, 1923 y Lithohoplites Spath, 1925 en el Tithoniano de Cuba. In: (Ed.), Atlas of Palaeobiogeography. Elsevier, Amsterdam, London, pp. 4th International Symposium Cephalopods Present and Past. Granada, 287–296. Spain, Abstracts, p. 125. Cecca, F., 1999. Palaeobiogeography of Tethyan ammonites during the Titho- Myczyn´ski, R., 1999. Some ammonites genera from the Tithonian of western nian (latest Jurassic). Palaeogeography, Palaeoclimatology, Palaeoecology Cuba and their palaeogeographic importance. Studia Geologica Polonica 147, 1–37. 114, 93–102. Enay, R., 1972. Paléobiogéographie des Ammonites du Jurassique terminal Myczyn´ski, R., Psczółskowski, A., 1994. Tithonian stratigraphy and micro- (Tithonique = Volgien = Portlandien l.s.) et mobilité continentale. Geobios facies in the Sierra del Rosario, western Cuba. Studia Geologica Polonica 5, 355–407. 105, 7–38. Enay, R., 1980. Paléobiogéographie et Ammonites jurassiques : « rythmes Olóriz, F., 1987. El significado biogeográfico de las plataformas mexicanas en el fauniques » et variations du niveau marin; voies d’échanges, migrations et Jurásico superior. Consideraciones sobre un modelo eco-evolutivo. Revista domaines biogéographiques. Livre jubilaire de la Société géologique de de la Sociedad Mexicana de Paleontología 1, 219–247. France 1830–1980. Mémoires hors série de la Société géologique de Olóriz, F., 1992. North-central and eastern Mexico. In: Westermann, G.E.G. France 1980, 10, 261–281. (Ed.), The Jurassic of the Circum-Pacific, part 3, Regional Geology and Fernández-Carmona, J., 1995. Jurásico de las provincias occidentales de Cuba. Stratigraphy, Meso-America. Cambridge University Press, New York, pp. In: Riccardi, A.C., Iglesias Llanos, M.P. (Eds.),Jurásico de Cuba y América 100–107. del Sur, PICG 322, Correlación de Eventos Jurásicos en América del Sur, Olóriz, F., Villaseñor, A.B., González-Arreola, C., Westermann, G.E.G., 1990. Buenos Aires. Asociación Geológica Argentina Serie D, 3, Sociedad Las plataformas mexicanas durante el Jurásico Superior. Un ejemplo de Cubana de Geología, Boletín 1, 1995, Cuba, pp. 7–11. áreas de recepción en biogeografía. In: Civis Llovera, J., Flores Villarejo, Hammer, Ø., Harper, D.A.T., 2006. Paleontological Data Analysis. Blackwell J.A. (Eds.), Actas de Paleontología, IV Jornadas de Paleontología, Sala- Publishing Ltd, India. manca. Ediciones Universidad de Salamanca, pp. 277–287. Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001. PAST: Paleontological Statistics Olóriz, F., Villaseñor, A.B., González-Arreola, C., Westermann, G.E.G., Software Package for Education and Data Analysis. Palaeontologia Electro- 1999. Ammonite Biostratigraphy and Correlations in the Upper Juras- nica 4, 9 p. http://palaeo-electronica.org/2001_1/past/issue1_01.htm. sic-Lowermost Cretaceous La Caja Formation of North-Central Mexico Hernández De La Fuente, R., 1996. Sedimentología y diagénesis del Kimme- (Sierra de Catorce, San Luís Potosí). In: Olóriz, F., Rodríguez-Tovar, F.J. ridgiano en la región de Jonotla, Puebla y Tlacolula, Veracruz. M.D. thesis, (Eds.), Advancing Research on Living and Fossil Cephalopods. Kluwer Facultad de Ingeniería, Universidad Nacional Autónoma de México, Academic/Plenum Publishers, New York, pp. 463–492. México (unpublished). Ortega-Gutiérrez, F., Mitre-Salazar, L.M., Roldán-Quintana, J., Aranda- Housˇa, V., De la Nuez, M., 1973. Las faunas de ammonites del Tithoniano y del Gómez, J.J., Morán-Zenteno, D., Alaniz-Álvarez, S.A., Nieto-Samaniego, Infracretaceo de Cuba. Actas de la Academia de Ciencias de Cuba. Instituto A.F., 1992. Texto explicativo de la quinta edición de la carta geológica de la de Geología 3, 18–19. república mexicana Escala 1: 2,000,000. Universidad Nacional Autónoma Housˇa, V., De la Nuez, M., 1975. Ammonite fauna of the Tithonian and de México, México. Lowermost Cretaceous of Cuba. Colloque sur la Limite Jurassique–Crétacé, Pozˇariska, K., Brochwicz-Lewin´ski, W., 1975. The nature and origin of Lyon–Neuchâtel, septembre 1973. Mémoire du Bureau de Recherche Mesozoic and early Cenozoic faunal provinces. Mitteilungen der Geo- Géologique et Minière 86, 57. logische und Paläontologisches Institut, Universität Hamburg 44, 207– Hyatt, A., 1889. Genesis of the Arietitidae. Smithsonian Contributions to 216. Knowledge 26, 1–238. Pszczółkowski, A., Myczyn´ski, R., 2003. Stratigraphic constraints on the Late Ilovaiski, D., 1917. Les ammonites du jurassique supérieur du pays de Liapine. Jurassic-Cretaceous paleotectonic interpretations of the Placetas belt in Ouvrages de la Section de Géologie de la Société Impériale des Amis des Cuba. In: Bartolini, C., Buffler, R.T., Blickwede, J. (Eds.), The Circum- Sciences Naturelles de Moscou 1, 1–180. Gulf of Mexico and the Caribbean Hydrocarbon habitats, basin formation, Imlay, R.W., 1942. Late Jurassic fossil from Cuba and their economic signifi- and tectonics, 79. AAPG Memoir, Tulsa, pp. 545–581. cance. Geological Society of America Bulletin 53, 1417–1477. Salvador, A., 1987. Late Triassic–Jurassic paleogeography and origin of the Imlay, R.W., 1965. Jurassic marine fauna differentiation in North America. Gulf of Mexico Basin. Association of Petroleum Geologist Bulletin 71, Journal of Paleontology 39, 1023–1038. 419–451. Imlay, R.W., 1980. Jurassic paleobiogeography of the conterminous United Semenov, V.P., 1898. Versuch einer Anwendung der statistischen Methode States in its continental setting. Geological Survey Professional Paper 1062, zum Studium der Vertheilung der Ammoniten in dem russischen Jura. 1–134. Annuaire géologique et minéralogique de la Russie. Nova Alexandria 2, ICZN (International Code on Zoological Nomenclature), 2000. International 1–101. Commission on Zoological Nomenclature, 4th Edition. (Spanish version). Schweigert, G., Zeiss, A., 1998. Berckhemeria n. gen. (Passendorferiinae), eine Judoley, C.M., Furrazola, G., 1968. Estratigrafía y fauna del Jurásico de Cuba. neue Ammonitengattung aus dem Unter-Tithonium (Hybonotum-Zone) von Publicación Especial Instituto Cubano de Recursos Mineros, La Habana. Süddeutschland. Neues Jahrbuch für Geologie und Paläontologie. Monats- López-Caballero, G.I., 2006. Estudio bioestratigráfico de la fauna de ammonites hefte 9, 559–576. del Jurásico Superior del área de Mazatepec, Puebla. Thesis, Facultad de Spath, L.F., 1925. Ammonites and Aptychi. Monographs of the Geological Ciencias, Universidad Nacional Autónoma de México (unpublished). Department of the Hunterian Museum, University of Glasgow v. I. On López-Ramos, E., 1979a. Geología de México. 2. 2 edición, México, D.F. Collection of Fossils and Rocks from Somaliland made by Wyllie and López-Ramos, E., 1979b. Geología de México. 3. 2 edición, México, D.F. Smellie 7, 111–164. Morán-Zenteno, D., 1994. Geology of the Mexican Republic. American Spath, L.F., 1926. On new ammonites from the English Chalk. Geological Association of Petroleum Geologist. Studies in Geology 39, 1–160. Magazine 63, 77–83. Myczyn´ski, R., 1989. Ammonite biostratigraphy of the Tithonian of western Steinmann, G., Doderlein, L., 1890. Elemente der Paläontologie. Engelmann, Cuba. Annales Societatis Geologorum Poloniae 59, 43–125. Leipzig. 132 A.B. Villaseñor, F. Olóriz / Geobios 42 (2009) 117–132

Stinnesbeck, W., Adatte, T., Remane, J., 1993. Mazatepec (Estado de Puebla, Villaseñor, A.B., Olóriz, F., González-Areola, C., 2003. First record of the México)–Reevaluación de su valor como estratotipo del límite Jurásico- genus Simocosmoceras Spath, 1925, Ammonitina, in Mexico. Biostrati- Cretácico. Revista Española de Micropaleontología 25, 63–79. graphic and Palaeogeographic interpretation. GFF 125, 49–56. Suter, M., 1990. Geología de la hoja de Tamazunchale, Estado de Hidalgo, Westermann, G.E.G., 1984. Summary of Symposium papers on the Jurassic- Querétaro y San Luis Potosí. Instituto de Geología, Universidad Nacional Cretaceous Biochronology and Paleogeography of North America. Geolo- Autónoma de México. gical Association of Canada Special Paper 27, 307–315. Uhlig, V., 1911. Die marine Reiche des Jura und der Unterkreide. Mitteilungen Wright, C.W., Callomon, J.H., Howarth, M.K., 1996. Cretaceous Ammonoidea. der Geologischen Gesellschaft. Wien 4, 329–448. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology. Part L. Verma, H.M., Westermann, G.E.G., 1973. The Tithonian (Jurassic) ammonite Mollusca 4. Geological Society of America, Boulder, and University of fauna and stratigraphy of Sierra de Catorce, San Luis Potosí, Mexico. Kansas Press, Lawrence, xviii+362 pp. Bulletin of the American Paleontology 63, 107–320. Zeiss, A., 1968. Untersuchungen zur Paläontologie der Cephalopoden des Villaseñor, A.B., Olóriz, F., 2001. Combined record of Simocosmoceras and Unter-Tithon der Südlichen Frankenalb. Bayerische Akademie der Wissen- Simoceras (Ammonitina) in Mexico.In: 1st International Conference on schaften Mathematish-Naturwissenschaftliche Klasse Abhandlungen Neue Paleobiogeography and paleoecology, Piacenza, and Castell’Arquato, Folge 132, 7–190. Abstracts. pp. 131–132. Ziegler, B., 1971. Biogeographie der Tethys. Jahreshefte der Gesellschaft für Villaseñor, A.B., Olóriz, F., González-Areola, C., 2000. Simoceras aesinense Naturkunde in Württemberg 126, 229–243. (Meneghini). A biomarker for signiﬁcant ﬂoodings during Early Tithonian. Zittel, C.A. von, 1884. Cephalopoda. Handbuch der Paläontologie 2, Geological Society America Abstracts with Programs A-469. München.