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The Oldest Camel Footprints from Mexico ABSTRACT the Oldest Camel Footprints from Mexico INTRODUCTION 352 352 Eocene-Early of Tezoatlán, Oaxaca

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The Oldest Camel Footprints from Mexico ABSTRACT the Oldest Camel Footprints from Mexico INTRODUCTION 352 352 Eocene-Early of Tezoatlán, Oaxaca Boletín de la Sociedad Geológica Mexicana / 2018 / 351 The Oldest Camel Footprints from Mexico Eduardo Jiménez-Hidalgo, Rosalía Guerrero-Arenas ABSTRACT Eduardo Jiménez-Hidalgo ABSTRACT RESUMEN [email protected] Rosalía Guerrero-Arenas Laboratorio de Paleobiología, Instituto de Camel footprints have a well pre- En los estratos del Neógeno norteamericano, Recursos, campus Puerto Escondido, Uni- served record in Neogene strata from incluyendo a México, existe un registro de versidad del Mar, Km 2.5, Carretera Puerto North America, including Mexico, huellas de camélidos bien preservado; sin Escondido – Oaxaca, C.P. 71985, Oaxaca, Mexico. however their Paleogene record is embargo, el registro del Paleógeno es mucho much less known. Here, we describe menos conocido. En este artículo describimos the oldest camel footprints from las huellas de camello más antiguas de Mexico, located in the outskirts of México, procedentes de los alrededores de Tezoatlán de Segura y Luna, north- Tezoatlán de Segura y Luna, noroeste de western Oaxaca, southern Mexico. Oaxaca, sur de México. Las impresiones de The camel footprints are preserved huellas están preservadas como epirelieves as concave epirelief in fine-grained cóncavos en areniscas tobáceas del Eoceno tuffaceous sandstone of the late tardío-Oligoceno temprano de la Formación Eocene-early Oligocene Huajuapan Huajuapan. Las huellas se identificaron Formation. These footprints display como Lamaichnum guanicoe, debido a los the diagnostic features of Lamaichnum rasgos que presentan, los cuales incluyen: guanicoe, such as bidigital tracks with huellas bidigitígradas con bordes anteriores pointed anterior ends and rounded acuminados y bordes posteriores redondea- posterior ends, with digit impressions dos, impresiones de dígitos que no convergen that do not converge anteriorly and anteriormente e impresiones de dígitos footprints that have digit imprints conectadas. Los productores potenciales de connected. Potential track makers las huellas incluyen a Paratylopus, Paralabis include Paratylopus, Paralabis and y Pseudolabis. La superficie axial cóncava Pseudolabis. The concave axial surface y las impresiones de uñas de las huellas and nail imprints of the Oaxacan oaxaqueñas, indican que los productores footprints, indicate that the track compartían algunas características anató- makers shared some anatomical foot micas podiales con los camellos actuales. Lo features with extant camels, sug- anterior, sugiere que algunas características gesting that some characteristics of anatómicas del pie de los camellos recientes the peculiar camel foot had already habían evolucionado ya en el Eoceno tar- evolved by the late Eocene-early dío-Oligoceno temprano. Oligocene. Palabras clave: Camelidae, Keywords: Camelidae, Lamaichnum guanicoe, Eoceno- Lamaichnum guanicoe, Eocene- Oligoceno, Tezoatlán, Oaxaca, BOL. SOC. GEOL. MEX. 2018 Oligocene, Tezoatlán, Oaxaca, Mixteca. VOL. 70 NO. 2 Mixteca. P. 351 ‒ 359 http://dx.doi.org/10.18268/BSGM2018v70n2a5 Manuscript received: March 31, 2017. Corrected manuscript received: July 15, 2017. Manuscript accepted: September 29, 2017. The Oldest Camel Footprints from Mexico 352 / Boletín de la Sociedad Geológica Mexicana / 2018 1. Introduction the Plio-Pleistocene of Jalisco (San Juan de los Lagos), associated with bird, proboscidean and Camel footprints are relatively well recorded in felid ichnites (Rodríguez-de la Rosa and Guz- North American Neogene strata, geographically mán-Gutiérrez, 2012); from the Plio-Pleistocene INTRODUCTION ranging from western Canada to central Mexico of southern Puebla (Tepexi de Rodríguez), where (Lucas and Hunt, 2007). In contrast, the Paleo- the camel footprints are associated with bird foot- gene record is scarce; published records include prints, two kinds of felid footprints, proboscidean those from the Oligocene of South Dakota and and small ruminant footprints (Cabral-Perdomo, 193 2013; Jiménez-Hidalgo et al., 2015a); and from the 19 199. Plio-Pleistocene of southeastern Puebla (Tehu- In Mexico, camel ichnites have been formally re- acán), where camel footprints are associated with ported from the Miocene of Durango (El Salto), canid ichnites (Dávalos-Álvarez et al., 2007; Ro- associated with a turkey-like tracks (Lockley and dríguez-de la Rosa and Guzmán-Gutiérrez, 2012) 1 (Figure 1). Figure 1 Map of Mexico showing the states with camel footprint records and a hypsometric map of the study area in Tezoatlán de Segura y Luna, Oaxaca, southern Mexico. 1. El Salto, Miocene of Durango; 2. San Juan de los Lagos, Plio-Pleistocene of Jalisco; 3. Tepexi de Rodríguez, Plio-Pleistocene of Puebla; 4. Tehuacán, Plio-Pleistocene of Puebla. The red stars indicate the ichnofossil localities of late Eocene-early Oligocene age in the outskirts of Tezoatlán, Oaxaca. The Oldest Camel Footprints from Mexico Boletín de la Sociedad Geológica Mexicana / 2018 / 353 INTRODUCTION / GEOLOGICAL CONTEXT Figure 2 Late Eocene-early Oligocene camel impressions from the surroundings of Juquila de León, Oaxaca. 1. MGRN-01, isolated slab with one footprint; 2. MGRN-02, isolated slab with two footprints (A and B); 3. MGRN-03, isolated slab with five footprints (A to E). Scale is in cm. Paleontological research carried out since 2009 in 2. Geological context the Paleogene sediments from northwestern Oax- aca, has allowed us to study several new Eocene and Oligocene fossil localities. Two of these lo- - calities are near Tezoatlán de Segura y Luna, in east of Tezoatlán de Segura y Luna, northwestern northwestern Oaxaca, southern Mexico (Figure Oaxaca (Figure 1), which is part of the Huajuapan 1), where some camel ichnites were discovered. P The aim of this paper is to describe the camel ich- that consists of sandstone, conglomerate, breccia, nites from Tezoatlán and to make some comments 199 about their paleobiological implications. and Ríos-Martínez, 2009). The fossiliferous beds can be correlated with the lower unit of Martiny et al. (2000), which consists The Oldest Camel Footprints from Mexico 354 / Boletín de la Sociedad Geológica Mexicana / 2018 - . 33. 1. 3. 1. Ma are available for the lower unit (Martiny et al., MATERIALS AND METHODS / SYSTEMATIC ICHNOLOGY 2000), which suggest a late Eocene-early Oligo- cene age for the ichfossiliferous beds. 3. Materials and Methods The studied material consists of 11 footprints in concave epirelief (Figures 2 and 3). Eight foot- prints are preserved in three slabs collected from Rosario Nuevo (MGRN), located in the town of Rosario Nuevo. Three additional ichnites were discovered in a sandstone bed near the road to El Naranjo. Measurements were taken as maximum length and width and are expressed in mm. We followed the camel ichnotaxonomy of Lucas and Hunt (2007). Potential producer’s limb length was estimated based on measurements of Prothero (1996) and the body mass estimations of potential producers were derived from the data of Prothero (1996) and 199. 4. Systematic ichnology Class Mammalipedia Vialov, 1966 Order Artiodactipedida Vialov, 1966 Morphofamily Pecoripedidae Remeika et al., 1995 Ichnogenus Lamaichnum Aramayo and Bianco, 1987 Lamaichnum guanicoe Aramayo and Bianco, 1987 Referred material. MGRN-01, isolated slab with one footprint; MGRN-02, isolated slab with 3 Figure 3 Late Eocene-early Oligocene camel impressions from well preserved footprints. Three footprints pre- the road to El Naranjo, Oaxaca. 1. Footprint H1; 2. Footprint H2, which appears to represent an underprint; 3. Footprint H3; note the push up of sediment in the rear of the print. Scale is in cm. The Oldest Camel Footprints from Mexico Boletín de la Sociedad Geológica Mexicana / 2018 / 355 Table 1. Measurements of Lamaichnum guanicoe footprints from that the potential track maker was Lama guanicoe, the late Eocene-early Oligocene of Tezoatlán, Oaxaca. as Aramayo and Bianco (1987) suggested when Specimen Length (mm) Width (mm) Depth (mm) the ichnospecies was erected. The oldest record MGRN-01 38 12 of lamine camels is early Miocene in age (Honey MGRN-02.A 45 40 10 et al. 199 1 SYSTEMATIC ICHNOLOGY / PALEOBIOLOGICAL IMPLICATIONS MGRN-02.B 40 16 MGRN-03.A 65 26 than the estimated age of the ichnofossiliferous MGRN-03.B 60 24 beds. Potential track makers should be members MGRN-03.C 45 39 21 MGRN-03.D 50 42 13 MGRN-03.E 55 42 the late Eocene-early Oligocene (Honey et al., H1 65 61.3 12 1998), as will be discussed below. H2 31 33 H3 64 59 14 Mean 53.3 43.8 16.4 SD* 11.6 9.6 5.8 5. Paleobiological implications of the * Standard deviation. Oaxacan footprints served in a sandstone bed located near the road The published Mexican record of camel ichnites to El Naranjo. includes Miocene and Plio-Pleistocene localities in Description. Footprints are longer than wide northwestern, western and central Mexico (Figure (length/width ratio = 1.21) (Table 1), with an 1). The Lamaichnum guanicoe footprints from Tezo- ovoid outline, with two digit impressions and P- rounded posterior ends. The digit impressions are gene record in the country and are amongst the parallel or slightly divergent and are separated by . 193 19 199. Two nail impressions are evident in most of the Characteristics of the camel footprints from Te- . zoatlán, Oaxaca do indicate some of the foot’s . producer anatomy. The concave axial surface of are 10 to 26 mm in depth (Table 1). Some ichnites show extramorphological variation, such as
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