The fossil fishes of the archaeological site of Palenque, Chiapas, southeastern Mexico

The fossil fishes of the archaeological site of Palenque, Chiapas, southeastern Mexico

Journal of Archaeological Science: Reports 17 (2018) 462–476 Contents lists available at ScienceDirect Journal of Archaeological Science: Reports jo...

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Journal of Archaeological Science: Reports 17 (2018) 462–476

Contents lists available at ScienceDirect

Journal of Archaeological Science: Reports journal homepage: www.elsevier.com/locate/jasrep

The fossil fishes of the archaeological site of Palenque, Chiapas, southeastern Mexico

T



Jesús Alvarado-Ortegaa, , Martha Cuevas-Garcíab, Kleyton Cantalicea a

Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad Universitaria, Delegación Coyoacán, Ciudad de México, 04510, México Dirección de Registro Público de Monumentos y Zonas Arqueológicas, Instituto Nacional de Antropología e Historia, Av. Victoria 110, Copilco El Bajo, Ciudad de México, 04510, México b

A B S T R A C T Since the end of the nineteenth century, different remains of fossilized fishes had been observed or recovered from the Mayan City of Palenque. Although some efforts had been made to identify these objects, their taxonomical nature and possible origin within the geological environment of this archaeological site are poorly understood. In this work, we review these fossils recovered in Palenque and other two Mayan cities, Agua Clara and the El Lacandón, which are deposited into the collections of the Instituto Nacional de Antropología e Historia (Mexico); we describe the morphological characteristics that allow us to identify them taxonomically. This research also reports the results of a paleontological exploration through the different Paleocene-Miocene sedimentary geological units in the surroundings of Palenque; the fossils thus collected belong to the same groups of those found within the archaeological Maya context. These results suggest that such geological units are the sources of all these fossils. In this exercise, three types of anthropogenic features produced by the Maya people were discovered on the fossils. Paint stains and plaster spots suggest that these fossils were painted. The pronounced differential wearing marks in some regions along the cutting edges of shark teeth and ray tail spines show that these fossils were used as cutting tools. Besides, the surfaces excavated around fossils preserved in slabs demonstrate that some Maya people intended on discovering and recognizing these objects, as modern paleontologists do.

1. Introduction At the end of the nineteenth century, Sapper (1896) reported the occurrence of “fossil fish remains in a very fine-grained limestone which resembles the lithographic stone of Solenhofen… used in the construction of Palenque”. Fifty-five years later, Mulleried (1951) provided a superficial description of two fossils bearing rocks collected in this Maya site into the Chiapas State, southeastern Mexico. He recognized the Sapper's sample in a slab carrier of fish remains that “probably” belongs to the fossil genus Pycnodus Agassiz, 1833, as well as other unidentified fishes and some undetermined foraminifera. In concordance with the geological occurrence of Pycnodus known at that time, the Eocene age was suggested for this slab. The second Mulleried's sample, collected in an unreported building of Palenque site, bears “macro and micro-foraminifera (Orbitollites), gastropods, bivalves, and echinoid spines” that suggested the Middle Oligocene–Early Miocene age. The authors of the present paper unsuccessfully tried to locate the specimens referred above. Our search included the collections of the ⁎

Instituto Nacional de Antropología e Historia (INAH), Instituto de Geología (Igl, UNAM), and Instituto de Biología (IB, UNAM). Hence, now it is not possibly to analyze those samples because neither Sapper (1896) nor Mulleried (1951) provided data about the final deposit of such materials. Since the firsts archaeological field investigations on Palenque, RuzLhuillier (1958a–d, 1959) [under the advisement of Dr. Roberto Llamas, then Director of the IB, UNAM] and other authors reported the occurrence of fossil shark teeth and stingray spines from different primary (offerings and tombs) and secondary (building debris and ceremonial dumps) deposits in this ancient city (Borhegyi, 1961; Acosta, 1976; Fernández, 1991; González-Cruz, 1993; Olvera-Carrasco, 1997; López Bravo et al., 2003; Venegas-Duran, 2005). According to CuevasGarcía (2008), the fossil nature of these materials was not originally recognized in some of these reports; hence, the possible patterns in the use of fossil elements in Maya rituals of Palenque had not been the target of accurate researches. The studies of these elements have implications for our understanding on the Mayan cosmology. The discovery of these fossils should encourage the launch of new research

Corresponding author. E-mail address: [email protected] (J. Alvarado-Ortega).

https://doi.org/10.1016/j.jasrep.2017.11.029 Received 4 August 2017; Received in revised form 12 November 2017; Accepted 20 November 2017 2352-409X/ © 2017 Elsevier Ltd. All rights reserved.

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Fig. 1. Geologic map of the northern are of Chiapas showing the Sedimentary geological units around the archeological Paleque City and the paleontological sites prospected during in the present research. 1, División del Norte quarry (IGM-loc 3869); 2, Small ravine near Building XXVI of Palenque; 3, Gilberto Meneses quarry; 4, Belisario Domínguez (IGM-loc 3870); 5, Motiepá; 6, Estrella de Belén (IGM-loc 3900); 7, Chancalá (IGM-loc 3963); 8–10, El Gato (IGM-loc 3636); 11, Sánchez site; 12, Luis site (IGM-loc 3899); Río Chacamax (IGM-loc 3880); Don Eber site (IGM-loc 3879).

as the northern edge of “Sierra Madre Oriental de Chiapas” (Dengo, 1968) and the Reverse-Fault Province defined by Meneses-Rocha (2001). Thus, probably the most emblematic building of this archaeological site, the Temple of the Inscriptions looks northeast towards the Gulf Coastal Plain. The geological environment of Palenque area involves outcrops of the Tenejapa-Lacandón unit; as well as the Lomut and Tulijá formations (Figs. 1, 2). The paleontological prospection performed along the present research included all these geological units. Fourteen paleontological sites were identified; now, some of them are recorded as IGM-loc (Table 1).

lines, so far overshadowed by the undue recognition of these paleontological materials found in surrounding Maya region of Chiapas. In 2008, a joint INAH-UNAM research program was launched. The challenges of this archaeological-paleontological project were: 1) Discovering the taxonomical identity of the fossils recovered from Palenque and two surrounding archeological sites, the El Lacandón and Agua Clara, located to 20 km to the west and 26 km to the south of Palenque, respectively (Fig. 1); 2) Identifying the possible source of these fossils within the geological framework in northern Chiapas (Fig. 2); 3) Describing the archeological context associated to the findings of this fossils; and 4) Exploring the possible meaning of these objects within the different activities and beliefs recorded in the archaeological remains of this emblematic Maya site. The aim of the present manuscript is to provide the descriptions of these fossil remains collected in these archeological sites (Figs. 3–10, Table 1), update their taxonomical identity, and identify their possible source within the surrounding geological environment of Palenque.

1.1.1. Tenejapa–Lacandón Unit (L-TU) This sedimentary unit was informally named by Alvarado-Ortega et al. (2015), when they report the fossil fishes from the Belisario Domínguez and División del Norte quarries. These authors could not recognize the geological formation to which the sediments of these sites belong. This is because these sites share characteristics that previously were described for the Lacandón Formation or for Tenejapa Formation. Also, the limits of these coeval formations are not seen in the available geological maps (Servicio Geológico Mexicano, 2006a, 2006b). Vinson (1962) named the Lacandón Formation based on a sequence of limestone sediments exposed at the west of Petén, Guatemala. Although, Quezada-Muñetón (1987) reported this formation in Chiapas; he named the Tenejapa Formation based on a sequence of cream-yellow brownish marls, limestones, and calcareous breccias with small bioclasts, flint bands and nodules. Based on lithological, paleontological, and structural data, Quezada-Muñetón (1987) concluded that these

1.1. The geological survey Here, the geologic setting of Palenque is described to emphasize the possible geological source of those fossils referred in this study. In this section we also report, name, and provide the characteristics of the paleontological sites discovered in the present research. Palenque is between the coordinates 17°29′21″-17°28′46″ N and 92°03′00″-92°02′46″ W. Its altitude is 105–223 masl. This pre-Hispanic Maya city is on the north side of the “El Mirador” hill, which represents the fold in the northwestern corner of the “Zona Sola” anticline, as well 463

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Fig. 2. General views of the paleontological sites considered in the present work (also see Fig. 1 and Table 2 of the present manuscript). A, Small ravine near Building XXVI of Palenque; B and C, Ambar bearing sands at Estrella de Belén (IGM-loc 3900); D, Rock fragment with fossils from Chancalá (IGM-loc 3963); E, Sedimentary level with ostreids at the El Gato site (IGMloc 3636); F, general view of the Sánchez site.

1.1.2. Lomut Formation Quezada-Muñetón (1987) applied the name of Lomut Formation to a sequence of Eocene marine sediments exposed near Ocosingo that are extended along the center and north of Chiapas, including the Palenque area (Fig. 1). This sedimentary unit consists of quartz sands and gravels up to 2 m. thick, which are interbedded with thin layers of fossiliferous gray-green shales and biogenic cream-yellow dolomites, limestones and marls (Ham-Wong, 1979). Fossils and lithology of the Lomut Formation suggests that it was deposited under marine shallow conditions along coastal lagoons or near shore continental platforms. This formation is laterally interdigitated with sediments of the El Bosque Formation; its bottom lies transitionally over the L-TU; and its top is covered in discordance for sediments of the Tulijá Formation (Quezada-Muñetón, 1987). Fossils from this formation include a large array of microfossils, bivalves, gastropods, echinoderms, corals, algae, bryozoans, and mammals (Pérez-Martínez and López-Palomino, 2011). In this project, fossils from the Lomut Formation were collected in the Motiepá site; these include millimetric shark teeth of uncertain affinity.

early-middle Paleocene formations were deposited under marine condition along the same inclined platform, in which the Lacandón sediments were deposited under inshore and shallow conditions (southeastern Chiapas and northern Guatemala) while the Tenejapa deposits occurred under offshore and deep conditions (northwestern Chiapas). Fossil fish from the L-TU were collected in three quarries, División del Norte (IGM-loc 3869), Belisario Domínguez (IGM-loc 3870), and a small ravine near the Building XXVI of Palenque once thought as a quarry exploited by the Maya (Figs. 1, 2; Table 1). Isolated fish bones were also observed in slabs in the Gilberto Meneses quarry; however, these were not collected because their poor preservation and the large size of the slabs that carried them. Locating these sites was a simple task because the local people extract slabs from some centimetric-decimetric thick layers of the L-TU with commercial proposes. These strong slabs are used in the construction (on floors and walls) and in the artisanal reproduction of the low-reliefs found in the ruins of Palenque. The fossil assemblage collected in L-TU is mainly represented by fishes. These include the osteoglossiform Phaerodus Leidy, 1873; the pycnodontiform Pycnodus Agassiz, 1833; perciformes as Eekaulostomus cuevasae Cantalice and Alvarado-Ortega, 2016, Kelemejtybus castroi Cantalice and Alvarado-Ortega, 2017; as well as clupeiformes, gonorynchids, eels, sea basses, and other unidentified teleosts (Cuevas-García and Alvarado-Ortega, 2009; Cuevas García and Alvarado Ortega, 2012; Alvarado-Ortega et al., 2015; among others). Other fossils from LT-U are crabs, coprolites, fragments of a turtles, and carbonized plants. A small dentary bone of an unidentified pycnodontiform fish was recovered from the small ravine near the Building XXVI. Although some scattered fish bones and scales are preserved on the slabs extracted from the Gilberto Meneses quarry; these are so scarce and badly preserved that they were not collected.

1.1.3. Estrella de Belén Amber sands In 2009, we found a small mine of sands near the Estrella de Belén village (IGM-loc 3900), where the local people recover amber (Figs. 1, 2). Here, quarzitic sand strata 1–2 m thick are interbedded with thin green-reddish clays. Although the tops and bottom of this strongly folded strata are unknown; these, bear abundant carbonized plant remains and nodules of pyrite. The paleontological and lithological characteristics of this amber site do not match with those of the Tulijá Formation, as it is already mapped (SGM, 2006a, 2006b); hence, the geological designation of this amber bearing sands in the Estrella de Belén quarry, as part of the Miocene Tulijá Formation, is challenged in 464

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Fig. 3. Fossil teeth of Carcharocles megalodon collected in different archeological context in the Maya sites of Palenque, Agua Clara, and the El Lacandón, in Chiapas, southeastern Mexico (see Table 1). A, MUPAL 2730; B, MUPAL 2571; C, Specimen 120 (Olvera-Carrasco, 1997); D, MUPAL 2568; E, REG 1637 PJ 9; F, MUPAL 2770; G, MUPAL 2578; H, MUPAL 2573; I, MUPAL 2575. J, MUPAL 3291; K, REG 1638 PJ 3. Abbreviatures: Cv, carinal or lateral view; Lb, labial view; Lg, lingual view; Rv, root view. Scale bars = 10 mm.

with the present data because the Estrella de Belén amber site belongs to the Salto de Agua Municipality (Table 1). Other amber localities, present in the center and northwest of Chiapas (Solórzano-Kraemer, 2010), constitute deposits of the Late Oligocene- Early Miocene sediments of the La Quinta Formation (= Simojovel Formation), Lower-Middle Miocene of the Mazantic Shale, and Middle Miocene of Balumtum Sandstone (Castañeda-Posadas

this work (Fig. 2). After the discovery of amber from Estrella de Belén, SolórzanoKraemer (2010) reported the discovery of a new amber locality within the Palenque Municipality based on a personal communication of a local paleontologist that heard vague news about this new site. Currently, we are not aware of any amber mine within the limits of the Municipality of Palenque. Therefore, such a report must be updated 465

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Tulijá” for a fossiliferous calcareous sequence exposed along the Tulijá river, which is extended up to Palenque (Fig. 1). The Tulijá Formation consists of dark brown-bluish clay strata interbedded with white gray-reddish millimetric laminated limestones and sandy clays. These 30 to 150 cm thick sequence shows frequent facies changes, in which a large array of marine fossils have been collected (foraminifera, ostracods, gastropods, bivalves, echinoderms, corals, crabs, and vertebrate remains). Such fossil assemblage and lithology suggest that during the Early Miocene, the Tulijá formation sediments were deposited in a marine environment near shore, under shallow and high energy conditions which may include transgressive and regressive episodes along the inner platform and/or coastal lagoons (Meneses-Rocha, 2001; Riquelme et al., 2012; among others). Usually the abundance of invertebrate fossils is so high that they really form embedded coquinas. During this research, fossils of the Tulijá Formation were collected in eight sites (Figs. 1, 2; Table 1). Although, these fossils include a large array of invertebrates as foraminifera, ostracods, bivalves, corals, echinoderms, crabs, possible tubes of polychaete worms, and sponges; different vertebrate remains have been collected in seven sites, including three spots at the El Gato (IGM-loc 3636), Río Chacamax (IGMloc 3880), Chancalá (IGM-loc 3963), Don Eber site (IGM-loc 3879), and Luis site (IGM-loc 3899). No fossils were collected in the Sánchez site near the Lacandón ruins (Table 1). These vertebrates consist of isolated teeth of sharks, rays, and teleostean fishes, fish scales, as well as, sirenid bones. Among invertebrates, it is remarkable the presence of banks of the ostreid Crassostrea Sacco, 1897 in life position (Velasquillo-García, 2011) and the abundant of remains of portunid crabs, including one almost complete specimen of Necronectes proavitus (Rathbun, 1918) (Martínez-Ortiz et al., 2017), already identified in these paleontological sites.

Fig. 4. MUPAL 2569, fossil tooth of Negaprion sp., lemon shark, collected in the archeological site of Palenque, Chiapas, southeastern Mexico. Abbreviatures: Lb, labial view; Lg, lingual view; Rv, root view. Scale bar = 10 mm.

2. Material and methods 2.1. Institutional abbreviations The fossils referred in this manuscript are formally deposited into the following scientific collections: IGM, Colección Nacional de Paleontología, Instituto de Geología, Universidad Nacional Autónoma de México. IHNFG, Paleontological Museum “Eliseo Palacios Aguilera” belonging to Instituto de Historia Natural y Ecología, Chiapas. INAH, Instituto Nacional de Antropología e Historia. MUPAL, Museo de Palenque. Numbers without acronym (i.e. 10.335155) represent numbers of INAH inventory. Specimens with no INAH-inventory numbers from the Agua Nueva and El Lacandón sites are indicated with field numbers (i.e. REG 1637 PJ 9). The paleontological sites reported in the present study are recorded into the catalog of paleontological localities of IGM, under the acronym IGM-loc.

2.2. Material analyzed Fig. 5. Myliobatiform caudal fin spines recovered in Palenque. A, REG 1638 PJ 7; B, MUPAL 2579; C, REG 1638 PJ 1. Abbreviatures: Dv, dorsal view; Lv, lateral view; Vv, Ventral view. Scale bars = 5 mm.

The present study includes 68 fossil fish specimens recovered in the archaeological sites of Palenque, Agua Clara, and the El Lacandón (Cuevas-García, 2008: 483–484) (Table 1). Unfortunately, only 42 of these are available within the INAH collections; the remaining specimens seems be lost (L1 to L16 in main text and Table 1). The available specimens are identified with the formal numbers of the INAH inventory, catalog numbers of MUPAL, and in the specimens from the Agua Clara and El Lacandón sites were considered the field control numbers (Table 1, Figs. 3 to 10). The specimens MUPAL 2787 and MUPAL 2731 are marl slabs bearing numerous fish remains; in this work, each of such fossils are identified with progressive numbers (Figs. 6, 7).

and Cevallos-Ferriz, 2007; Riquelme et al., 2014; among others). After a crude comparison, it is possible to suggests that the Estrella de Belén amber sands correspond to those Miocene coarse quartz sands of La Quinta Formation. According to local people, sediments of this new site amber also bears organic inclusions, as insects and plant remains.

1.1.4. Tulijá Formation The author of this geological unit is obscured (Sáenz-Pita and LópezPalomino, 2011); however, López (1964) used the name “Mioceno 466

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Fig. 6. MUPAL 2587, 1 fossil fish bearing slab recovered in the archeological site of Palenque. 1–2, Pycnodus; 3, Kelemejtubus castroi; 4–7, teleosts, possible gonorynchiforms; 8, fish remain of uncertain affinity; 9, serranid. Scale bar = 100 mm.

Fig. 7. Fossil fish bearing slab MUPAL 2737, collected in Palenque, Chiapas, southeastern Mexico. Abbreviatures: 1–7, Pycnodus; 8–15, Kelemejtubus castroi; 16, isolated lower jaw of indeterminate fish; 17, serranid; 18–23, teleosts, possible gonorynchiforms; 24–26, fish remain of uncertain affinity. Scale bar = 200 mm.

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Fig. 8. Wear marks in fossil fish remains collected in Palenque. A, MUPAL 2730, Carcharocles megalodon tooth with both cutting edges disturbed, small arrows show the serrations eroded probably by repetitive reciprocating movements and the large arrows show a couple of notch probably generated by hitting; B, Unaltered serrations in the tooth carinae of C. megalodon, close op of MUPAL 2575; C, MUPAL 2578 (ventral view above and dorsal view in the bottom), a myliobatiform caudal fin spine showing a noticiable notch on its lateral serrated border eroded probably by repetitive reciprocating movements. Scale bar in C = 5 mm.

Fig. 9. Preparation marks on fossil specimen of the slab MUPAL 2587.

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Fig. 10. Color stains and plaster patches features on the slabs MUPAL 2587. Abbreviatures: b, blue r, red; p, plaster. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

2.3. Comparative materials

3. Results

Fossils collected out of the archeological context mentioned in this paper were studied with comparative proposes. Some of them fossils were previously analyzed by different authors. These fossils include: Serranidae, undefined species, IGM 4550, from División del Norte (Alvarado-Ortega et al., 2015). Pycnodus sp., IGM 4551, IHNFG-4890, both from División del Norte (Alvarado-Ortega et al., 2015).

This section consists of two parts. The first part deals with the description and the taxonomic affinities of the referred fossil specimens (Figs. 3–8; Tables 1, 2). The anthropogenic features observed in some of these fossils are documented in the second part (Figs. 8–10). 3.1. Fossil fishes from the Mayan archeological contexts 3.1.1. The “megatooth” shark teeth It is remarkable the long and unfinished dispute among shark taxonomist concerning the correct name or names to be applied to the most famous species among the “megatooth” sharks. Originally, this species was named as Carcharodon megalodon Agassiz, 1834, and considered as part of the family Lamnidae, together with the living “great white shark” Carcharodon carcharias Linnaeus, 1758; however lately, it was relocated into the family Otodontidae and the genus Carcharocles Jordan and Hannibal, 1923 (Casier, 1960; Muizon and de Vries, 1985; among others). Hence, this species is referred as Carcharodon megalodon by some authors (i.g. Applegate and Espinosa-Arrubarrena, 1996; Purdy et al., 2001) or as Carcharocles megalodon by others (i.g. Cappetta, 1987;

2.4. Taxonomy and morphological nomenclature The names of the taxonomic groups of those fishes referred here, as well as the nomenclature of their conserved bone structures and described in this work, follow the authors cited in each case.

2.5. Geological Nomenclature The names and maps of all the sedimentary geological units indicated in this work are registered in the Maps and Stratigraphic Lexicon of Mexico published by the Mexican Geological Service (Fig. 1). 469

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Table 1 Fossil fish remains recovered into the archeological context of the Mayan cities of Palenque (at the top), the El Lacandón (at the center), and Agua Clara (in the bottom), which are included in the present study. All the specimens are deposited in the INAH collection except for those referred as L1 to L16 that seems to be lost. Catalog number (INAH inventory)

Original taxonomical identification

Archeological context of the discovery

MUPAL 2568

Shark tooth

MUPAL 2569 MUPAL 2570 MUPAL 2571

Fossil tooth Fossil tooth Fossil shark tooth

MUPAL 2572

Not identified

MUPAL 2573 MUPAL 2575

Not identified Not identified

MUPAL 2576

Bone or tooth

MUPAL 2578 MUPAL 2579 MUPAL 2587 (10.335185)

Stingray spine of “Dasyatis sp.” Stingray spine Slab with fishes

MUPAL 2730 (10.335155)

Shark tooth

MUPAL 2731

Slab with fishes

MUPAL 3291 (10.335156)

Shark tooth

MUPAL 3292

Stingray spine

120

Shark tooth

L1

Tooth of “Odantaspis cuspidate”

L2 to L8

Tertiary elasmobranch vertebrae

L9

Carcharodon tooth

L10 L11

Probably tertiary Myliobatis serratus or M. magister Tertiary Myliobatis sp.

L12 to L14 L15 and L16

Fossil stingray spines A shark tooth and a stingray spine

REG 1638 PJ 1

Shell ring/shark tooth

REG 1638 PJ 3

Tooth

REG 1638 PJ 7

Stingray spine

REG 1637 PJ 9

Lithic material

Associated with the human bones of right arm found in Tomb 7, Building 5, Grup B (González-Cruz, 1993). Content of Tomb 1, Building 2-B, Grup C; collected together with specimen MUPAL 2770 (Fig. 4) Content of Tomb 1, Building 2-B, Grup C; collected together with specimen MUPAL 2569. Borehole within Ball Stone Set (Venegas-Duran, 2005: 58–59). Field data: “Pit 7, Layer II, Level 2, Bag 485”, VDBJ (on the fossil). Secondary archeological context (Venegas-Duran, 2005). Field data: “P.C.U. 04, Capa II, nivel 5, pozo 65, 1147, VDBJ” and 12 (on the fossil). Dump of The Palace. Field data: “The garbage dump, March 23, 1992” Dump of The Palace, Field data: “E1 Basurero, Capa XI (100− 110), Bolsa 443, No. 0452, excavators LSL y AGC”. Collected in June 28, 1991. Secondary archeological context, floor of north side of corridor 2–3 of Building 3, Group B. Field data: “Project Group B, Layer B, Bag 35, No. B. 044”. Landfill in the floor of Room 6, Building 3 at Grup B. Associated with the human bone, Tomb 7, Building 5, Group B. Among the debris of The Palace (Ruz-Lhuillier, 1959). Note: This slab show nine fish remains (Figs. 6, 9, and 10) Into the radial altar located at the center of the plaza between Temple of the Cross, Temple of the Foliated Cross, and Temple of the Sun (Ruz-Lhuillier, 1958b). (Figs. 3, 8) Tap of the tomb in the Building 3, Bat Grup (González-Cruz, 1993). Note: This slab has remains of at least 25 fishes (Fig. 7). Into the cylindrical box with a cross-shaped cavity found in Temple V of North Grup (Ruz-Lhuillier, 1958d). Tumb III, Count Temple (Ruz-Lhuillier, 1958c, p. 208, fig. 14-v, lám. XLVIII-m). According to this author, the spine is similar with those found before in the Temple of the Cross and Temple of Foliated Cross. Secondary context (collapsed debris), Temple II of Group North, Palenque (Olvera-Carrasco, 1997, p. 254, 260, 275). Part of the Offering II placed into a pot with lid, found in the floor of Temple of the Cross, together with shark vertebrae (L2 to L8) and a stingray spine (L12) (Ruz-Lhuillier, 1958a, p. 79; García-Moll, 2007, p. 262, lam. XVIII). Part of the Offering II placed into a pot with lid, found into the Temple of the Cross, together with a shark tooth (L1) and a stingray spine (L 12) (Ruz-Lhuillier, 1958a, p. 79; García-Moll, 2007, p. 262, lam. XVIII). Part of the offering deposited within a vessel taped with a bowl, unearthed in the floor of Temple of Foliated Cross (Ruz-Lhuillier, 1958a, p. 85–88) Part of the Offering I, deposited a rouded pot, found in the Temple of the Cross (Ruz-Lhuillier, 1958a, p.76–79, fig. 4h-h´, lam VIIIa; García-Moll, 2007, p. 262, lam. XVIIIa). Part of the Offering II preserved in a pot with lid, found in the floor of Temple of the Cross, deposited together with the specimens L1 and L2 to L8 (Ruz-Lhuillier, 1958a, p. 76–79, fig. 4i, lam. XVIIIb; García-Moll, 2007, p. 262, lam. XVIIIb). A tomb in Palenque (Acosta, 1976, p. 37) Part of the Offering preserved in a pot, together with bird bones and jade remains (Fernández, 1991, p. 240). Preserved in three parts associated to Burial 2 of Unit 10 at the El Lacandón site. Field data: “March 1 of 2001, U-10, Element 3, Tomb 2, Lot 13, UCM” (Cuevas-García, 2008, p. 483–484) Near the Structure 45-A, Unit 4 into the El Lacandón. Field data: “March 1, 2001; Square H-3, Layer 1, Level 1, Bag 158, Lot 17, NZNR”. Preserved into the Burial 2 of Unit 8 at the El Lacandón. Field data: “May 15, 2001; El Lacandón 01, Square AE2/AE3, L II, Bag 127, Lot 32, RLB”; 127 (in the fossil). Secondary archaeological deposit, Pit 5, excavated in the Plaza of archeological site of Agua Clara.

The most complete teeth studied here are robust and triangular structures (Fig. 3), in which the crown consists of a single smooth cusped, not very thick, with two sharp cutting edges (carinae) intensely serrated, almost flat and in the lingual surface, and slightly curved in the labial. In contrast, the roots consist of two almost symmetrical lobes, deeply separated by a central notch, about as width as the crown base, and uniformly thick except in the center where a prominent torus rises around the central foramen. Both carinae show a line of rounded serrations, uniformly sized and closely spaced. All teeth studied herein, complete or fragmented, share the presence of these serrations, which make it possible to assume that these fossils belong to the same taxonomic group (Table 3). A recent morphometric study of megatooth shark teeth, including Carcharodon and relatives, concludes that Carcharocles megalodon as other “megatooth” species with small crown cuplets have teeth with cutting edges entirely covered with a line of rounded serrations,

Nyberg et al., 2006). Twelve megatooth shark teeth of extinct Carcharocles megalodon from different Maya archeological contexts are identified in the present research (Fig. 3, Table 1). These specimens are MUPAL 2730, MUPAL 3291, MUPAL 2568, MUPAL 2575, MUPAL 2573, MUPAL 2770, MUPAL 2576, MUPAL 1571, and Specimen 120 from Palenque (RuzLhuillier, 1958b, 1958d; Olvera-Carrasco, 1997; González-Cruz, 1993; Venegas-Duran, 2005). Additional specimens are REG 1637 PJ 9 and REG 1638 PJ 3, respectively recovered in the Agua Clara and the Lacandón sites. The lost specimen L9 from Palenque (Table 1) and previously identified as a Carcharodon tooth by Ruz-Lhuillier (1958a, p. 85–88), probably also belongs to C. megalodon. Since this is an heterodont shark, the nomenclature defined by Applegate and EspinosaArrubarrena (1996) and Purdy et al. (2001) is used here to name the position of these teeth along the tooth set. Table 2 summarizes the measurements of most complete teeth. 470

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Table 2 Names, ages, and locations of the paleontological sites around the Maya city of Palenque, prospected in the INAH-UNAM archeological-paleontological project reported in this work this project (sites are numbered as in Fig. 1). Paleontological sites

IGM-loc

Geological unity (age)

Geographical coordinates N - W

Municipality

14 13 12 11 10 9 8 7 6 5 4 3 2 1

3879 3880 3899 – 3636 3636 3636 3963 3900 – 3870 – – 3869

Tulijá Fm. (Miocene)

17°29′29.7″–92°3′46.2” 17°28′49.43″–91°58′28.64” 17°29′20.8″–91°58′7.3” 17°28′14.0″–91°52′8.1” 17°29′29.4″–92°56′29.5” 17°29′59.4″–92°56′30.7” 17°29′21.9″–92°56′33.1” 17°19′11.2″–91°41′33.3” 17°22′58.66″-91°57′40.77” 17°29′21.9″-92°3′0.70” 17°25′28.6″-91°58′46.8” 17°26′9.3″-91°59′14.2″

Palenque

Don Eber site Río Chacamax Luis site Sánchez site El Gato 3 El Gato 2 El Gato 1 Chancalá Estrella de Belén amber sands Motiepá Belisario Domínguez Gilberto Meneses Near Building XXVI División del Norte

La Quinta Fm. (Lower Miocene) Lomut Fm. (Eocene) Tenejapa–Lacandón unit (Paleocene)

Salto de Agua Palenque Salto de Agua Palenque

17°27′59.0″-92°1′59.9″

(Fig. 4). This has both ends of the root broken; however, its crown reveals enough characteristics identifying it as a left lateral tooth of the upper of jaw. The root and crown heights of this tooth are 14 and 19 mm respectively. In the triangular crown, the tip is curved distally, the enamel is smooth, and the labial surface is almost flat and bears a shallow middle basal-apical groove extended along one third of the crown height; in contrast, its labial surface is gently convex. In the crown, the mesial cutting edge is convex and forms a long shallow shoulder with a small notch, the distal cutting edge is slightly concave close to the tip and curved backward near the root. Both cutting edges are entirely smooth, except in the distal shoulder that has tiny serrations. In the root, the end of both lobes and large part of the basal border are missing. In cross section, the root is slightly thicker that the crown base. The middle part of the root protrudes lingually around central foramen. An inconspicuous transverse groove runs from the central foramen toward the root base. Negaprion groups four marine species. The living species N. brevirostris (Poey, 1868) and N. acutidens (Rüppell, 1837) are present along the tropical areas of the Indo-Pacific and Atlantic oceans. Fossils of N. brevirostris are known from Early Miocene-Pleistocene sediments of America, Europe, Asia, and Australia (Williams, 1999). In contrast, the extinct species show a disjoint temporal distribution, N. amekiensis (White, 1926) is present in Eocene deposits of Nigeria while N. eurybatrodon (Blake, 1862) has been collected in Neogene sediments of France, Portugal, Peru, Venezuela, Pakistan, and USA (Cappetta, 1987; Sánchez-Villagra and Aguilera, 2006). According to Purdy et al. (2001:156), N. brevirostris is synonymous with N. eurybatrodon. The crown tooth of MUPAL 2569 shows three diagnostic characteristics of Negaprion. These are the cutting edges are smooth except for the weak serrations in the distal shoulder, the shallow transverse groove, and the small notch in the long lateral shoulders. MUPAL 2569 6 differs from teeth of N. amekiensis that has teeth with edges completely serrated (Cappetta, 1987). Unfortunately, MUPAL 2569 is undifferentiable from N. acutidens and N. brevirostris, which are diagnosed on external body characteristics and vertebrae accounts; therefore, this fossil is identified only at generic level. Previously in México, fossil teeth of Negaprion were collected in middle Miocene deposits of Baja California (Minch et al., 1979; ArandaManteca, 1994). In the present project, teeth of Negaprion were collected in deposits of the Tulijá Formation at Río Chacamax (IGM-loc 3880) and Chancalá (IGM-loc 3880) sites (Fig. 1, Table 1). This represents the first report of Negaprion in marine deposits of Chiapas.

Table 3 Measurements (in millimeters) and tooth position of the most complete Carcharocles megalodon teeth studied in this work. Abbreviations: HR, height of the root; MHW, maximum height of the crown; MRT, maximum thickness of the crown in cross section; MTC, maximum thickness of the crown in cross section; MWC, maximum width of the crown; TP, position on tooth set. Specimen

TP

MHC

MWC

MTC

MTR

HR

MUPAL 2730 MUPAL 2568 REG 1637 PJ 9 MUPAL 2571

ii LL1 LL3 or LL4 UL4 or UL5

90 40 28 31

50 ≈40 32–35 ≈31

11 9 7 12.4

16 13 9 12.5

19–20 – 11–12 12.4

uniformly sized and spaced (Nyberg et al., 2006, p. 812–814, Figs. 8–9). In contrast, this analysis also revealed that teeth of Carcharodon carcharias and relatives have carinae armored with triangular and pointed serrations distributed with a comparative irregular order. In all the specimens described here with a complete crown, this is triangular and single cusped, somewhat flat, and with carinae bearing a line of small rounded serration uniformly sized and distributed; these characteristics support their taxonomical identification as Carcharocles megalodon. In MUPAL 3291 is a left second anterior lower tooth or “ii” because its root notch is obtuse (≈118°), the crown is equilateral triangular, and the tip inward curved. The position of REG 1637 PJ 9 in the left or right side of the jaw is unknown; however, its equilateral triangular crown with both carinae slightly concave suggests that this is a first lower lateral tooth or “LL1”. In 120 and MUPAL 2568 teeth the crown is triangular, slightly wider than high, and straight lateral edges; characteristics that suggest they are third or fourth lower lateral tooth (“LL3” or “LL4”). REG 1641 PJ1shows the typical heart-shaped crown of the fourth and fifth upper lateral tooth of Carcharocles megalodon, “UL4” or “UL5” (Purdy et al., 2001, fig. 391). Other teeth referred here are so incomplete and fragmented that their position within the tooth set are unidentifiable (Fig. 3). According to different authors (Adnet et al., 2010; Pimiento and Clements, 2014; among others), Carcharocles megalodon lived during a little more than thirteen million years between the middle Miocene (15.9 my) and late Pliocene (2.6 my). Into the geological environment of Palenque, teeth attributable to this species have been collected in the marine Miocene deposits of the Tulijá Formation exposed in the Río Chacamax (IGM-loc 3880) and Chancalá (IGM-loc 3880) (Fig. 1, Table 1).

3.2.1. The case of lost “Odantaspis” and the seven “elamobrach vertebrae” Some of the fossil fish elements (L1-L8 in Table 1), previously recovered in Palenque and identified by Ruz-Lhuillier (1958a), seem to be lost. Therefore, it is unable to carry out a suitable description of these

3.2. The lemon shark tooth The lemon sharks are formally grouped into the genus Negaprion Whitley, 1940. The specimen MUPAL 2569 clearly belongs to this genus 471

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(Table 1), previously reported Ruz-Lhuillier (1958a, Figs. 4h-h and i, lam. XVIII a and b) and García-Moll, (2007, lam. XVIII a and b) share the characteristics already described above. The anterior observation is also applicable to the large number of recent or non-fossil stinging tail spines also found into the archeological context of Palenque, which in some cases were burned down up to the complete carbonization. It is not uncommon for vertebrate fossils to be strongly impregnated with iron minerals; a natural process that causes significant color changes in the fossils since clear-bone color toward the reddish, brown, and in extreme cases different shades of black, as it is the case of the specimen REG 1637 PJ2 (Fig. 8). Nonetheless, the nonfossil and fossil nature of the archeological artifacts made of bone found in this project is easily verifiable; recent specimens partially or totally carbonized usually have a grainy texture and are much lighter; in contrast, the fossil specimens are crystalline and heavier. Finally, the specimen MUPAL 3292, reported by Ruz-Luihiller (1958c: 208, Fig. 14-v, lám. XLVIII-m) as a “fossil stingray spine” from the Temple of the Count, Palenque, is not a fossil remain. This issue is a representation of a stingray spine carving on bone, perhaps of a recent mammal. The myliobatiform tail spines are relatively fragile fossils, difficult to collect in large fragments or complete samples. The presence of almost complete specimens of this kind in Palenque and the El Lacandón sites reveals a significant collection effort by the Maya people. Although in this project, these fossils were not collected; their possible presence into Miocene sediments of the Tulijá Formation is evidenced by the presence of myliobatiform teeth recovered in the Río Chacamax (IGMloc 3880) and Chancalá (IGM-loc 3880) sites (Fig. 1, Table 1).

fossils; however, published data and illustrations allow us to provide additional commentaries (see García-Moll, 2007, p. 262, lam. XVIII). The single tooth of these fossils (L1 in Table 1) do not have a useful metric scale and shows only one of its surfaces. Unfortunately, RhuzLhuillier (1958a, p. 79) had a misspelling when he reported these fossils as part of the genus “Odantaspis”; later, such a mistake was repeated by other authors (Polaco and Guzmán, 1997; García-Moll, 2007; CuevasGarcía, 2008). The correct name of this genus erected by Agassiz (1838) is Odontaspis. Although Llamas (in Ruz-Lhuillier, 1958a) correctly identified this tooth as O. cuspidate; a briefly review of the nomenclatural history of this sand-tiger shark indicates that its proper name is Carcharias cuspidata (Agassiz, 1843). Fisrtly named as Lamna cuspidata (Agassiz, 1843), repeatedly this species has been assigned to other genera; consequently, and its name has been repeatedly changed due the nomenclatural combinations. In fact, this species has already been named as Odontaspis cuspidata (Eastman, 1901), Lamna cuspidata (Hay, 1902), Carcharias (Odontaspis) cuspidata (Zbyszewski and de Almeida, 1950), Odontaspis (Synodontaspis) cuspidata (Schultz, 1968), Synodontaspis cuspidata (Cappetta, 1987), and Carcharias cuspidata (Purdy et al., 2001). The available illustrations of those seven vertebrae referred as “Tertiary fossil elasmobranch vertebrae” (Ruz-Lhuillier, 1958a, p. 79, lam. XVIII; García-Moll, 2007, p. 262, lam. XVIII), do not show enough features to confirm or reject their taxonomical identity nor their fossil nature (L2-L8 in Table 1). 3.2.2. Myliobatiforms stinging tail spines Fossils MUPAL 2578 and MUPAL 2579 recovered in Palenque and respectively identified as spines of “Dasyatis” and “Eagleray”, as well as REG 1638 PJ 7 and REG 1638 PJ 1 from the El Lacandón and El Lacandón, and labeled in the field as “stingray spine” and “shell ring/ shark tooth”, represent stinging tail spines of cartilaginous fishes belonging to superorder Batoidea, commonly known as rays (Table 1, Fig. 5). Unfortunately, six similar specimens from Palenque are lost (L 10 to L14, and L 16 in Table 1). Among batoidea, members of three families belonging to the order Myliobatiformes (Dasyatidae, Rhinopterydae and Myliobatidae) bear one or more stinging spines on the dorsal surface of tail. The precise specific identification of the fossils referred to above requires a deep analysis. Although Schwartz (2005, 2007, 2008) claimed that the stingray tail spines of rays can be used to identified and even describe new species; today, there are no conclusive characteristics that can be applied in the generic or specific recognition of the rays based on isolated or incomplete tail spines (Purdy et al., 2001; Laurito and Valerio, 2008; Cuny and Piyapong, 2007). Therefore, in this work it is suggested that all stingray spines described in the manuscript must be simply referred as myliobatiform caudal spines. None of the fossil spines mentioned above is complete; however, they have common characteristics suggesting their belonging to the same myliobatiform group. These are long flat and stout rod structures divided in two sections. In this spine, the basal and toothless section tapers in lateral view, is slightly wider in dorsoventral view, and has somewhat convex lateral edges. In contrast, the serrated section of this spine is almost uniformly thick in lateral view, uniformly wide in dorsoventral view except for its tapered terminal third, and its lateral edges have a row of serrations uniformly shaped and distributed that decrease in size towards both ends of the spine (in the middle part of REG 1638 PJ 7 these serrations are about 1 mm in length). In these spines both dorsal and ventral surfaces are smooth with the longitudinal axis slightly protruding; the dorsal surface shows a single middle longitudinal shallow groove along and the ventral surface displays a couple of deep lateral grooves running next to serrated lateral edges. Among these fossils, REG 1638 PJ 7 is almost complete of 87 mm in length. Other specimens are smaller or fragmented that their lengths are imprecise (REG 1638 PJ is about 47 mm, MUPAL 2578 is about 45 mm, and MUPAL 2579 is about 77 mm). The specimens L10 and L11

3.2.3. Pycnodus The fossils referred in this section are preserved on the surfaces of the slabs MUPAL 2587 and MUPAL 2731 recovered in Palenque (Figs. 6, 7; Table 1). These slabs show numerous specimens of different fish groups; therefore, to facilitate their identification in this and the following manuscript sections, each of these specimens is identified with a particular number, clearly marked in the Figs. 6 and 7. Numbers 1 and 2 in MUPAL 2587 plus 1 to 7 in MUPAL 2731, represent complete or partial specimens of the genus Pycnodus Agassiz, 1833, actually allocated within the order Pycnodontiformes and family Pycnodontidae. The massive mortality of Pycnodus is shown for the first time in MUPAL 2587 and MUPAL 2731; hence, by itself this finding deserves an accurate study that goes beyond the scope of this work. These fishes display similar morphological characteristics, including one or more of the diagnostic features, suggesting their belonging to the same species of Pycnodus. These characteristics include the presence of winged ribs, massive teeth arranged in rows on vomer and prearticular, modified stout scales along the pre-dorsal and pre-anal edges of the body, which are diagnostic for the order Pycnontiformes (e.g. Poyato-Ariza and Wenz, 2002). These have the piniculus-like process in the parietal, which is a distinctive feature of the family Pycnodontidae (Machado and Brito, 2006) as well as the bifid stout scale above the cloacal opening, the incompletely ossified or reduced rod-like scales in the dorsal and abdominal areas of the trunk, and the presence of only 10 to 15 scales along the pre-anal body edge that characterize the subfamily Pycnodontinae (Poyato-Ariza, 2013). The genus Pycnodus differs from other pycnodontins because the mentioned bifid cloacal scale is not bordered by modified comma-shaped scales and its caudal peduncle is comparatively short or bad differentiated (there are less than three neural and haemal spines between the beginning of caudal endoskeleton and the end of anal and dorsal fins), and exhibit a welldefined dermocranial fenestra; these characters are also present in the specimens from Palenque referred here. Pycnodus groups numerous Cretaceous-Eocene nominal species described based mainly on extremely incomplete specimens; hemce, Poyato-Ariza (2013) restricted the genus to only one species, P. apodus Agassiz, 1833, from the Eocene deposits of Monte Bolca. Alvarado472

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pectoral girdle.

Ortega et al. (2015) reported the occurrence of Pycnodus sp. recovered from both Belisario Dominguez (IGM-loc 3870) and División de Norte (IGM-loc 3870) quarries belonging to the Paleocene marine sediments of the T-LU, which also are present around and constitute the basement of Palenque. During this research a dentary bone of Pycnodus was collected near the Building XXVI of Palenque (Fig. 1 and Table 1).

3.2.6. Other fishes Other fossils preserved on the slabs MUPAL 2587 and MUPAL 2731 cannot be confidently assigned to any taxonomic fish group (Figs. 6, 7). The specimens 4 to 7 on MUPAL 2587 and 18 to 23 on MUPAL 2731 are elongated fishes, in which the dorsal and pelvic fins are opposed to each other, the anal fin is placed farther back, and the snout is reduced. These characteristics are similar with those of the teleostan order gonorynchiformes. Unfortunately, the poor preservation of these specimens and the impossibility to prepare these archeological remains preclude the verification of this observation. Complete gonorynchiform specimens have already been collected in the Paleocene marine deposits of Belisario Domínguez and División del Norte quarries (Fig. 1, Table 2); however, at this moment nothing can be added beyond the present observations because these fossils are under study. Finally, other fish remains of uncertain taxonomic affinity are numbered as 8 in MUPAL 2587 and 24 to 26 in MUPAL 2731.

3.2.4. Kelemejtubus castroi The perciformes fishes numbered as 3 in MUPAL 2587 and 8 to 15 in MUPAL 2731 represent the species Kelemejtubus castroi Cantalice and Alvarado-Ortega, 2017 (Figs. 6, 7), recently described based on specimens from the Paleocene deposits of the Tenejapa-Lacandón Unit exploted in the Belisario Domínguez and the Northern Division quarries (Fig. 1, Table 2). A remarkable feature of this elongated percomorph species is the longest maxillary and the presence of stout conical teeth on the jaws, with a well-developed canine on tip of the dentary. Other features, such as the absence of supramaxilla, two spines on the posterior portion of the preopercle, and two separated lobes on the dorsal fin, are also observed in the specimens referred here.

3.3. Anthropogenic alterations of fossils

3.2.5. Serranid perciform Fish specimens numbered as 9 in MUPAL 2587 and 17 in MUPAL 2731 are identified here as “serranids” (Figs. 6, 7). Swainson (1839) named the family Serranidae to include the genus Serranus Cuvier, 1817, and related modern “groupers” and “sea basses”; however, for a long time the family was a “taxonomic wastebasket” (Johnson, 1983, Adolfssen et al., 2017). Serranids have a poor understood history that probably goes back beyond 65 mybp (Betancur-R et al., 2013). Although the family now is well defined and nested into the order Perciformes; its composition and naturalness has been repeatedly questioned because the extreme high diversity of the family entails some problems of their relations and taxonomy that have not yet been solved (see Gosline, 1966; Wiley and Johnson, 2010; Smith and Craig, 2007; Ma et al., 2016; among others). The specimen, IGM 4550, is a serranid fish from the marine Paleocene sediments of the T-LU exposed in the Division del Norte quarry (IGM-loc 3869) identified by Alvarado-Ortega et al. (2015). Today a larger number of better preserved specimens from this quarry and its coeval Belisario Domínguez quarry (IGM-loc 3870) are part of an accurate morphological and taxonomic study, carried out by the authors of this work. Although the most conspicuous features of the serranid group (opercle with three posterior spines and preopercle serrated) are not exposed in any of the fossils from Palenque preserved on MUPAL 2587 and MUPAL 2731; these share other morphological characteristics with IGM 4550 and other serranids already identified from Division of the North and Belisario Domínguez quarries; these characteristics suggest that all these fossils belong to the same species. These fossils share the body shape (the body is somewhat chubby, relatively short and high, and its maximum body depth is about 40% of the standard length (SL)); proportion of head and body (the head represents about 40% of SL); the composition, distribution and shape of paired and unpaired fins (the pelvic girdle is located just below and behind the pectoral girdle, the pectoral fin consists of eleven soft rays, the pelvic fin includes one stout spine plus five soft rays; the long dorsal fin has two continuous lobes; the anterior dorsal fin lobe is formed by eight thick thorns while the posterior lobe of this fin has one thick spine plus 11 soft rays; the anal fin ray is relatively short and includes three stout spines and probably only seven soft ray); as well as the number of vertebrae (24 including10 abdominals and 14 caudals), and the number and distribution of predorsal bones (= predorsal formula (0/0/0 + 2/1 + 1/). These fossils also have the diagnostic characteristics noted by different authors of the order Perciformes (Patterson, 1964; Wiley and Johnson, 2010), including a pelvic fin consisting of one spine and five soft rays and located close to the pectoral girdle, as well as pelvic bones in contact with the

A mandatory indication in the INAH protocols pointed out to us at the beginning of this study, which is applicable in the collection and conservation of archeological objects, is that not one of the archeological objects (including fossils) is be submitted, without prior authorization, to preparation or preservation procedures that could alter its form and composition. Therefore, since there is no record of alteration of any of the fossils studied here, we consider that all their characteristics were acquired in the prehispanic time. In addition to their biological features, some of these fossils show unnatural alterations, which deserve special attention due their possible anthropogenic nature (Figs. 8–10). In this framework, such alterations are considered potential keys to a better understand of the Mayas thought about the fossils. Three differ alteration kinds are identified in this study, including the wear marks in cutting edges, paint stains and plaster patches, as well as preparation marks. 3.3.1. Wear marks in cutting edges It is noteworthy that some of the cutting edges in a shark tooth and a sting ray spine referred show areas clearly altered, in which the edges are concave and the serrations are eroded (Fig. 8). These alterations are present in both carinae of the shark tooth MUPAL 2730 that also has two deep triangular notches closely placed near to the crown tip probably produced by hitting the same object twice. These morphological alterations on MUPAL 2730 differ from those dental pathologies found in fossil and living sharks that include deformations in crown and root (notches in the carinae, crowns surfaces soft or strongly twisted, enamel cracked, and excess of dentine) (Hubbell, 1996; Purdy et al., 2001, among others). The right lateral edge of the sting ray spine MUPAL 2578 also shows unnatural alterations; its profile is concave, and its serrations are eroded (Fig. 8). The use of different teeth of species of fossil and living shark as tools, ornaments, and weapons by ancient human groups is rare but have been documented along America (Holmo, 1969; Cione and Bonomo, 2003); usually, in these teeth the roots are carved to make possible to tie them in the tip of long objects and get spearheads or arrows, or are perforated to include them in necklaces (López and Sánchez García, 2004; González, 2005; Lowery et al., 2011; Haines et al., 2008; Moyer and Bemis, 2017). Shark teeth and sting ray spines from Palenque referred here do not show these craft changes; however, the localized disturbance and the concave smooth altered profiles of these fossils clearly represent artificial features. These alterations represent abrasion marks as those generated on the edges of cutting tools (serrated knives or saws) after the extreme wear, whose mechanics of use follows a reciprocating movement (Diniz and de Oliveira, 2008). 473

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García, 2008). Although some of the fossils described here were found in the socalled garbage dumps, outside the most striking archeological contexts such as offerings and tombs; this observation reveals that these fossils might have some utility and were discarded as they deteriorated or ruptured. Other fossils were directly collected from primary archeological contexts, in burials and offerings, which denote they had a special meaning and usefulness. Here, the noticeable wear marks on fossilized shark teeth and the spines of rays demonstrate their used as cutting artifacts (Figs. 8 and 9). Based on the distribution of colorful stains on the slab MUPAL 2587, it is impossible to determine if these rocks were part of an elaborated pictorial design; however, this finding points to the effort to highlight this object. The carved surfaces around the fossils and some high reliefs on slabs studied here (Fig. 9) also show the interest to discover their skeletons. The present findings and observations lead us to recognize the knowledge and practice of recovering fossils in the immediate geological environment that the Mayas of Palenque had; their (archaic) attempts for preparing and discovering them are evidenced; as well as the anthropogenic features preserved in these fossils revealing their utility and inclusion in ornamental and ritual activities. Cuevas-García (2007, 2008) pointed out a relevant fact that deserves further investigations, the Maya gods represented in the incense burners of Palenque are represented with large shark teeth, fins or fish gills next to their mouths, as well as marine shells or bivalves as earmuffs. Although it is difficult to determine what kind of shark teeth the Maya attempted to represent in these burners; such representations show large size tri-cuspids teeth, with a large central cusp, resembling teeth of some megatooth sharks. Undoubtedly, these sculptures may be indicators of the importance of fossils in the Maya religious thinking.

3.3.2. Preparation marks Fig. 9 shows an orderly array of uniformly wide scratches on the surfaces of both bearing fishes slabs MUPAL 2731 and MUPAL 2587. These marks are in parallel and around the osseous structures of the fossils; these seem to be the result of the same mechanical preparation procedures applied in the modern paleontology. First, the trained fossil preparators remove the sediments around the fossilized structures with the intention of recognize their general shape and extension; this step is done with rough or general purposes tools (such as chisels or strong needles). Posteriorly, the fine details of the fossils are discovered removing the superficial sediments with finer tools (such as fine needles used below microscope). Although these slabs do not show marks of fine preparation; these scratches demonstrate the intentional effort to discover their fossils. Same marks are also present on some of the protruding areas of these rocks showing that “fossil preparator” was looking for additional fossils covered in the slabs. 3.3.3. Color stains and plaster patches In pre-Hispanic Mesoamerica the use of colors had a significant religious connotation; thus, blue is linked to the celestial diurnal sphere and the aquatic world while red is related to the blood and sacrificial acts (Nicholson, 1985; Gastélum, 2006). Curiously, corners of the slab MUPAL 2587 show some small stains of these two colors (Fig. 10). Unfortunately, these colorful spots are so scarce and scattered that it is not possible to recognize if they were part of a decorative frame or a more elaborate pictorial representation; hence, there is no opportunity to link this rock and its fossils with any ideological reference of Palenque. The same fossil bearing slab shows another notable feature, it has large spots of plaster (Fig. 10). Given its porous and irregular appearance, it is difficult to interpret the occurrence of this plaster; in more usual cases, such as when the plaster cements rocky blocks (as in the walls) or is used to fill or produce superficial fine details (as in the sculptures or high reliefs coated with plaster), this material shows a strong compaction or forms well-defined overlapping layers, which in both cases implies the loss of porosity.

Acknowledgements We are in debt with the institutions and people that made this research possible. We owe our gratitude to Alberto Montejo and Nicolás Hurtado and their families, as well as Miguel Demeza, Ulises and Maritza, for their support during the field works. Our colleagues and students, A. Cristin, A. Alaniz, M. del P. Melgarejo, M. J. Nuñez, G. Solano, E. Valsequillo, C. Nuñez, and Y. M. Alves, help us along this research. L.P. Machado and J.M. Contreras help us with the photos included in this work. Financial supported of this research was provided by the INAH (Grant 2008-2009) and the UNAM (grants DGAPA-PAPIIT IN106011, IN207314, and IN209017).

4. Discussion and conclusion The presence of fossils, mainly shark teeth, in the prehispanic archeological contexts of America is not a new subject; however, Borhegyi (1961) pointed out the lack of clues to explain the meaning and function, as well as to recognize the sites and extraction procedures, of fossil and recent stingray spines and shark teeth during the pre-Hispanic period in Mexico and other parts of America (Guatemala, Costa Rica, and Panama). This research provides evidences to recognize that, at least, in the case of the Palenque area, the Maya people recovered fossils from the nearby geological context. The fossil fish remains found within the primary and secondary archeological contexts of this ancient city correspond to different geological temporalities. The results of field prospection performed in this research show that the Palenque environment has geological units bearing or potentially carrying all the species represented by these fossils. The pycnodonts, serranids, percomorphs, and other fishes preserved in the slabs MUPAL 2587 and MUPAL 2731 came from the Peleocene sediments of the T-LU while fossils of sting rays and sharks come from the Miocene coastal marine deposits of the Tulijá Formation. In the Caribbean archeological records there is a basic difference in the use of shark teeth (Borhegyi, 1961; Maxwell, 2000). Small perforations were made in these teeth to use them as pendants elements of necklaces, earrings or bracelets; others, had to have other utility as cutting tools or weapons. Evidences so far accumulated suggest that in the central and Maya areas of Mexico, shark teeth were deposited at ritual contexts, as in funeral and votive offerings, after their use as punches for self-sacrifice, piercing the tongue, ear lobes, nose or penis, and probably once used in ceremonies related with these acts (Cuevas-

References Acosta, J.R., 1976. Exploraciones en Palenque durante 1972. Anales del Instituto Nacional de Antropología e. Historia 7 (5), 5–42. Adnet, S., Balbino, A.C., Antunes, M.T., Marin-Ferrer, J.M., 2010. New fossil teeth of the white shark (Carcharodon carcharias) from the early Pliocene of Spain. Implication for its paleoecology in the Mediterranean. Neues Jahrb. Geol. Palaontol. Abh. 256 (1), 7–16. Adolfssen, J.S., Milan, J., Friedman, M., 2017. Review of the Danian vertebrate fauna of southern Scandinavia. Bull. Geol. Soc. Den. 65 (1), 1–23. Agassiz, L., 1833–1843. Recherches sur les Poissons fossiles. 5 Neuchâtel, Petitpierre. Alvarado-Ortega, J., Cuevas-García, M., Melgarejo-Damián, M. del P., Cantalice, K.M., Alaniz-Galvan, A., Solano-Templos, G., Than-Marchese, B.A., 2015. Paleocene fishes from Palenque, Chiapas, southeastern Mexico. Palaeontol. Electron. 18 (2), 1–22. Applegate, S.P., Espinosa-Arrubarrena, L., 1996. The fossil history of Carcharodon and its possible ancestor, Cretolamna: a study in tooth identification. In: Klimley, A.P., Ainley, D.G. (Eds.), Great White Sharks: The Biology of Carcharodon carcharias. Academic Press, San Diego, California, pp. 19–36. Aranda-Manteca, F.J., 1994. Vertebrados marinos fósiles de la Misión, Baja California, México. Master degree Thesis. Universidad Autónoma de Baja California, México. Betancur-R, R., Broughton, R.E., Wiley, E.O., Carpenter, K., López, J.A., Li, C., Holcroft, N.I., Arcila, D., Sanciangco, M., Cureton, J.C., Zhang, F., Buser, T., Campbell, M.A., Ballesteros, J.A., Roa-Varon, A., Willis, S., Borden, W.C., Rowley, T., Reneau, P.C., Lu, G., Grande, T., Arratia, G., 2013. The tree of life and a new classification of bony fishes. In: PLOS Currents Tree of Life. Blake, C.C., 1862. Sharks' teeth at Panama. In: The Geologist. 5. pp. 316. Borhegyi, S.F., 1961. Shark teeth, stringray spines, and shark fishing in ancient Mexico

474

Journal of Archaeological Science: Reports 17 (2018) 462–476

J. Alvarado-Ortega et al.

of groupers: clade diversification patterns and processes. Mol. Phylogenet. Evol. 100, 21–30. Machado, L.P.C., Brito, P.M., 2006. The new genus Potiguara (Actinopterygii: Pycnodontiformes) from the upper cretaceous of northeast Brazil. J. Vertebr. Paleontol. 26 (1), 1–6. Martínez-Ortiz, A.C., Alvarado-Ortega, J., Cuevas-García, M., 2017. Ocurrencia de un decápodo braquiuro extinto, Necronectes proavitus (Rathbun, 1918), en los yacimientos marinos de la Formación Tulijá (Mioceno temprano) en las cercanías de Palenque, Chiapas, sureste de México. Paleontología Mexicana 6 (1), 1–15. Maxwell, D., 2000. Beyond maritime symbolism. Anc. Mesoam. 11 (1), 91–98. Meneses-Rocha, J.J., 2001. Tectonic evolution of the Ixtapa graben, an example of a Strike-slip Basin of southeastern Mexico: implications for regional petroleum systems. In: Bartolini, C., Buffler, R.T., Cantu-Chapa, A. (Eds.), The Western Gulf of Mexico Basin: tectonics, Sedimentary Basins, and Petroleum Systems. Memoir 75. The American Association of Petroleum Geologists, pp. 183–216. Minch, A.J., Schulte, C.K., Hofman, G., 1979. A middle Miocene age for the Rosarito Beach formation in northweastern Baja California, Mexico. Geol. Soc. Am. Bull. 81, 3149–3154. Moyer, J.K., Bemis, W.E., 2017. Shark teeth as edged weapons: serrated teeth of three species of selachians. Zoology 120, 101–109. Muizon, C., de Vries, T.J., 1985. Geology and paleontology of late Cenozoic marine deposits in the Sacaco area (Peru). Geol. Rundsch. 74, 547–563. Mulleried, F.K.G., 1951. Algunos fósiles marinos del Terciario inferior y medio de Palenque, Chiapas. In: Revista de la Sociedad Mexicana de Historia Natural XII. 1-4. pp. 209–227. Nicholson, H.B., 1985. Polychrome on Aztec sculpture. In: Boone, E.H. (Ed.), Painted Architecture and Polychrome Monumental Sculpture in Mesoamerica. Dumbarton Oaks, Washington DC, pp. 145–171. Nyberg, K.G., Ciampaglio, C.N., Wray, G.A., 2006. Tracing the ancestry of the great white shark, Carcharodon carcharias, using morphometric analyses of fossil teeth. J. Vertebr. Paleontol. 26 (4), 806–814. Olvera-Carrasco, M.T., 1997. In: Arroyo-Cabrales, J., Polaco, O. (Eds.), La arqueoictiofauna de Palenque, Chiapas, México. Homenaje al professor Ticul Álvarez, Colección Científica, INAH, México, pp. 253–278 (Coords.). Patterson, C., 1964. A review of Mesozoic acanthopterygian fishes, with special reference to those of the English chalk. Philos. Trans. R. Soc. Lond. Ser. B Biol. Sci. 213–482. Pérez-Martínez, M.B., López-Palomino, I., 2011. Lomut, Formación. Léxico Estrátigráfico Mexicano, Servicio Geológico Mexicano. pp. 3. Pimiento, C., Clements, C.F., 2014. When did Carcharocles megalodon become extinct? A new analysis of the fossil record. PLoS One 9 (10), e111086. Poey, F., 1868. Synopsis piscium cubensium. Catálogo razonado de los peces de la Isla de Cuba. Repertorio Físico-Natural de la Isla de Cuba. 2. pp. 279–484. Polaco, O.J., Guzmán, A.F., 1997. Arqueoictiofauna Mexicana. Serie Arqueología. INAH, México, pp. 99. Poyato-Ariza, F.J., 2013. Sylvienodus, a new replacement genus for the cretaceous pycnodontiform fish “Pycnodus” laveirensis. C.R. Palevol 12 (2), 91–100. Poyato-Ariza, F.J., Wenz, S., 2002. A new insight into pycnodontiform fishes. Geodiversitas 24 (1), 139–248. Purdy, R.W., Schneider, V.P., Applegate, S.P., McLellan, J.H., Meyer, R.L., Slaughter, B.H., 2001. The Neogene sharks, rays, and bony fishes from Lee Creek mine, aurora, North Carolina. Smithson. Contrib. Paleobiol. 90, 71–202. Quezada-Muñetón, J.M., 1987. El Cretácico medio-superior, y el límite Cretácico superior-Terciario inferior en la Sierra de Chiapas. Boletín de la Asociación Mexicana de Geólogos Petroleros 39 (1), 1–98. Rathbun, M.J., 1918. Decapod crustaceans from the Panama region. In: United States National Museum Bulletin. 103. pp. 123–184. Riquelme, F., Alvarado-Ortega, J., Cuevas-García, M., Ruvalcaba-Sil, J.L., Linares-López, C., 2012. Calcareous fossil inclusions and rock-source of Mayan lime plaster from the temple of the inscriptions, Palenque, Mexico. J. Archaeol. Sci. 39, 624–639. Riquelme, F., Northrup, P., Ruvalcaba-Sil, J.L., Stojanoff, V., Siddons, D.P., AlvaradoOrtega, J., 2014. Insights into molecular chemistry of Chiapas amber using infraredlight microscopy, PIXE/RBS, and Sulfur K-edge XANES spectroscopy. J. Appl.Phy. A 116, 97–109. Rüppell, E., 1837. Newe Wirbelthiere zu der Fauna von Abyssinien gehörig. (Fische des). Ruz-Lhuillier, A., 1958a. Exploraciones arqueológicas en Palenque: 1953. Anales del INAH 10 (39), 69–116. Ruz-Lhuillier, A., 1958b. Exploraciones arqueológicas en Palenque: 1954. Anales del INAH 10 (39), 117–184. Ruz-Lhuillier, A., 1958c. Exploraciones arqueológicas en Palenque: 1955. Anales del INAH 10 (39), 185–240. Ruz-Lhuillier, A., 1958d. Exploraciones arqueológicas en Palenque: 1956. Anales del INAH 10 (39), 241–299. Ruz-Lhuillier, A., 1959. Guía oficial de Palenque. Instituto Nacional de Antropología e História, México, pp. 70. Sacco, F., 1897. Pelecypoda (Ostreidae, Anomiidae e Dimyidae) of L. Bellardi & Federico Sacco, 1872–1904. I moluschi dei terreni Terziarii de Piemonte e dela Liguria. Carlo Clausen. Torino, pp. 102. Sáenz-Pita, M.R., López-Palomino, I., 2011. Tulijá. Formación. Léxico Estrátigráfico Mexicano, Servicio Geológico Mexicano, pp. 3. Sánchez-Villagra, M.R., Aguilera, O.A., 2006. Neogene vertebrates from Urumaco, Falcón state, Venezuela: diversity and significance. J. Syst. Palaeontol. 4, 213–220. Sapper, K., 1896. Sobre la geografía física y la geología de la península de Yucatán: Instituto Geologico de México, Boletín. 3 (57 p). Schultz, O., 1968. Die Selachierfauna (Pisces, Elasmobranchii) aus den Phosphoritsanden (Unter-Miozän) von Plesching bei Linz. Oberösterreich. Naturkundl. Jb. Stadt Linz (Linz) 14, 61–102.

and central America. Southwest. J. Anthropol. 17 (3), 273–296. Cantalice, K.M., Alvarado-Ortega, J., 2016. Eekaulostomus cuevasae gen. and sp. nov., an ancient armored trumpetfish (Aulostomoidea) from Danian (Paleocene) marine deposits of Belisario Domínguez, Chiapas, southeastern Mexico. Palaeontol. Electron. 18 (3), 1–24. Cantalice, K.M., Alvarado-Ortega, J., 2017. Kelemejtubus castroi gen. et sp. nov., an ancient percomorph (Teleostei, Actinopterygii) from the Paleocene marine deposits near Palenque, Chiapas, southeastern Mexico. J. Vertebr. Paleontol. 37, 12 e1383265. Cappetta, H., 1987. Handbook of Paleoichthyology. Chondrichthys II: Mesozoic and Cenozoic Elasmobranchii. Gustav Fischer Verlag, Stuttgart and New York. Casier, E., 1960. Note sur la collection des poissons Paléocènes et Eocènes de L'Enclaver e Cabinda (Congo). Annales du Musée Royal du Congo Belge A 3 (1, 2), 1–48. Castañeda-Posadas, C., Cevallos-Ferriz, S.R., 2007. Swietenia (Meliaceae) flower in Late Oligocene–Early Miocene amber from Simojovel de Allende, Chiapas, Mexico. Am. J. Bot. 94 (11), 1821–1827. Cione, A.L., Bonomo, M., 2003. Great white shark teeth used as pendants and possible tools by early-middle Holocene terrestrial mammal hunter-gatherers in the eastern pampas (southern South America). Int. J. Osteoarchaeol. 13 (4), 222–231. Cuevas García, M., Alvarado Ortega, J., 2012. El mar de la creación primordial: un escenario mítico y geológico en Palenque. Arqueología Mexicana 113, 32–37. Cuevas-García, M., 2008. Paisaje Paleontológico en Palenque. In: Laporte, J.P., Arroyo, B., Mejía, H.E. (Eds.), 2008. XXI Simposio de Investigaciones Arqueológicas en Guatemala Memoirs. 1. pp. 483–490. Cuevas-García, M., Alvarado-Ortega, J., 2009. Estudio Arqueológico y Paleontológico de los Fósiles marinos que proceden del sitio de Palenque, Chiapas 1st Field-Season Report (2008). INAH, México, pp. 54. Cuny, G., Piyapong, C., 2007. Tail spine sharacteristics of stingrays (order Myliobatiformes): a comment to Schwartz (2005). Electr. J. Ichth. 1, 15–17. Cuvier, G., 1817. Le règne animal. t.1. Deterville, Paris. Dengo, G., 1968. Estructura geológica, historia tectónica y morfológica de América Central. In: Centro Regional de Ayuda Técnica. Agencia para el Desarrollo Internacional (AID), México. Diniz, A.E., de Oliveira, A.J., 2008. Hard turning of interrupted surfaces using CBN tools. J. Mater. Process. Technol. 195 (1), 275–281. Eastman, C.R., 1901. Pisces. In: Clark, W.B., Martin, G.C. (Eds.), The Eocene Deposits of Maryland. Maryland Geological Survey, Baltimore, pp. 98–115. Fernández, M.A., 1991. Las ofrendas del Templo del Sol y de la Cruz Enramada. Temporada de trabajos en la zona arqueológica de Palenque, Chiapas, del 25 de mayo al 10 de septiembre de 1942. In: García-Moll, R. (Ed.), En Palenque 1926–1945. Antologías, Serie Arqueología, INAH, México, pp. 239–298. García-Cuevas, M., 2007. Los incensarios efigie de Palenque: Deidades y rituales mayas (Vol. 1). Serie Testimonios y Materiales Arqueológicos para el Estudio de la Cultura Maya 1. Universidad Nacional Autónoma de México 350 pp. García-Moll, R., 2007. Palenque 1947–1958. Alberto Ruz Lhuillier. INAH, México. Gastélum, L.G., 2006. Una aproximación a la temática del color em el México antiguo. Cuicuilco 13 (36), 151–175. González, M., 2005. Use of Pristis spp. (Elasmobranchii: Pristidae) by hunter-gatherers on the coast of São Paulo, Brazil. Neotropical Ichthyology 3 (3), 421–426. González-Cruz, A., 1993. Trabajos Arqueológicos en Palenque. Chiapas. Informe de Campo VI Temporada, INAH, (México), pp. 8. Gosline, W.A., 1966. The limits of the fish family Serranidae, with notes on other lower percoids. Proc. Calif. Acad. Sci. 33, 91–111. Haines, H.R., Willink, P.W., Maxwell, D., 2008. Stingray spine use and Maya bloodletting rituals: a cautionary tale. Lat. Am. Antiq. 19 (1), 83–98. Ham-Wong, J.M., 1979. Prospecto Nazareth, Chiapas. Petróleos Mexicanos, Zona Sur. Informe Geológico 745, 8–18 (unpublished). Hay, O.P., 1902. On a collection of upper cretaceous fishes from Mount Lebanon, Syria, with descriptions of four new genera and nineteen new species. Bull. Am. Mus. Nat. Hist. 19, 395–452. Holmo, O., 1969. Dientes de tiburón. El utensilio prehistórico y un comentario folklóriko. Casa de la Cultura Ecuatoriana, Guayaquil, pp. 22. Hubbell, G., 1996. Using tooth structure to determine the evolutionary history of the white shark. In: Klimley, A.P., Ainley, D.G. (Eds.), Great White Sharks: The Biology of Carcharodon carcharias. Academic Press, New York, pp. 9–18. Johnson, G.D., 1983. Niphon spinosus: a primitive epinepheline serranid, with comments on the monophyly and intrarelationships of the Serranidae. Copeia 1983, 777–787. Jordan, D.S., Hannibal, H., 1923. Fossil sharks and says of the Pacific slope of North America. Bull. South. Calif. Acad. Sci. 22, 27–63. Laurito, C.A., Valerio, A.L., 2008. Ictiofauna de la localidad de San Gerardo de Limoncito, Formación Curré, Mioceno Superior, cantón de Coto Brus, provincia de Puntarenas, Costa Rica. Revista Geológica de América Central 39, 65–85. Leidy, J., 1873. Notice of remains of fishes in the bridger tertiary formation of Wyoming. Proc. Acad. Natl. Sci. Phila. 25, 97–99. Linnaeus, C.V., 1758. Systema Naturae, 10th Ed. 1 L. Salvii, Stockholm. López, O.R., 1964. Informe geológico del área SW del alto Tulijá, Chiapas. Petróleos Mexicanos, Zona Sur. Informe Geológico 474 (unpublished). López Bravo, R., López Mejía, J., Venegas Durán, B., 2003. Entre el Motiepá y el Picota: la primera temporada del Proyecto Crecimiento Urbano de la antigua ciudad de Palenque. Lakamha 2 (9), 10–15. López, A.R., Sánchez García, L.C., 2004. Costumbres funerarias en el Conchalito, La Paz, Baja California Sur. Anales de Antropología 38, 139–178. Lowery, D., Godfrey, S.J., Eshelman, R., 2011. Integrated geology, paleontology, and archaeology: native American use of fossil shark teeth in the Chesapeake Bay region. Archaeol. East. N. Am. 39, 93–108. Ma, K.Y., Craig, M.T., Choat, J.H., van Herwerden, L., 2016. The historical biogeography

475

Journal of Archaeological Science: Reports 17 (2018) 462–476

J. Alvarado-Ortega et al.

Facultad de Estudios Superiores Iztacala, Universidad Nacional Autónoma de México, Estado de México. Venegas-Duran, B.J., 2005. En busca de los orígenes de Palenque: Investigaciones recientes del proyecto Crecimiento Urbano de la Antigua ciudad de Palenque (PCU). Mayab. 18. pp. 57–67. Vinson, G.L., 1962. Upper cretaceous and tertiary stratigraphy of Guatemala. Bull. Am. Assoc. Pet. Geol. 62, 425–456. White, E.I., 1926. Eocene fishes from Nigeria. In: Bulletin of the Geological Survey of Nigeria. 10. pp. 1–82. Whitley, G.P., 1940. The fishes of Australia. Part I. The sharks, rays, devil-fish, and other primitive fishes of Australia and New Zealand. In: R. Zool. Soc. N. S. W., Austral. Zool. Handbook, pp. 1–280. Wiley, E.O., Johnson, G.D., 2010. A teleost classification based on monophyletic groups. In: Nelson, J.S., Schultze, H.-P., Wilson, M.V.H. (Eds.), Origin and phylogenetic interrelationships of Teleosts. Verlag Dr. Friedrich Pfeil, München, pp. 123–182. Williams, G.S., 1999. A listing of fossil sharks and rays of the world, Version 32 (Available). http://www.afn.org/~afn02877/neosel.htm. Zbyszewski, G., de Almeida, F.M., 1950. Os peixes miocénicos portugueses. Comunicações da Comissão do Serviço Geológico de Portugal. 21. pp. 309–412.

Schwartz, F.J., 2005. Tail spine characteristics of stingrays (order Myliobatifromes) found in the Northeast Atlantic, Mediterranean, and black seas. Electron. J. Ichthyol. 1, 1–9. Schwartz, F.J., 2007. Tail spine characteristics of stingrays (order Myliobatiformes) frequenting the FAO fishing area 61 (20°N 120 E–50°N 150 E) of the Northwest Pacific Ocean. Raffles Bull. Zool. 14, 121–130. Schwartz, F.J., 2008. A survey of tail spine characteristics of stingrays frequenting African, Arabian to Chagos-Maldive archipelago waters. Smithiana Bulletin 8, 41–52. Servicio Geológico Mexicano, 2006a. Carta Geológico-Minera Estados de Chiapas y Tabasco; escala 1:500 000. Servicio Geológico Mexicano, Secretaria de Economía. Servicio Geológico Mexicano, 2006b. Carta Geológico-Minera Tenosique E15-9; Chiapas, Tabasco y Campeche, escala 1: 150 000. Servicio Geológico Mexicano, Secretaria de Economía. Smith, W.L., Craig, M.T., 2007. Casting the percomorph net widely: the importance of broad taxonomic sampling in the search for the placement of serranid and percid fishes. Copeia 2007 (1), 35–55. Solórzano-Kraemer, M.M., 2010. Mexican amber. In: Penney, D. (Ed.), Biodiversity of Fossils in Amber from the Major World Deposits. Siri Scientific Press, pp. 42–56. Swainson, W., 1839. The natural history and classification of fishes, amphibians, & reptiles, or monocardian animals. 2. Spottiswoode & Co., London, pp. 1–448 + vi. Velasquillo-García, E., 2011. Ostras fósiles de Palenque. Chiapas. Bachelor's Thesis.

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