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Mammoths in Central America: New records from Guatemala
Quaternary International 443 (2017) 122e128
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Mammoths in Central America: New records from Guatemala vila A b, ** H. Gregory McDonald a, *, Sylvia Lorena Da a
Bureau of Land Management, Utah State Office, 400 West 200 South, Salt Lake City, UT, 84101, USA n de Fo siles, Museo Historia Natural, Universidad de San Carlos de Guatemala, Calle Mariscal Cruz 1-56, Zona 10, Ciudad de Guatemala, Coleccio Guatemala b
a r t i c l e i n f o
a b s t r a c t
Article history: Received 7 December 2015 Received in revised form 8 December 2016 Accepted 18 December 2016 Available online 13 July 2017
The remains of the Columbian Mammoth, Mammuthus columbi, are reported from four localities in Guatemala based on seven dental records. These localities are widely separated and include one spec n, Sayaxche , Departamento de Pete n in the northern part of the country, one imen from Río La Pasio specimen from Chinautla, Departamento de Guatemala in the southwest and four specimens from the n, Departamento de Zacapa in the vicinity of Estanzuela, Departamento de Zacapa and one from Teculuta southeast. Based on the known records the distribution of Mammuthus in Central America tends to be concentrated on the Pacific side of the isthmus and suggests the former presence of a savanna corridor that formed during an interval of decreased precipitation. © 2016 Published by Elsevier Ltd.
Keywords: Mammuthus columbi Mammoth Central America Guatemala Pleistocene
r e s u m e n Los primeros vestigios de Mamuts, Mammuthus columbi, se han reportado de cuatro localidades en n, Sayaxche Departamento de Guatemala, muy distantes entre ellas: uno procedente de Río La Pasio n, en la parte septentrional del país, otro de Chinautla, Departamento de Guatemala en el suroeste, y Pete n, Departamento de Zacapa, en el sureste. Estos nuevos registros cuatro de Estanzuela, y uno de Teculuta brindan un número total de siete basados en las evidencias dentales conocidas de Mammuthus de n de Mamuts, en America Central, los regristros Guatemala. Basados en el conocimiento de la distribucio se centran en la vertiente del pacifico del Itsmo, sugieriendo la presencia de un corridor seco de sabana. © 2016 Published by Elsevier Ltd.
1. Introduction While mammoths are widely distributed in North America from Canada through the United States (Agenbroad, 2005) and into Mexico (Arroyo-Cabrales et al., 2003) their record in Central America is minimal with only a few scattered records in five of the seven countries. In Mexico mammoths have been studied in more rez-Crespo et al., 2012; detail (Arroyo Cabrales et al., 2010; Pe rrez-Bedolla et al., 2016) but much less is known about Gutie them in Central America. The presence of Mammuthus in Central America was first reported by Leidy (1886) as part of a small Pleistocene fauna from
* Corresponding author. ** Corresponding author. E-mail addresses: [email protected] (H.G. McDonald), [email protected] vila A). (S.L. Da http://dx.doi.org/10.1016/j.quaint.2016.12.018 1040-6182/© 2016 Published by Elsevier Ltd.
Nicaragua and the genus has subsequently been found in Guatemala, El Salvador, Honduras, and Costa Rica (Laurito and Aguilar, 2007; Lucas and Alvarado, 2010). The recovery of additional mammoth remains in Guatemala provides a stronger geographic link between Mexico and these more southern records and improves our understanding of the southernmost portion of the genus' distribution. As there are other Pleistocene proboscideans, Cuvieronius and Mammut, present in Central America (Arroyo-Cabrales et al., 2007; Lucas and Alvarado, 2010) and Cuvieronius is known from Guatemala (Mead et al., 2012), we have only considered those records of mammoth based on dentition. We recognize that other records of mammoth may exist represented by only the post-cranial skeleton, such as the specimen from Las Banderas, Nicaragua, reported by Lucas (2004). We do not consider these here given the lack of detailed anatomical studies of differences in the post-cranial skeleton of mammoths and the other genera of proboscideans present in Central America and the uncertainty related to taxonomic identifications based on post-cranial
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specimens. The taxonomic assignment to species of specimens of Mammuthus has varied in the past and often reflected the changing opinions on the valid species of North American Mammuthus based on both morphological or molecular criteria (Lister and Sher, 2015; Enk et al., 2016). Based strictly on tooth morphology, records of Mammuthus from Central America can be referred to M. columbi, although some earlier records were referred to the species M. imperator (Gutierrez, 1963) and Lucas and Alvarado (2010) recognize a record of M. meridionalis from El Salvador. Based on the limited available stratigraphic and chronologic information available for most specimens it seems most likely that most of the Mammuthus records in Central America can be considered to be late Pleistocene in age. The new specimens from Guatemala have the meristic and morphological features currently used to diagnose Mammuthus columbi and are assigned to this species. 2. Abbreviations INGUAT e Instituto Guatemalteco de Turismo ME e Museo de Estanzuela, Estanzuela MUSHNAT e Usac e Museo de Historia Natural de la Universidad de San Carlos de Guatemala, Guatemala City MNHN e Museo Nacional de Historia Natural (Estatal). Guatemala, City While some workers use a system of identifying mammoth dentition as M/m 1e6 for the teeth to distinquish them based on the sequence of eruption, we have identified the teeth according to their inferred homology as a premolar or molar. All measurements are in millimeters. 3. Description of material Order Proboscidea Illiger, 1811 Family Elephantidae Gray, 1821 Mammuthus Brookes, 1828 Mammuthus columbi (Falconer), 1857
3.1. Guatemala The area around Estanzuela is perhaps the richest in Pleistocene vertebrates for Guatemala in terms of the number of specimens of Pleistocene fauna known from a restricted location within the country. However, despite the large number of specimens the species richness is low and limited to megafauna. Along with the mammoth, Cuvieronius hyodon, Eremotherium laurillardi, Glyptotherium cf. G. floridanus, Neochoerus sp., Mixotoxodon larensis and Equus sp. are known from the area. All of these taxa are known from the Pleistocene and each one has a long chronological range limiting more precise age assessment of the fauna at this time. While exact locality information is not available, there are four specimens of Mammuthus known from the general vicinity of Estanzuela, making it the largest concentration of mammoth from a general area in Central America. Specimen: Right upper second molar, ME 121 (INGUAT PV-H36), housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Zacapa, (Fig. 1). ctor Enrique Pinto, EstanLocality: 10 Recado talud de casa He zuela, Departamento de Zacapa, Guatemala. The general location of Estanzuela is 14.9989 N, 89.7078 W, elevation 189 m. (Locality 5, Fig. 7). Description: The tooth, collected by Benjamin Cabrera and donated to the museum is complete and retains part of the attached
Fig. 1. Mammuthus columbi, right upper second molar. Estanzuela, Departamento de Zacapa. ME 121 (INGUAT PV-H-36) in A. lingual, B. occlusal view.
maxilla (Fig. 1). The occlusal surface includes all of the plates comprising the tooth and the entire tooth was erupted with slightly greater wear of the anterior part of the tooth. The total number of plates is 10 and the number of plates per 10 cm is 6. The length of tooth's occlusal surface is 203.7 mm and the width is 93.5 mm. Plate width varies between 8.5 and 11 mm and enamel thickness of the plates between 2.9 and 3.9 mm. Identification of the tooth as a second molar is based on the criteria outlined by Graham (1986) utilizing the number of plates, and the length and width of the tooth. Specimen: Tooth fragments with the occlusal surface, INGUAT PV-H-36 3, housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Departamento de Zacapa. No locality information, possibly from the vicinity of Estanzuela. (Locality 5, Fig. 7). Description: The fragments preserve parts of the occlusal surface of a mammoth tooth. The lack of associated locality data limits their usefulness. Based on their preservation, they most likely came from near Estanzuela, Departamento de Zacapa. Specimen: upper left fourth premolar or first molar (Fig. 2), MNHN Fs-001, identified as molar in the Museo National de Historia Natural. The specimen was collected by Barnum Brown in 1950 and donated to the museum. Locality: The tooth was collected in the vicinity of Estanzuela, Departamento de Zacapa. but specific locality information was not provided. The general location of Estanzuela is 14.9989 N; 89.7048 W, elevation 189 m. (Locality 5, Fig. 7). Description: The specimen is complete and has 10e11 plates, with 10 plates exposed and worn. The number of plates in
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Fig. 3. Mammuthus columbi, lower right molar, first or second. MNHNUSC uncatalogued, identified as molar B, Estanzuela, Departamento de Zacapa. in A. occlusal, B. lingual view.
town, no other data Departamento de Zacapa, approximate coordinates are 14.9897 N; 89.6975 W, elevation 189 m. (Locality 8, Fig. 2. Mammuthus columbi, upper left fourth premolar or first molar, MNHNUSC uncatalogued (molar A), Estanzuela, Departamento de Zacapa in A. labial, B. occlusal view.
10 cm ¼ 7. Length of the occlusal surface ¼ 158.5 mm; Maximum length of the tooth perpendicular to the plates is 188.5 mm; Width of the plates ranges between 7.7 and 12.55 mm. Thickness of the enamel of the plates 2.95 to 3 mm. Specimen: lower right molar, either a first or second. MNHN Fs002, identified as molar in the Museo National de Historia Natural (Fig. 3). The specimen was collected by Barnum Brown in 1950 and donated to the museum. Locality: The tooth was collected in the vicinity of Estanzuela, Departamento de Zacapa. but specific locality information was not provided. The general location of Estanzuela is 14.9989 N; 89.7048 W, elevation 189 m. (Locality 5, Fig. 7). Description: The tooth had not yet erupted and none of the plates show any wear. As the tooth has 12 plates, it is either a first or second molar, since there are more plates than are present in the premolars and not as many as the third molar (Haynes, 1991 Table A5A). The number of plates in 10 cm ¼ 7; Maximum length of the tooth perpendicular to the plates ¼ 125.7 mm; Length of the occlusal surface ¼ 146.6. Maximum width of the occlusal surface ¼ 52.5 mm; Width of the plates ranges between 7.99 and 12.41 mm. Thickness of the enamel of the plates 2.35 to 2.94 mm. Specimen: Lower first right molar. No catalog number, housed in the Colecciones Zoologicas de la Universidad del Valle de Guatemala, Guatemala City (Fig. 4). n, from the banks of Motagua River near the Locality: Teculuta
Fig. 4. Mammuthus columbi, lower first right molar. Uncataloged, Colecciones Zoologicas de la Universidad del Valle de Guatemala, Guatemala City; Teculutan Departamento de Zacapa in A. occlusal, B. labial view.
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Fig. 7). Description: The specimen is missing the anterior margin of the tooth. The tooth preserves 10 complete and half of an 11th plate, and based on the size of the missing portion probably is only missing two plates. The number of plates in 10 cm ¼ 6; Maximum length of tooth ¼ 187.5 mm; length of the occlusal surface is 180 mm; width of the plates 6.6e14.05 mm; width of the enamel of the plates ¼ 2.69e3.45 mm. Specimen: Lower left third molar, ME 110 (INGUAT PV-H-35), housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Departamento de Zacapa (Fig. 5). n, region de Sayaxche , DepartaLocality: Fondo de Río La Pasio n, Guatemala. The general location of the Sayaxche mento de El Pete region is 16.5314 N; 90.1892 W, elevation about 118 m. (Locality 7, Fig. 7). n by A Pleistocene fauna was collected from the Río La Pasio Barnum Brown and the specimens are housed at the American Museum of Natural History in New York. While the collection is n the collection actually represents referred to as from Río La Pasio numerous individual localities along the river and should not be treated as a single associated collection. Given the length of the Río n the Brown collections are best referred to as originating La Pasio from Santa Amelia, 16.2211 N, 90.0464 W; elevation 128 m. Except for the peccary, Dicotyles tajacu (Woodburne, 1969), the fauna has not been studied. Examination of these collections by Richard White on the behalf of the authors has indicated that no mammoth remains are present in the Brown collections. The Brown collec from tions are from farther upstream from the locality in Sayaxche which the specimen described here was recovered and indicates that there are numerous separate localities along the river containing Pleistocene fauna. The specimen described here was collected by Arturo Arbizú and donated to the Museum of Estanzuela on April 22, 1977 and there are no associated faunal remains with it in the collections. Description: The total number of plates in the tooth is 17 and the number of plates worn on the occlusal surface is 14 indicating the tooth was not fully erupted. The plate count in 10 cm is 7. The complete tooth has a total length of 340 mm. The length of occlusal surface is 186.4 mm and the width is 87.8 mm. The plate thickness
Fig. 5. Mammuthus columbi, lower left third molar. ME 110 (INGUAT PV-H-35). Río La n, Departamento de El Pete n in A. occlusal, B. labial view. Pasio
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ranges from 8.9 to 11 mm and enamel thickness is 2.7 mm. Specimen: Upper second molar, USAC 1,075, (Registro IDAEH 1.1.3.1) (Fig. 6) housed in the MUSHNAT-USAC. Locality: Chinautla, Departamento de Guatemala, old city of Chinautla. The general location of Chinautla, 14.0700 N, 90.4994 W, elevation 1227 m. The specimen was found in 1996, during a Irineo Montoya construction of a house, was recovered by Jose along with some parts of the mandible. A tooth of Equus sp. and a horn of cf. Euceratherium was found associated with the specimen vila, Cesar Augusto Nun ~ ez and and were collected by Lorena Da Sergio Ericastilla. (Locality 6, Fig. 7). Description: The occlusal surface extends the entire length of the tooth and is composed of nine plates. It is possible that the tooth had at least 10 plates because it is broken at the distal margin. Number of plates exposed on the occlusal surface ¼ 9, number of plates in 10 cm of the tooth ¼ 7, Maximum length of the tooth perpendicular to the plates is- 193 mm, length of the occlusal surface - 164 mm, maximum width of the occlusal surface is - 93.75 mm; width of plates ¼ 5.90e14.90 mm, width of enamel ¼ 2.15e3.90 mm. 3.2. Other Central American records of Mammuthus 3.2.1. Costa Rica n, Provincia de Puntarenas, Hacienda de El Silencio, Mata Limo Gutierrez, 1963; Espinoza, 1976; Alverado, 1986; Lucas et al., 1997 incomplete molar (right lower second or third) Lucas et al. (1997) reported the specimen is now missing (Locality 1, Fig. 7). 3.2.2. El Salvador n de Bank of Río Jerusalen, Hacienda San Lorenzo, Jurisdiccio Nueva Granada, Departmento de Usulutan e specimen in Museo Nacional of El Salvador. (Stirton and Gealey, 1949). (Locality 2,
Fig. 6. Mammuthus columbi, right lower first molar. MNHNUSC (USAC 1075) (Registro IDAEH 1.1.3.1). Chinautla, Departamento de Guatemala in A. occlusal and B. lingual view.
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Fig. 7. Distribution map of Mammuthus columbi in Central America. Costa Rica 1. Haciendo del Silencio, Provincia de Puntarenas; El Salvador 2, Río Jerusalen, Departamento de Usulutan, 3. Cerro Pacho, Departamento de Chalatenango, 4. Río Tomayate, Departamento de San Salvador; Guatemala 5. Estanzuela, Departamento de Zacapa, 6. Chinautla, n, Departamento de El Pete n, 8. Teculutan, Departamento de Zacapa; Honduras 9. Orillas del Humuya, Departamento de Comayagua; Departamento de Guatemala, 7. Río La Pasio Nicaragua 10. El Palmar, Departamento de Rivas, 11. Masachapa, Departamento de Managua, 12(?) Las Banderas, Departamento de Managua.
Fig. 7). n de Laguna, Seca, Jurisdiccio n de Foot of cerro Pacho, Canto n, Departamento de Chalatenango. A molar Nueva Concepcio (P268a) and a fragment of the tusk (P268.2) (Laurito and Aguilar, 2007). (Locality 3, Fig. 7). Along the margins of the Río Tomayate, 2.3 m above the upper level of the fossiliferous strata described by Cisneros (2005), Municipio de Apopa, 12 km north of the city of San Salvador. Fragment of a molar (2-ss-ap-30-893) (Laurito and Aguilar, 2007). (Locality 4, Fig. 7).
associated with the specimen and the report did not provide any specifics as to how the determination was made that they were mammoth and not one of the other proboscideans present in Central America we have identified this record in Fig. 7 with a question mark (?) based on our criterion of only utilizing records based on clearly diagnostic teeth. We do recognize that it may represent an additional record of Mammuthus in Central America and that there may be many additional unrecognized records represented by the post-cranial skeleton. 4. Discussion
3.2.3. Honduras Orillas del Humuya, on the Río Humuya, 3 km south of the bridge for the Carretera Nacional north of Comayagua, Departamento de Comayagua, 14.5 N, 87.6 W, 600 m, UF 17728 incomplete first lower molar. (Webb and Perrigo, 1984), (Locality 9, Fig. 7). 3.2.4. Nicaragua Río El Chorro, Departamento de Esteli. As Leidy (1886) did not provide details of the locality and assignment to this location follows Cisneros (2005) but it is not included in Fig. 7. El Palmar, 7 km south of Rivas, Departamento de Rivas, Museo Nacional de Nicaragua, 11.45 N, 85.9 W, elevation 71 m, partial skeleton including lower jaw (Lucas, 2004; Lucas et al., 2008). (Locality 10, Fig. 7). Masachapa, Departamento de Managua, Pacific Coast, Museo Nacional de Nicaragua, 11.8 N, 86.5 W, elevation 0 m, teeth. (Lucas, 2004; Lucas et al., 2008) (Locality 11, Fig. 7). Las Banderas, Departamento de Managua, 12.3 N, 86 W, elevation 87 m, (Lucas, 2004; Lucas et al., 2008). (Locality 12, Fig. 7). Mammoth bones were reported from Las Banderas, between Lake Nicaragua and Lake Managua, Departamento de Managua (Lucas, 2004; Lucas et al., 2008). Since no teeth were reported
Arroyo-Cabrales et al. (2003) in their review of records of mammoth in Mexico listed three records of mammoths in Chiapas, n in northern the state that borders the department of Pete Guatemala. There are no records for the Yucatan Peninsula (states of Tabasco, Campeche, Yucatan and Quintana Roo) and the current absence of mammoth in Belize may either reflect the absence of open savanna habitat in this area during the Pleistocene, or merely the lack of sufficient field work in this and adjacent regions in Mexico that border the Caribbean. In addition to the three records of mammoth in Chiapas there are two and six records in the northern adjacent states of Veracruz and Oaxaca respectively (Arroyo-Cabrales et al., 2003). While it might be expected that the population densities of mammoths would decrease southward (Fig. 7) reflecting more limited availability of suitable open country habitat and more closed forest, any patterns that can be observed based on the current state of knowledge more likely reflect that lack of extensive collecting in these regions rather then any indication of the relative abundance of mammoths in Central America. Hence, it is possible that mammoths may eventually be found in Panama as discussed below. Given the lack of any comprehensive summaries of the
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Pleistocene faunas for most of the countries in Central America detailed comparisons are not possible but the few examples are illustrative of the differences in relative abundance of mammoths compared to the other proboscideans in Central America. Laurito (1988) listed 15 records of gomphotheres in Costa Rica, compared to the single mammoth record. Cisneros (2005) in a comparison of 11 Central American faunas listed eight with gomphothere and two with mammoth. This pattern is the same in Guatemala, gomphotheres are known from at least 11 locations: Estanzuela, Chivacabe, San Rafelito, Mixco, Malacancito, Chinautla, Tívoli, Ciudad Real, El n compared to the four loRosario-Ipala, Jutiapa, and Río La Pasio cality records of mammoth, although both genera are found in Estanzuela and Chinautla (Arroyo-Cabrales et al., 2007). Based on our understanding of the differences in habitat preference of gomphotheres and mammoths the difference in their relative abundance in Central America is suggestive of a greater extent of forest compared to open country. Arroyo-Cabrales et al. (2003) noted that in Mexico much of the distribution of Mammuthus follows a pattern close to that of other Nearctic taxa. In Central America the pattern is less clear as there are few faunas that include Nearctic taxa along with mammoth. Most finds of mammoth consist of isolated specimens and are not part of a larger faunal assemblage. The few available records of faunal assemblages that include mammoth in Central America tend to include mostly representatives of taxa originating in South America and the only other taxa of North American origin are Cuvieronius and Equus. As a grazer Equus is often associated with Mammuthus in many North American faunas, but Cuvieronius is generally considered to be a forest browser and rarely found in the same fauna as Mammuthus (Lucas et al., 1999). Besides the two localities in Guatemala reported here, there is a Middle Pleistocene (Irvingtonian) fauna from Chalatenango, El Salvador with M. meridionalis and Cuvierionius sp. (Lucas and Alvarado, 2010). Given the paucity of data on the Pleistocene fauna of Central America any analysis of patterns of faunal associates with mammoth in Central America will be limited and for the moment can only be addressed in a general way. Among the taxa of South American origin associated with Mammuthus is Mixotoxodon. Based on the extreme hypsodonty of its dentition it might be inferred that Mixotoxodon like Mammuthus was an inhabitant of open savanna. The two taxa co-occur at Orillas del Humuya, Honduras as well as at Estanzuela, Guatemala and n, Guatemala, although the association is possibly at Río La Pasio less certain. Stable isotope studies of Mixotoxodon from Orillas del Humuya indicate that the dominant local habitat was C3 forests but that some C4 grasslands may have been present and at El Hatillo, Panama, which does not include mammoth in the fauna, it inhabited a canopied forest habitat (MacFadden, 2005). During the Last Glacial Maximum, it appears that the modern summer precipitation regime in the Yucatan had collapsed (Metcalfe et al., 2000) and a similar change in the climate regime appears to have occurred throughout Central America. Leyden (1984) and Leyden et al. (1993) documented that the Central American lowlands were cooler and more arid during Marine Isotope Stage 3 (~36e24 ka) and continued to cool with increasing aridity during Stage 2 (~24e13 ka), with the lowest lake levels at this time and the presence of savanna and juniper scrub habitat. Based on their work in Panama, Piperno and Jones (2003) indicated that during the late Pleistocene Central America was more arid and generally cooler then today and savanna grasslands expanded at the expense of tropical forested habitats, which became restricted to higher elevations. This may have provided the ideal situation for the southward expansion of mammoths into Central America based on our understanding of the distribution of mammoths in North America and their association with grassland environments.
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However, the distribution of open savanna in Central America may not have been continuous but rather consisted of isolated patches within a more forested environment, hence the presence of forest browsers such as Cuvieronius and the sloth, Eremotherium in some of the faunas that include Mammuthus. This may simply reflect the vegetational response to drier Pleistocene climates in the Pacific watershed lowlands with an annual precipitation of 2000 mm and less, and forests that were more prone to fragmentation (Piperno and Jones, 2003). Unfortunately, there are no radiocarbon dates for any of the Mammuthus from Central America which prevents placing them within a more precise chronological context. n and Río La Except for the records from Estanzuela, Teculuta n in Guatemala, which are located closer to the Caribbean side Pasio of the isthmus, the distribution of mammoths in Central America is primarily restricted to the Pacific side (Fig. 7) and may indicate where the major portions of savannah habitat in Central America existed during the Pleistocene. This distribution may reflect differences in seasonal rainfall on each side of the mountains extending through the center of the isthmus that separates the lowlands along the Atlantic and Pacific coasts, a pattern that exists today. Today Central America is embedded in an easterly trade wind flow for most of the year with dry months on the Pacific slopes coinciding with stronger low-level easterly flow, while ~ a and wetter months are associated with weaker winds (Pen Douglas, 2002). During the Pleistocene the long-term existence of similar patterns may have contributed to the spread of the more arid grassland habitat preferred by mammoths. Direct evidence for this is provided by the phytolith record from Monte Oscuro (9 20 000 N, 79 310 000 West), a crater lake located 10 m above sea level on the Pacific coastal plain of Panama. During the Late Pleistocene the lake bed was dry and savanna-like vegetation expanded at the expense of tropical deciduous forest reflecting a significant reduction in precipitation that would have resulted in the observed changes in vegetation (Piperno and Jones, 2003). Piperno and Jones (2003) noted that there was a persistence of forests during the late Pleistocene, but that the few known glacial-age paleovegetational sequences below 1000 m in tropical America have no modern analog. While most records of mammoth in Central America are at lower elevations well below 1000 m (Fig. 8) it should be noted that the presence of mammoth at Chinautla, Guatemala at 1232 m may indicate some flexibility in habitat use or simply the patchiness of the forest habitat that did exist. That these forests have no modern analog (Piperno and Jones, 2003) may also be a critical factor that requires further investigation. While Monte Oscuro is south of the current known distribution of mammoth in Central America it does indicate that potential
Fig. 8. Elevational distribution of Mammuthus in Central America by latitude.
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mammoth habitat extended farther south. Other pollen studies from lakes in Mexico and Guatemala show that vegetational shifts from forests (C3- dominated) to grasslands (C4-dominated) ecosystems in the lowland Neotropics are unlikely to occur without a significant reduction in precipitation (Huang et al., 2001). Given that mammoths in Central America were at the extreme southern margin of their distribution, their extirpation from the region was probably the result of increased rainfall, the resulting expansion of tropical rainforest into lower elevations, and the loss of suitable habitat. 5. Summary Currently there are 12 localities of the Columbian mammoth, Mammuthus columbi, documented from Central America, based on dentition, and one possible record based on the post-cranial skeleton. With the descriptions of the new specimens in this paper the number of records of mammoth from Guatemala is larger than any other Central American country. The distribution of mammoth in Central America is essentially limited to the Pacific side of the isthmus. We infer a southern expansion of suitable open grassland habitat for mammoth in the Pleistocene in association with stronger low-level easterly flow of wind creating more arid conditions on the Pacific side of the isthmus. Subsequent loss of those habitats may be related to elevational lowering of tropical rainforest as precipitation increased. Acknowledgements It is a pleasure to contribute a paper on mammoths to a volume dedicated to Larry Agenbroad, who was always ready to share his knowledge of them. He will be missed. We thank Richard White for checking the Barnum Brown collections from Guatemala in the American Museum of Natural History and providing information from their archives. Dr. Jack Shuster, curator at Universidad del Valle de Guatemala allowed us to study the specimen in his care. We thank the reviewers and editors for their insights and suggestions which have improved the quality of the manuscript. References Agenbroad, L.D., 2005. North American proboscideans: mammoths: the state of knowledge, 2003. Quat. Int. 126e128, 73e92. siles de Costa Rica. Rev. Geol. Alverado, G.E., 1986. Hallazgos de megamamíferos fo Ameríca Cent. 4, 1e46. Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Ferrusquía-Villafranca, I., 2010. A perspective on mammal biodiversity and zoogeography in the late pleistoxico. Quat. Int. 212, 187e197. cene of Me Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Guzman, A.F., 2003. The distribution of the genus Mammuthus in Mexico. Deinsea 9, 27e39. Arroyo-Cabrales, J., Polaco, O.J., Laurito, C., Johnson, E., Alberdi, M.T., Zamora, A.L.V., 2007. The proboscideans (mammalia) from mesoamerica). Quat. Int. 169e170, 17e23. Cisneros, J.C., 2005. New Pleistocene vertebrate fauna from El Salvador. Rev. Bras. Paleontol. 8, 239e255. Enk, J., Devault, A., Widga, C., Saunders, J., Szpak, P., Southon, J., Rouillard, J.-M., Shapiro, B., Golding, G.B., Zazula, G., Froese, D., Fisher, D.C., MacPhee, R.D.E., Poinar, H., 2016. Mammuthus population dynamics in late Pleistocene north America: divergence, phylogeography, and introgression. Front. Ecol. Evol. 4, 42. http://dx.doi.org/10.3389/fevo.2016.00042.
gicas en El Bosque. Instituto Geografica Espinoza, J.A., 1976. Evaluaciones arqueolo Nacional managua. Minist. Obras Publicas 1, 22e45. Falconer, H., 1857. On the species of mastodon and elephant occurring in the fossil state in great britain. Part I. Mastodon. Q. J. Geol. Soc. 13, 307e360. Graham, R.W., 1986. Descriptions of the dentitions and the stylohyoids of Mammuthus columbi from the colby site. In: Frison, G.C., Todd, L.C. (Eds.), The Colby Mammoth Site: Taphonomy and Archaeology of a Clovis kill Site in Northern Wyoming. University of New Mexico Press, Albuquerque, NM, pp. 171e190. Gutierrez, F., 1963. Hallazgo de restos de un mamut. Inst. Geogr. Nac. Costa Rica Inf. Semest. 3, 41e47. rrez-Bedolla, M., García-Zepeda, M.L., Lo pez-García, R., Arroyo-Cabrales, J., Gutie ndez-Herrera, E., Fuentes-Farías, A.L., 2016. Diet and Marín-Leyva, A.H., Mele habitat of Mammuthus columbi (Falconer, 1857) from two Late Pleistocene localities in central western Mexico. Quat. Int. 406 (Part B), 137e146. Haynes, G., 1991. Mammoths, Mastodonts, and Elephants: Biology, Behavior, and the Fossil Record. Cambridge University Press, Cambridge, UK, 413 pp. Huang, Y., Street-Perrott, F.A., Metcalfe, S.E., Brenner, M., Moreland, M., Freeman, K.H., 2001. Climate change as the dominant control on glacialeinterglacial variations in C3 and C4 plant abundance. Science 293, 1647e1651. siles de Costa Rica y su contexto en la Laurito, C.A., 1988. Los proboscídeos fo rica central. Vínculos 14, 29e58. Ame Laurito, C., Aguilar, D.H., 2007. El registro de Mammuthus (Proboscidea, Elerica central. Rev. Geol. Ame rica phantidae) en la República de El Salvador, Ame Cent. 34, 73e81. Leidy, J., 1886. Toxodon and other remains from Nicaragua. In: Proceedings of the Academy of Natural Sciences of Philadelphia (Unnumbered Volume), pp. 275e277. Leyden, B.W., 1984. Guatemalan forest synthesis after Pleistocene aridity. Proc. Natl. Acad. Sci. 81, 4856e4859. Leyden, B.W., Brenner, M., Hodell, D.A., Curtis, J.H., 1993. Late Pleistocene climate in the central American lowlands. Geophys. Monogr. 78, 165e178. Lister, A., Sher, A., 2015. Evolution and dispersal of mammoths across the Northern hemisphere. Science 350, 805e809. Lucas, S.G., 2004. New information on the Pleistocene mammals of Nicaragua. Curr. Res. Pleistocene 21, 97e98. Lucas, S.G., Alvarado, G.E., 2010. Fossil proboscidea from the Upper Cenozoic of Central America: taxonomy, evolutionary and paleobiogeographic significance. rica Cent. 42, 9e42. Rev. Geol. Ame Lucas, S.G., Alvarado, G.E., Vega, E., 1997. The Pleistocene mammals of Costa Rica. J. Vertebrate Paleontol. 17, 413e427. Lucas, S.G., Garcia, R., Espinoza, E., Alvardo, G.E., de Mendoza, L.H., Vega, E., 2008. The fossil mammals of Nicaragua. N. M. Mus. Nat. Hist. Sci. Bull. 44, 417e429. Lucas, S.G., Morgan, G.S., Estep, J.W., Mack, G.H., Hawley, J.W., 1999. Co- occurrence of the proboscideans Cuvieronius, stegomastodon, and Mammuthus in the lower Pleistocene of southern New Mexico. J. Vertebrate Paleontol. 19, 595e597. MacFadden, B.J., 2005. Diet and habitat of toxodont megaherbivores (mammalia, Notoungulata) from the late quaternary of South and Central America. Quat. Res. 64, 113e124. Mead, J.I., Baez, A., Swift, S.L., Jon Lohse, J., Paiz, L., 2012. Late Pleistocene mammals , huehuetenango, Guatemala. Rev. Mex. Ciencias Geol. 29, from chivacabe 319e329. Metcalfe, S.E., O'Hara, S.L., Caballero, M., Davies, S.J., 2000. Records of late pleistocene-holocene climate change in Mexico e a review. Quat. Sci. Rev. 19, 699e721. ~ a, M., Douglas, M.W., 2002. Characteristics of wet and dry spells over the Pacific Pen side of Central America during the rainy season. Mon. Weather Rev. 130, 3054e3073. rez-Crespo, V.A., Arroyo-Cabrales, J., Benammi, M., Johnson, E., Polaco, O.J., SanPe tos-Moreno, A., Morales-Puente, P., Cienfuegos-Alvarado, E., 2012. Geographic variation of diet and habitat of the Mexican populations of Columbian Mammoth (Mammuthus columbi). Quat. Int. 276e277, 8e16. Piperno, D.R., Jones, J.G., 2003. Paleoecological and archaeological implications of a Late Pleistocene/Early Holocene record of vegetation and climate from the Pacific coastal plain of panama. Quat. Res. 59, 79e87. Stirton, R.A., Gealey, W.K., 1949. Reconnaissance geology and vertebrate paleontology of El Salvador, Central America. Geol. Soc. Am. Bull. 60, 1731e1754. Webb, S.D., Perrigo, S.C., 1984. Late cenozoic vertebrates from Honduras and el alvador. J. Vertebrate Paleontol. 4, 237e254. Woodburne, M.O., 1969. A late Pleistocene occurrence of the collared peccary, Dicotyles tajacu, in Guatemala. J. Mammal. 50, 121e125.
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Quaternary International journal homepage: www.elsevier.com/locate/quaint
Mammoths in Central America: New records from Guatemala vila A b, ** H. Gregory McDonald a, *, Sylvia Lorena Da a
Bureau of Land Management, Utah State Office, 400 West 200 South, Salt Lake City, UT, 84101, USA n de Fo siles, Museo Historia Natural, Universidad de San Carlos de Guatemala, Calle Mariscal Cruz 1-56, Zona 10, Ciudad de Guatemala, Coleccio Guatemala b
a r t i c l e i n f o
a b s t r a c t
Article history: Received 7 December 2015 Received in revised form 8 December 2016 Accepted 18 December 2016 Available online 13 July 2017
The remains of the Columbian Mammoth, Mammuthus columbi, are reported from four localities in Guatemala based on seven dental records. These localities are widely separated and include one spec n, Sayaxche , Departamento de Pete n in the northern part of the country, one imen from Río La Pasio specimen from Chinautla, Departamento de Guatemala in the southwest and four specimens from the n, Departamento de Zacapa in the vicinity of Estanzuela, Departamento de Zacapa and one from Teculuta southeast. Based on the known records the distribution of Mammuthus in Central America tends to be concentrated on the Pacific side of the isthmus and suggests the former presence of a savanna corridor that formed during an interval of decreased precipitation. © 2016 Published by Elsevier Ltd.
Keywords: Mammuthus columbi Mammoth Central America Guatemala Pleistocene
r e s u m e n Los primeros vestigios de Mamuts, Mammuthus columbi, se han reportado de cuatro localidades en n, Sayaxche Departamento de Guatemala, muy distantes entre ellas: uno procedente de Río La Pasio n, en la parte septentrional del país, otro de Chinautla, Departamento de Guatemala en el suroeste, y Pete n, Departamento de Zacapa, en el sureste. Estos nuevos registros cuatro de Estanzuela, y uno de Teculuta brindan un número total de siete basados en las evidencias dentales conocidas de Mammuthus de n de Mamuts, en America Central, los regristros Guatemala. Basados en el conocimiento de la distribucio se centran en la vertiente del pacifico del Itsmo, sugieriendo la presencia de un corridor seco de sabana. © 2016 Published by Elsevier Ltd.
1. Introduction While mammoths are widely distributed in North America from Canada through the United States (Agenbroad, 2005) and into Mexico (Arroyo-Cabrales et al., 2003) their record in Central America is minimal with only a few scattered records in five of the seven countries. In Mexico mammoths have been studied in more rez-Crespo et al., 2012; detail (Arroyo Cabrales et al., 2010; Pe rrez-Bedolla et al., 2016) but much less is known about Gutie them in Central America. The presence of Mammuthus in Central America was first reported by Leidy (1886) as part of a small Pleistocene fauna from
* Corresponding author. ** Corresponding author. E-mail addresses: [email protected] (H.G. McDonald), [email protected] vila A). (S.L. Da http://dx.doi.org/10.1016/j.quaint.2016.12.018 1040-6182/© 2016 Published by Elsevier Ltd.
Nicaragua and the genus has subsequently been found in Guatemala, El Salvador, Honduras, and Costa Rica (Laurito and Aguilar, 2007; Lucas and Alvarado, 2010). The recovery of additional mammoth remains in Guatemala provides a stronger geographic link between Mexico and these more southern records and improves our understanding of the southernmost portion of the genus' distribution. As there are other Pleistocene proboscideans, Cuvieronius and Mammut, present in Central America (Arroyo-Cabrales et al., 2007; Lucas and Alvarado, 2010) and Cuvieronius is known from Guatemala (Mead et al., 2012), we have only considered those records of mammoth based on dentition. We recognize that other records of mammoth may exist represented by only the post-cranial skeleton, such as the specimen from Las Banderas, Nicaragua, reported by Lucas (2004). We do not consider these here given the lack of detailed anatomical studies of differences in the post-cranial skeleton of mammoths and the other genera of proboscideans present in Central America and the uncertainty related to taxonomic identifications based on post-cranial
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specimens. The taxonomic assignment to species of specimens of Mammuthus has varied in the past and often reflected the changing opinions on the valid species of North American Mammuthus based on both morphological or molecular criteria (Lister and Sher, 2015; Enk et al., 2016). Based strictly on tooth morphology, records of Mammuthus from Central America can be referred to M. columbi, although some earlier records were referred to the species M. imperator (Gutierrez, 1963) and Lucas and Alvarado (2010) recognize a record of M. meridionalis from El Salvador. Based on the limited available stratigraphic and chronologic information available for most specimens it seems most likely that most of the Mammuthus records in Central America can be considered to be late Pleistocene in age. The new specimens from Guatemala have the meristic and morphological features currently used to diagnose Mammuthus columbi and are assigned to this species. 2. Abbreviations INGUAT e Instituto Guatemalteco de Turismo ME e Museo de Estanzuela, Estanzuela MUSHNAT e Usac e Museo de Historia Natural de la Universidad de San Carlos de Guatemala, Guatemala City MNHN e Museo Nacional de Historia Natural (Estatal). Guatemala, City While some workers use a system of identifying mammoth dentition as M/m 1e6 for the teeth to distinquish them based on the sequence of eruption, we have identified the teeth according to their inferred homology as a premolar or molar. All measurements are in millimeters. 3. Description of material Order Proboscidea Illiger, 1811 Family Elephantidae Gray, 1821 Mammuthus Brookes, 1828 Mammuthus columbi (Falconer), 1857
3.1. Guatemala The area around Estanzuela is perhaps the richest in Pleistocene vertebrates for Guatemala in terms of the number of specimens of Pleistocene fauna known from a restricted location within the country. However, despite the large number of specimens the species richness is low and limited to megafauna. Along with the mammoth, Cuvieronius hyodon, Eremotherium laurillardi, Glyptotherium cf. G. floridanus, Neochoerus sp., Mixotoxodon larensis and Equus sp. are known from the area. All of these taxa are known from the Pleistocene and each one has a long chronological range limiting more precise age assessment of the fauna at this time. While exact locality information is not available, there are four specimens of Mammuthus known from the general vicinity of Estanzuela, making it the largest concentration of mammoth from a general area in Central America. Specimen: Right upper second molar, ME 121 (INGUAT PV-H36), housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Zacapa, (Fig. 1). ctor Enrique Pinto, EstanLocality: 10 Recado talud de casa He zuela, Departamento de Zacapa, Guatemala. The general location of Estanzuela is 14.9989 N, 89.7078 W, elevation 189 m. (Locality 5, Fig. 7). Description: The tooth, collected by Benjamin Cabrera and donated to the museum is complete and retains part of the attached
Fig. 1. Mammuthus columbi, right upper second molar. Estanzuela, Departamento de Zacapa. ME 121 (INGUAT PV-H-36) in A. lingual, B. occlusal view.
maxilla (Fig. 1). The occlusal surface includes all of the plates comprising the tooth and the entire tooth was erupted with slightly greater wear of the anterior part of the tooth. The total number of plates is 10 and the number of plates per 10 cm is 6. The length of tooth's occlusal surface is 203.7 mm and the width is 93.5 mm. Plate width varies between 8.5 and 11 mm and enamel thickness of the plates between 2.9 and 3.9 mm. Identification of the tooth as a second molar is based on the criteria outlined by Graham (1986) utilizing the number of plates, and the length and width of the tooth. Specimen: Tooth fragments with the occlusal surface, INGUAT PV-H-36 3, housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Departamento de Zacapa. No locality information, possibly from the vicinity of Estanzuela. (Locality 5, Fig. 7). Description: The fragments preserve parts of the occlusal surface of a mammoth tooth. The lack of associated locality data limits their usefulness. Based on their preservation, they most likely came from near Estanzuela, Departamento de Zacapa. Specimen: upper left fourth premolar or first molar (Fig. 2), MNHN Fs-001, identified as molar in the Museo National de Historia Natural. The specimen was collected by Barnum Brown in 1950 and donated to the museum. Locality: The tooth was collected in the vicinity of Estanzuela, Departamento de Zacapa. but specific locality information was not provided. The general location of Estanzuela is 14.9989 N; 89.7048 W, elevation 189 m. (Locality 5, Fig. 7). Description: The specimen is complete and has 10e11 plates, with 10 plates exposed and worn. The number of plates in
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Fig. 3. Mammuthus columbi, lower right molar, first or second. MNHNUSC uncatalogued, identified as molar B, Estanzuela, Departamento de Zacapa. in A. occlusal, B. lingual view.
town, no other data Departamento de Zacapa, approximate coordinates are 14.9897 N; 89.6975 W, elevation 189 m. (Locality 8, Fig. 2. Mammuthus columbi, upper left fourth premolar or first molar, MNHNUSC uncatalogued (molar A), Estanzuela, Departamento de Zacapa in A. labial, B. occlusal view.
10 cm ¼ 7. Length of the occlusal surface ¼ 158.5 mm; Maximum length of the tooth perpendicular to the plates is 188.5 mm; Width of the plates ranges between 7.7 and 12.55 mm. Thickness of the enamel of the plates 2.95 to 3 mm. Specimen: lower right molar, either a first or second. MNHN Fs002, identified as molar in the Museo National de Historia Natural (Fig. 3). The specimen was collected by Barnum Brown in 1950 and donated to the museum. Locality: The tooth was collected in the vicinity of Estanzuela, Departamento de Zacapa. but specific locality information was not provided. The general location of Estanzuela is 14.9989 N; 89.7048 W, elevation 189 m. (Locality 5, Fig. 7). Description: The tooth had not yet erupted and none of the plates show any wear. As the tooth has 12 plates, it is either a first or second molar, since there are more plates than are present in the premolars and not as many as the third molar (Haynes, 1991 Table A5A). The number of plates in 10 cm ¼ 7; Maximum length of the tooth perpendicular to the plates ¼ 125.7 mm; Length of the occlusal surface ¼ 146.6. Maximum width of the occlusal surface ¼ 52.5 mm; Width of the plates ranges between 7.99 and 12.41 mm. Thickness of the enamel of the plates 2.35 to 2.94 mm. Specimen: Lower first right molar. No catalog number, housed in the Colecciones Zoologicas de la Universidad del Valle de Guatemala, Guatemala City (Fig. 4). n, from the banks of Motagua River near the Locality: Teculuta
Fig. 4. Mammuthus columbi, lower first right molar. Uncataloged, Colecciones Zoologicas de la Universidad del Valle de Guatemala, Guatemala City; Teculutan Departamento de Zacapa in A. occlusal, B. labial view.
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Fig. 7). Description: The specimen is missing the anterior margin of the tooth. The tooth preserves 10 complete and half of an 11th plate, and based on the size of the missing portion probably is only missing two plates. The number of plates in 10 cm ¼ 6; Maximum length of tooth ¼ 187.5 mm; length of the occlusal surface is 180 mm; width of the plates 6.6e14.05 mm; width of the enamel of the plates ¼ 2.69e3.45 mm. Specimen: Lower left third molar, ME 110 (INGUAT PV-H-35), housed in the Museo de Paleontología y Arqueología “Ingeniero Roberto Woolfolk Saravia”, Estanzuela, Departamento de Zacapa (Fig. 5). n, region de Sayaxche , DepartaLocality: Fondo de Río La Pasio n, Guatemala. The general location of the Sayaxche mento de El Pete region is 16.5314 N; 90.1892 W, elevation about 118 m. (Locality 7, Fig. 7). n by A Pleistocene fauna was collected from the Río La Pasio Barnum Brown and the specimens are housed at the American Museum of Natural History in New York. While the collection is n the collection actually represents referred to as from Río La Pasio numerous individual localities along the river and should not be treated as a single associated collection. Given the length of the Río n the Brown collections are best referred to as originating La Pasio from Santa Amelia, 16.2211 N, 90.0464 W; elevation 128 m. Except for the peccary, Dicotyles tajacu (Woodburne, 1969), the fauna has not been studied. Examination of these collections by Richard White on the behalf of the authors has indicated that no mammoth remains are present in the Brown collections. The Brown collec from tions are from farther upstream from the locality in Sayaxche which the specimen described here was recovered and indicates that there are numerous separate localities along the river containing Pleistocene fauna. The specimen described here was collected by Arturo Arbizú and donated to the Museum of Estanzuela on April 22, 1977 and there are no associated faunal remains with it in the collections. Description: The total number of plates in the tooth is 17 and the number of plates worn on the occlusal surface is 14 indicating the tooth was not fully erupted. The plate count in 10 cm is 7. The complete tooth has a total length of 340 mm. The length of occlusal surface is 186.4 mm and the width is 87.8 mm. The plate thickness
Fig. 5. Mammuthus columbi, lower left third molar. ME 110 (INGUAT PV-H-35). Río La n, Departamento de El Pete n in A. occlusal, B. labial view. Pasio
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ranges from 8.9 to 11 mm and enamel thickness is 2.7 mm. Specimen: Upper second molar, USAC 1,075, (Registro IDAEH 1.1.3.1) (Fig. 6) housed in the MUSHNAT-USAC. Locality: Chinautla, Departamento de Guatemala, old city of Chinautla. The general location of Chinautla, 14.0700 N, 90.4994 W, elevation 1227 m. The specimen was found in 1996, during a Irineo Montoya construction of a house, was recovered by Jose along with some parts of the mandible. A tooth of Equus sp. and a horn of cf. Euceratherium was found associated with the specimen vila, Cesar Augusto Nun ~ ez and and were collected by Lorena Da Sergio Ericastilla. (Locality 6, Fig. 7). Description: The occlusal surface extends the entire length of the tooth and is composed of nine plates. It is possible that the tooth had at least 10 plates because it is broken at the distal margin. Number of plates exposed on the occlusal surface ¼ 9, number of plates in 10 cm of the tooth ¼ 7, Maximum length of the tooth perpendicular to the plates is- 193 mm, length of the occlusal surface - 164 mm, maximum width of the occlusal surface is - 93.75 mm; width of plates ¼ 5.90e14.90 mm, width of enamel ¼ 2.15e3.90 mm. 3.2. Other Central American records of Mammuthus 3.2.1. Costa Rica n, Provincia de Puntarenas, Hacienda de El Silencio, Mata Limo Gutierrez, 1963; Espinoza, 1976; Alverado, 1986; Lucas et al., 1997 incomplete molar (right lower second or third) Lucas et al. (1997) reported the specimen is now missing (Locality 1, Fig. 7). 3.2.2. El Salvador n de Bank of Río Jerusalen, Hacienda San Lorenzo, Jurisdiccio Nueva Granada, Departmento de Usulutan e specimen in Museo Nacional of El Salvador. (Stirton and Gealey, 1949). (Locality 2,
Fig. 6. Mammuthus columbi, right lower first molar. MNHNUSC (USAC 1075) (Registro IDAEH 1.1.3.1). Chinautla, Departamento de Guatemala in A. occlusal and B. lingual view.
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Fig. 7. Distribution map of Mammuthus columbi in Central America. Costa Rica 1. Haciendo del Silencio, Provincia de Puntarenas; El Salvador 2, Río Jerusalen, Departamento de Usulutan, 3. Cerro Pacho, Departamento de Chalatenango, 4. Río Tomayate, Departamento de San Salvador; Guatemala 5. Estanzuela, Departamento de Zacapa, 6. Chinautla, n, Departamento de El Pete n, 8. Teculutan, Departamento de Zacapa; Honduras 9. Orillas del Humuya, Departamento de Comayagua; Departamento de Guatemala, 7. Río La Pasio Nicaragua 10. El Palmar, Departamento de Rivas, 11. Masachapa, Departamento de Managua, 12(?) Las Banderas, Departamento de Managua.
Fig. 7). n de Laguna, Seca, Jurisdiccio n de Foot of cerro Pacho, Canto n, Departamento de Chalatenango. A molar Nueva Concepcio (P268a) and a fragment of the tusk (P268.2) (Laurito and Aguilar, 2007). (Locality 3, Fig. 7). Along the margins of the Río Tomayate, 2.3 m above the upper level of the fossiliferous strata described by Cisneros (2005), Municipio de Apopa, 12 km north of the city of San Salvador. Fragment of a molar (2-ss-ap-30-893) (Laurito and Aguilar, 2007). (Locality 4, Fig. 7).
associated with the specimen and the report did not provide any specifics as to how the determination was made that they were mammoth and not one of the other proboscideans present in Central America we have identified this record in Fig. 7 with a question mark (?) based on our criterion of only utilizing records based on clearly diagnostic teeth. We do recognize that it may represent an additional record of Mammuthus in Central America and that there may be many additional unrecognized records represented by the post-cranial skeleton. 4. Discussion
3.2.3. Honduras Orillas del Humuya, on the Río Humuya, 3 km south of the bridge for the Carretera Nacional north of Comayagua, Departamento de Comayagua, 14.5 N, 87.6 W, 600 m, UF 17728 incomplete first lower molar. (Webb and Perrigo, 1984), (Locality 9, Fig. 7). 3.2.4. Nicaragua Río El Chorro, Departamento de Esteli. As Leidy (1886) did not provide details of the locality and assignment to this location follows Cisneros (2005) but it is not included in Fig. 7. El Palmar, 7 km south of Rivas, Departamento de Rivas, Museo Nacional de Nicaragua, 11.45 N, 85.9 W, elevation 71 m, partial skeleton including lower jaw (Lucas, 2004; Lucas et al., 2008). (Locality 10, Fig. 7). Masachapa, Departamento de Managua, Pacific Coast, Museo Nacional de Nicaragua, 11.8 N, 86.5 W, elevation 0 m, teeth. (Lucas, 2004; Lucas et al., 2008) (Locality 11, Fig. 7). Las Banderas, Departamento de Managua, 12.3 N, 86 W, elevation 87 m, (Lucas, 2004; Lucas et al., 2008). (Locality 12, Fig. 7). Mammoth bones were reported from Las Banderas, between Lake Nicaragua and Lake Managua, Departamento de Managua (Lucas, 2004; Lucas et al., 2008). Since no teeth were reported
Arroyo-Cabrales et al. (2003) in their review of records of mammoth in Mexico listed three records of mammoths in Chiapas, n in northern the state that borders the department of Pete Guatemala. There are no records for the Yucatan Peninsula (states of Tabasco, Campeche, Yucatan and Quintana Roo) and the current absence of mammoth in Belize may either reflect the absence of open savanna habitat in this area during the Pleistocene, or merely the lack of sufficient field work in this and adjacent regions in Mexico that border the Caribbean. In addition to the three records of mammoth in Chiapas there are two and six records in the northern adjacent states of Veracruz and Oaxaca respectively (Arroyo-Cabrales et al., 2003). While it might be expected that the population densities of mammoths would decrease southward (Fig. 7) reflecting more limited availability of suitable open country habitat and more closed forest, any patterns that can be observed based on the current state of knowledge more likely reflect that lack of extensive collecting in these regions rather then any indication of the relative abundance of mammoths in Central America. Hence, it is possible that mammoths may eventually be found in Panama as discussed below. Given the lack of any comprehensive summaries of the
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Pleistocene faunas for most of the countries in Central America detailed comparisons are not possible but the few examples are illustrative of the differences in relative abundance of mammoths compared to the other proboscideans in Central America. Laurito (1988) listed 15 records of gomphotheres in Costa Rica, compared to the single mammoth record. Cisneros (2005) in a comparison of 11 Central American faunas listed eight with gomphothere and two with mammoth. This pattern is the same in Guatemala, gomphotheres are known from at least 11 locations: Estanzuela, Chivacabe, San Rafelito, Mixco, Malacancito, Chinautla, Tívoli, Ciudad Real, El n compared to the four loRosario-Ipala, Jutiapa, and Río La Pasio cality records of mammoth, although both genera are found in Estanzuela and Chinautla (Arroyo-Cabrales et al., 2007). Based on our understanding of the differences in habitat preference of gomphotheres and mammoths the difference in their relative abundance in Central America is suggestive of a greater extent of forest compared to open country. Arroyo-Cabrales et al. (2003) noted that in Mexico much of the distribution of Mammuthus follows a pattern close to that of other Nearctic taxa. In Central America the pattern is less clear as there are few faunas that include Nearctic taxa along with mammoth. Most finds of mammoth consist of isolated specimens and are not part of a larger faunal assemblage. The few available records of faunal assemblages that include mammoth in Central America tend to include mostly representatives of taxa originating in South America and the only other taxa of North American origin are Cuvieronius and Equus. As a grazer Equus is often associated with Mammuthus in many North American faunas, but Cuvieronius is generally considered to be a forest browser and rarely found in the same fauna as Mammuthus (Lucas et al., 1999). Besides the two localities in Guatemala reported here, there is a Middle Pleistocene (Irvingtonian) fauna from Chalatenango, El Salvador with M. meridionalis and Cuvierionius sp. (Lucas and Alvarado, 2010). Given the paucity of data on the Pleistocene fauna of Central America any analysis of patterns of faunal associates with mammoth in Central America will be limited and for the moment can only be addressed in a general way. Among the taxa of South American origin associated with Mammuthus is Mixotoxodon. Based on the extreme hypsodonty of its dentition it might be inferred that Mixotoxodon like Mammuthus was an inhabitant of open savanna. The two taxa co-occur at Orillas del Humuya, Honduras as well as at Estanzuela, Guatemala and n, Guatemala, although the association is possibly at Río La Pasio less certain. Stable isotope studies of Mixotoxodon from Orillas del Humuya indicate that the dominant local habitat was C3 forests but that some C4 grasslands may have been present and at El Hatillo, Panama, which does not include mammoth in the fauna, it inhabited a canopied forest habitat (MacFadden, 2005). During the Last Glacial Maximum, it appears that the modern summer precipitation regime in the Yucatan had collapsed (Metcalfe et al., 2000) and a similar change in the climate regime appears to have occurred throughout Central America. Leyden (1984) and Leyden et al. (1993) documented that the Central American lowlands were cooler and more arid during Marine Isotope Stage 3 (~36e24 ka) and continued to cool with increasing aridity during Stage 2 (~24e13 ka), with the lowest lake levels at this time and the presence of savanna and juniper scrub habitat. Based on their work in Panama, Piperno and Jones (2003) indicated that during the late Pleistocene Central America was more arid and generally cooler then today and savanna grasslands expanded at the expense of tropical forested habitats, which became restricted to higher elevations. This may have provided the ideal situation for the southward expansion of mammoths into Central America based on our understanding of the distribution of mammoths in North America and their association with grassland environments.
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However, the distribution of open savanna in Central America may not have been continuous but rather consisted of isolated patches within a more forested environment, hence the presence of forest browsers such as Cuvieronius and the sloth, Eremotherium in some of the faunas that include Mammuthus. This may simply reflect the vegetational response to drier Pleistocene climates in the Pacific watershed lowlands with an annual precipitation of 2000 mm and less, and forests that were more prone to fragmentation (Piperno and Jones, 2003). Unfortunately, there are no radiocarbon dates for any of the Mammuthus from Central America which prevents placing them within a more precise chronological context. n and Río La Except for the records from Estanzuela, Teculuta n in Guatemala, which are located closer to the Caribbean side Pasio of the isthmus, the distribution of mammoths in Central America is primarily restricted to the Pacific side (Fig. 7) and may indicate where the major portions of savannah habitat in Central America existed during the Pleistocene. This distribution may reflect differences in seasonal rainfall on each side of the mountains extending through the center of the isthmus that separates the lowlands along the Atlantic and Pacific coasts, a pattern that exists today. Today Central America is embedded in an easterly trade wind flow for most of the year with dry months on the Pacific slopes coinciding with stronger low-level easterly flow, while ~ a and wetter months are associated with weaker winds (Pen Douglas, 2002). During the Pleistocene the long-term existence of similar patterns may have contributed to the spread of the more arid grassland habitat preferred by mammoths. Direct evidence for this is provided by the phytolith record from Monte Oscuro (9 20 000 N, 79 310 000 West), a crater lake located 10 m above sea level on the Pacific coastal plain of Panama. During the Late Pleistocene the lake bed was dry and savanna-like vegetation expanded at the expense of tropical deciduous forest reflecting a significant reduction in precipitation that would have resulted in the observed changes in vegetation (Piperno and Jones, 2003). Piperno and Jones (2003) noted that there was a persistence of forests during the late Pleistocene, but that the few known glacial-age paleovegetational sequences below 1000 m in tropical America have no modern analog. While most records of mammoth in Central America are at lower elevations well below 1000 m (Fig. 8) it should be noted that the presence of mammoth at Chinautla, Guatemala at 1232 m may indicate some flexibility in habitat use or simply the patchiness of the forest habitat that did exist. That these forests have no modern analog (Piperno and Jones, 2003) may also be a critical factor that requires further investigation. While Monte Oscuro is south of the current known distribution of mammoth in Central America it does indicate that potential
Fig. 8. Elevational distribution of Mammuthus in Central America by latitude.
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mammoth habitat extended farther south. Other pollen studies from lakes in Mexico and Guatemala show that vegetational shifts from forests (C3- dominated) to grasslands (C4-dominated) ecosystems in the lowland Neotropics are unlikely to occur without a significant reduction in precipitation (Huang et al., 2001). Given that mammoths in Central America were at the extreme southern margin of their distribution, their extirpation from the region was probably the result of increased rainfall, the resulting expansion of tropical rainforest into lower elevations, and the loss of suitable habitat. 5. Summary Currently there are 12 localities of the Columbian mammoth, Mammuthus columbi, documented from Central America, based on dentition, and one possible record based on the post-cranial skeleton. With the descriptions of the new specimens in this paper the number of records of mammoth from Guatemala is larger than any other Central American country. The distribution of mammoth in Central America is essentially limited to the Pacific side of the isthmus. We infer a southern expansion of suitable open grassland habitat for mammoth in the Pleistocene in association with stronger low-level easterly flow of wind creating more arid conditions on the Pacific side of the isthmus. Subsequent loss of those habitats may be related to elevational lowering of tropical rainforest as precipitation increased. Acknowledgements It is a pleasure to contribute a paper on mammoths to a volume dedicated to Larry Agenbroad, who was always ready to share his knowledge of them. He will be missed. We thank Richard White for checking the Barnum Brown collections from Guatemala in the American Museum of Natural History and providing information from their archives. Dr. Jack Shuster, curator at Universidad del Valle de Guatemala allowed us to study the specimen in his care. We thank the reviewers and editors for their insights and suggestions which have improved the quality of the manuscript. References Agenbroad, L.D., 2005. North American proboscideans: mammoths: the state of knowledge, 2003. Quat. Int. 126e128, 73e92. siles de Costa Rica. Rev. Geol. Alverado, G.E., 1986. Hallazgos de megamamíferos fo Ameríca Cent. 4, 1e46. Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Ferrusquía-Villafranca, I., 2010. A perspective on mammal biodiversity and zoogeography in the late pleistoxico. Quat. Int. 212, 187e197. cene of Me Arroyo-Cabrales, J., Polaco, O.J., Johnson, E., Guzman, A.F., 2003. The distribution of the genus Mammuthus in Mexico. Deinsea 9, 27e39. Arroyo-Cabrales, J., Polaco, O.J., Laurito, C., Johnson, E., Alberdi, M.T., Zamora, A.L.V., 2007. The proboscideans (mammalia) from mesoamerica). Quat. Int. 169e170, 17e23. Cisneros, J.C., 2005. New Pleistocene vertebrate fauna from El Salvador. Rev. Bras. Paleontol. 8, 239e255. Enk, J., Devault, A., Widga, C., Saunders, J., Szpak, P., Southon, J., Rouillard, J.-M., Shapiro, B., Golding, G.B., Zazula, G., Froese, D., Fisher, D.C., MacPhee, R.D.E., Poinar, H., 2016. Mammuthus population dynamics in late Pleistocene north America: divergence, phylogeography, and introgression. Front. Ecol. Evol. 4, 42. http://dx.doi.org/10.3389/fevo.2016.00042.
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