Lucas et al., eds., 2008, Neogene Mammals. New Mexico Museum of Natural History and Science Bulletin 44.

417
THE FOSSIL MAMMALS OF NICARAGUA

SPENCER G. LUCAS1, RAMIRO GARCIA2, EDGAR ESPINOZA2, GUILLERMO E. ALVARADO3,

LUIS HURTADO DE MENDOZA3 AND EDUARDO VEGA3
1
New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., Albuquerque, New Mexico 8714;
2
Museo Nacional de Nicaragua, Managua, Nicaragua;
3
Instituto Costarricense de Electricidada (ICE), Apdo. 10032-1000, San José, Costa Rica

Abstract—Fossil mammals of Pliocene and Pleistocene age are known from ten documented localities in Nicara-
gua. The only Pliocene mammal is a mysticete cetacean from marine strata of the El Salto Formation at the Mine
K-11 locality near San Rafael del Sur. Other Nicaraguan fossil mammal localities are of Pleistocene age (all are
probably late Pleistocene) from fluvial, lacustrine and volcaniclastic sediments and from ash-flow tuff deposits,
primarily of the Nicaraguan depression and the interior highlands. The complete Pleistocene mammal faunal list
from Nicaragua is: the ground sloth Eremotherium laurillardi, a procyonid carnivore, the capybara Neochoerus
aesopi (Leidy), the toxodont Mixotoxodon larensis, the gomphothere Cuvieronius hyodon, the elephantid
Mammuthus columbi, the horse Equus cf. E. occidentalis and the bison Bison sp. Reports of the deer Odocoileus
from Nicaragua are plausible but remain undocumented. Previous reports of Stegomastodon and Gomphotherium
are implausible. The association of Late Pleistocene (older than ~ 30,000 yBP) fossil mammals with supposed
human artifacts at the El Bosque locality is not demonstrated. Like other Central American Pleistocene mammal
records, the Pleistocene mammals of Nicaragua are almost all large edentates and ungulates and are a nearly equal
mixture of South American (capybara, sloth, toxodont) and North American (horse, bison, proboscidean) immi-
grants. The relative rarity of the common Central American gomphothere Cuvieronius and relative abundance of
Mammuthus may suggest a greater prevalence of savanna in western Nicaragua during the late Pleistocene than in
other Central American countries.

INTRODUCTION
Although the first published record of fossil mammals from Cen-
tral America was of fossils from Nicaragua (Leidy, 1886), much less is
known of Nicaraguan fossil mammals than from any other Central Ameri-
can country (Lucas and Alvarado, 1994; Lucas et al., 1997, 2007). This
largely reflects a lack of exploration and study. Indeed, after Leidy (1886),
only Espinoza (1976) has published detailed information on Nicaraguan
fossil mammals.
Here, we report the results of our efforts to study all of the fossil
mammals known from Nicaragua. This study was based on an examina-
tion of all known fossil mammal localities in the country in 1999 and
2001, which are documented here (Fig. 1). We also located and evaluated
specimens of fossil mammals from Nicaragua in museum collections.
These are in the Academy of Natural Sciences in Philadelphia (ANSP),
the Los Angeles County Museum of Natural History (LACM) and the
Museo Nacional de Nicaragua in Managua, Nicaragua (MNN). This
article thus presents a comprehensive review of knowledge of the fossil
mammals of Nicaragua and analyzes their significance for biostratigra-
phy, paleoecology, paleobiogeography and paleoanthropology.
PREVIOUS STUDIES
Knowledge of the fossil mammals of Nicaragua begins with Leidy
(1886), who described a small collection of Pleistocene mammals from
the northern part of the country. Nothing other than an abstract by
Howell and Macdonald (1969) was then published for nearly a century,
until Espinoza (1976) documented the excavation at the El Bosque local-
ity in northern Nicaragua, which he claimed established an association of
Pleistocene megafauna and human artifacts. In March-April 1982, Soviet
FIGURE 1. Generalized geologic map of Nicaragua (after Weinberg, 1992)
vertebrate paleontologist V. J. Reshetov visited Nicaragua to examine showing fossil mammal localities: 1 = Jalapa, 2 = El Bosque, 3 = Jinotepe, 4
vertebrate fossil localities and collections. His unpublished report = Sebáco, 5 = Matagalpa, 6 = Rio Viejo, 7 = Las Banderas, 8 = Masachapa, 9
(Reshetov, 1982) identified 15 sites, most of them of Pleistocene age. = El Palmar, 10 = Mine K-11.
However, as Reshetov noted, many of these localities were known only
from anecdotal information, and the fossils collected at most of them had
disappeared.
418
Lucas and Alvarado (1994) and Lucas et al. (1997, 2007) next
presented lists of the Pleistocene mammal taxa reported from Nicaragua,
based primarily on Espinoza (1976) and Reshetov (1982), though no
substantive documentation of these taxa had been published. Indeed,
other than Leidy (1886), no proper documentation (descriptions or illus-
trations) of any fossil mammals from Nicaragua has been published until
now.
GEOLOGICAL CONTEXT
Nicaragua is the largest nation in Central America, with a land area
of ~ 129, 500 km2. The country can be divided into four physiographic
provinces that correspond well to geological terrains (Fig. 1) (e.g.,
McBirney and Williams, 1965; Weyl, 1980; Weinberg, 1992; Ehrenborg,
1996; Marshall, 2007):
1. The Pacific coastal plain (the Sandino forearc of Marshall,
2007, fig. 3.1), primarily underlain by marine sediments of Cretaceous-
Miocene age that were deposited in the Nicoya basin. These rocks were
folded and eroded before being overlapped by Pliocene marine strata of
the El Salto Formation and by Pliocene volcanic rocks. The dominant
structural style is northwest-southeast trending folds that form low
coastal mountains.
2. The Nicaragua depression, a northwest-southeast-trending
strike-slip tectonic depression that is about 500 km long and 50 km wide
(e.g., Mann et al., 2007). Rocks filling the depression are up to two km
thick, and the exposed floor of the depression is covered by alluvial
sediments, an active chain of volcanoes and two great lakes—Lake
Managua and Lake Nicaragua—both of which drain into the Caribbean
via the Río San Juan.
3. The interior highlands (Chortis highlands of Marnn et al., 2007,
fig. 3.1), which is a mountainous region with elevations up to ~ 500 m
above sea level. The bedrock is primarily lavas, tuffs and ignimbrites of
the Oligocene Matagalpa Group and the Miocene-Pliocene Coyol Group.
4. The Atlantic coastal plain (Mosquito Coast lowlands of
Marshall, 2007, fig. 3.1), a tropical jungle primarily underlain by Quater-
nary alluvium.
Most of the fossil mammal localities known from Nicaragua are
from the Nicaragua depression. A few are from the interior highlands and
the Pacific coastal plain (Fig. 1).
FOSSIL LOCALITIES
Introduction
We have been able to confirm ten fossil mammal localities in Nica-
ragua (Fig. 1). Nine of these localities yield terrestrial mammal fossils of
Pleistocene age; one yields marine mammal fossils of Pliocene age.
Fossil Mammal Localities
1. Jalapa
The fossil mammals described by Leidy (1886) came from the
Jalapa (El Chorro) locality (Figs. 1, 2A). The site is at UTM 16P, 592348E,
1529984N (NAD 27) on the northeast bank of the small river called El
Chorro, southwest of Jalapa, in northernmost Nicaragua (Fig. 1). The
fossil bones at this site come from friable, crossbedded, pale yellowish
brown (10YR6/2), very fine to fine-grained micaceous sand that appears
to be a channel deposit of a Pleistocene river.
Leidy (1886) first reported fossil mammals from Jalapa that were
in the private collection of Mrs. Dr. B. F. Guerrero. Leidy identified the
sloth “Megatherium,” a proboscidean (“Mastodon andium”), the horse
Equus, the toxodont “Toxodon” (= Mixotoxodon) and a new species of
capybara, “Hydrochoerus (= Neochoerus) robustus.” Some of these
fossils (the horse, toxodont and capybara) are still housed in the ANSP
collection in Philadelphia (see below). Reshetov (1982, p. 9) reported
that in the MNN there are molar fragments of Stegomastodon from
Jalapa, but no fossils from Jalapa could be located in the MNN in 2001,
and we consider this record in error, as the Jalapa locality is late Pleis-
tocene, much younger than the known temporal range of Stegomastodon.
2. El Bosque
The most intensively collected and previously published Pleis-
tocene mammal locality in Nicaragua is the El Bosque locality, southwest
of Pueblo Nuevo (Figs. 1, 3). Discovered in 1974 and initially excavated
in 1975-1976, this site (Fig. 3) is on the western slope of a hill above the
Río Horcones at UTM 16P, 548923E, 1472989N (NAD 27). It covers
about 2000 m2 and is a lenticular deposit above Oligocene rhyolite tuff
(Page, 1978). Espinoza (1976), based on identifications by C. S. Churcher,
E. Lundelius and W. Miller, reported the following vertebrate taxa: the
megatheriid ground sloth Eremotherium (numerically dominating the as-
semblage), a megalonychid sloth, the gomphotheres Stegomastodon and
Gomphotherium, the horses Equus and Amerihippus santahelenae, the
deer Odocoileus, a toxodont, an aquatic turtle and a tortoise. None of
these identifications, however, was documented, and only fossils of the
sloth Eremotherium and a toxodont remain in the collection of the MNN.
Nevertheless, photographs provided to us by Wade Miller, who visited
the El Bosque site in 1974, also document the presence of the probos-
cidean Cuvieronius (see below).
At the El Bosque site, the fossil bones are in a green clay layer
below a colluvium of very pale orange (10YR8/2) and light brownish
gray (5YR6/1) rhyolitic boulders that range from 10 to 70 cm in diameter.
The bones are generally oriented approximately north-south, and the
colluvium appears to have derived from the hill immediately to the south-
east of the site. The bedrock of the hill (locally referred to as “Las
Mariposas”) is the same rhyolite as the boulders.
3. Jinotepe
South of Lake Aranás (north of Jinotepe), some proboscidean
postcrania were discovered while digging a water well but were subse-
quently lost. The bones came from a black clay at a depth of ~ 8 m at
UTM 16P, 605585E, 1450746N (NAD 27). The well was excavated in a
narrow valley in the mountain piedmont, so it seems likely that the
proboscidean bones were from a Pleistocene lake deposit.
4. Sebáco
The Sebáco (Palo Verde) locality is a man-made well just west of
the Río Grande de Matagalpa at UTM 16P, 590698E, 1412469N (NAD
27). The site is in a large flat valley, which we interpret as an ancient lake
bed. Bones of a mammoth (now lost) were exhumed in the well from a
bed of pale brown (5YR5/2) and medium dark gray (N4) sandy mud-
stone about 8 m below the surface.
5. Matagalpa
At Matagalpa, numerous bones and a molar of the proboscidean
Cuvieronius were collected from the bank of the Rio Viejo. These fossils
are documented in a photograph taken in 27 February 1940 by Omas
Medrano and stored in the MNN archive (see below), but the fossils are
apparently lost, and we have not relocated the fossil site.
6. Río Viejo
Howell and Macdonald (1969) published an abstract reporting
Pleistocene mammal bones found along the Río Viejo about 10 km west
of Ciudad Darío. They listed a toxodont, gomphothere, equid and Bison
at this locality. According to S. McLeod (written commun., 2007), this is
LACM locality 2025, and a small collection in the LACM (which we
have not studied) vouchers the taxa reported: LACM 19066, Equus,
right dentary w p2-m1; 19105, Bison, axis vertebra; 19106, Bison,
radius; 19107, Bison, cheek teeth; 19108, Proboscidea femur fragment;
and 19109, Proboscidea tooth fragment. We thus consider this a con-
firmed record of Bison sp. from Nicaragua.
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FIGURE 2. Photographs of selected fossil mammal localities in Nicaragua. A, Jalapa locality; fossils come from the light-colored fluvial sands in the
foreground along the bank of the Río Jalapa. B, Sea cliff at Masachapa locality; fossils are collected on the beach surface after they weather out of the cliff-
forming volcaniclastic sandstone. C-D, Overview of Las Banderas locality (C) and close-up of ignimbrite (D); the locality is an open pit mine where the
tuff is quarried for building stone. E, El Palmar locality in volcaniclastic sand along the bank of the Río Chacalapa. F, Mine K-11 locality, an abandoned
open-pit mine in shaly strata of the Pliocene El Salto Formation.

7. Las Banderas
The Las Banderas locality is an open pit mine (Figs. 2C, 4) just
west of the village of that name at UTM 16P, 612758E, 1362070N
(NAD 27), ~ 40 km east-northeast of Managua (Fig. 1). The pit is about
22 m deep and exposes a thick succession of ash-flow tuff (ignimbrite)
overlain by gravelly, volcaniclastic alluvium. The tuff has a pale yellow-
ish brown (10YR6/2) matrix and clasts of pinkish gray (5YR8/1) and
light brownish gray (5YR6/1) pumice and medium dark gray (N4) rhyo-
lite (Fig. 2D). It is mined for construction materials. The alluvium has a
matrix of pale brown (5YR5/2), medium-to very coarse-grained basaltic
sand. A fossil procyonid was collected from the tuff at the quarry floor in
1999, whereas mammoth bones are known from the overlying alluvium.
The bone-bearing tuff at the Las Banderas locality belongs to the
Las Banderas subunit of the Las Sierras unit of Ehrenborg (1996, 1999).
This is a pyroclastic flow unit that erupted from the Las Sierras volcanic
center during the Pleistocene.
420
8. Masachapa
The Masachapa locality is at the base of a sea cliff (Fig. 2B) at
UTM 16P, 552660E, 1302903N (NAD 27). The bone-bearing stratum is
a medium light gray (N6) and grayish orange (10YR7/4) coarse-grained to
pebbly basaltic sand. Bones weather from this alluvial deposit, which is
6-10-m thick and trough crossbedded, and wash onto the adjoining beach.
Specimens from the Masachapa locality belong to the ground sloth
Eremotherium and to Mammuthus columbi (see below). Espinoza (1976)
and Reshetov (1982) mention fossil bones from Matagalpa, and these
may have been from the same locality we document.

9. El Palmar
The El Palmar (Las Moras) locality is in southern Nicaragua north
of the highway between Rivas and Tola in the drainage of the Río Chacalapa
(Figs. 1, 2E). The site is in the western bank of San Luis Creek at UTM
16P, 619661E, 1266622N (NAD 27). A farmer and his son discovered
bones of a mammoth here and excavated them from a 6 x 2 m trench. The
fossil-bearing stratum is a friable sand that is a medium gray (N5) and
light brownish gray (5YR6/1), medium to very coarse grained basaltic
sand. These beds are trough cross bedded and contain imbricated pumice
clasts up to 15 cm in diameter; crossbeds and imbrication indicate south-
erly paleoflow of this volcaniclastic alluvium.

10. Mine K-11

FIGURE 3. Map of the 1970s excavation at the El Bosque locality (from
Espinoza, 1976, mapa 3).
Mine K-11 is a large, abandoned open pit mine (Figs. 2F, 4) south
of San Rafael del Sur at UTM 16P, 564899E, 1305906N (NAD 27). Two
lithologic units of the El Salto Formation are exposed at the pit: light
olive gray (5Y6/1) to grayish yellow green (5GY7/2) smectitic shale
overlain by oyster-bearing very pale orange (10YR8/2) and pale yellow-
ish orange (10YR8/6), hematitic coquinoid limestones and bioclastic sands.
Part of a fossil whale collected from the El Salto Formation at Mine K-11

FIGURE 4. Measured stratigraphic sections at the Las Banderas (left) and Mine K-11 (right) localities.

is on display at the Museo Comunitario de San Rafael del Sur. The El
Salto Formation has long been assigned an Early Pliocene age based on its
molluscan fossils (e.g., Woodring, 1973; Weyl, 1980; Kirby and Jackson,
2004), but more precise age data are not available.
Other Possible Localities
Reshetov (1982) reported several other fossil mammal localities
in Nicaragua, but none of these are documented by fossils or have been
successfully relocated. They are: Valle de Río Mayuca, Región de Santa
Barbara, El Recreo (a footprint locality very close to Acahualinca), Ciudad
Darío, El Hato (cetacean), San Juan del Sur (cetacean), Montelimar
(glyptodont, Holmesina, mammoth, horse) and Departamento de Zelaya.
Acahualinca Footprint Site
The human footprint site that is now the Acahualinca Tracks
Museum (Huellas de Acahualinca), and nearby footprint sites are in the
suburbs of Managua. Discovered in 1874, an extensive literature exists
on these sites (Flint, 1884, 1886, 1888, 1889, 1890; Johnson, 1884;
McA, 1886; Brinton, 1887; Peet, 1889, 1891; Crawford, 1891; Brown,
1947; Williams, 1952; Bryan, 1973; Bice, 1979; Weyl, 1980) and is
reviewed in detail by Lockley et al. (2007). The tracks are in volcaniclastic
strata deposited between 5800 and 6500 yBP and include those of hu-
mans, opposum, tapir, bison and bird. Although they are not old enough
to be considered fossil, we mention these footprints here because of their
longstanding notoriety as one of the most extensive human footprint
sites known (Lockley et al., 2007).Furthermore, they establish the pres-
ence of Bison in Nicaragua during the Holocene.
FOSSIL TAXA
Eremotherium laurillardi
Espinoza (1976) reported the giant ground sloth Eremotherium
from El Bosque, and we also document its presence at Masachapa. Leidy
(1886, p. 275) reported “Megatherium” from Jalapa based on “the greater
part of the distal extremity of a femur and a fragment of the mandible
with two teeth.” This may also be a record of Eremotherium, but the
specimens Leidy mentioned were never catalogued into the ANSP collec-
tion and cannot be located.
The material of Eremotherium from El Bosque is the left and right
femora (Fig. 5A-B), part of a right humerus, incomplete right ulna and
complete right radius (Fig. 5C), all possibly of a single individual, in the
MNN collection. Selected measurements of these bones are: femur length
= 670, diameter of head = 160, distal width = 260, radius length = 648,
proximal width = 48 and distal width = 158. Also, a distal left humerus of
a giant ground sloth is known from the Masachapa locality (Fig. 5D).
These bones closely resemble those of Eremotherium carolinense illus-
trated by Hoffstetter (1952, figs. 8, 9, 12).According to Cartelle and
DeIuliis (1995), all Late Pleistocene Eremotherium belong to the species
E. laurillardi, so we assign the Nicaraguan specimens to that species.
Procyonidae
A partial skull, lower jaws and postcranium of a procyonid carni-
vore were collected at the Las Banderas locality and are in the MNN
collection. This fossil represents a new taxon that will be described
elsewhere.
Neochoerus aesopi (Leidy)
Leidy (1886, p. 275, fig.1) coined the name Hydrochoerus robustus
for a left dentary fragment with an incomplete p4 (ANSP 12109) from
Jalapa (Fig. 6A-C). The p4 length = 11.4 mm. Subsequent authors reas-
signed this species to Neochoerus (Hay, 1926; Kraglievich, 1930; Spamer
et al., 1995). In his taxonomic revision of capybaras, Mones (1991, p.
60) considered Neochoerus robustus to be a junior subjective synonym
of Neochoerus aesopi. N. aesopi is a common late Pleistocene
421
(Rancholabrean-Lujanian) capybara known from Argentina, Ecuador, Peru,
Venezuela, Nicaragua, Guatemala, Mexico and the southeastern USA
(Mones, 1991).
Mixotoxodon larensis Van Frank
Espinoza (1976) reported a toxodont from El Bosque, which prob-
ably is a record of the common Central American/northern South Ameri-
can toxodont Mixotoxodon larensis. Indeed, a right femur and incomplete
innominate in the MNN collection (Fig. 5E-G) from the El Bosque local-
ity belong to a toxodont. These bones are relatively large (femur length =
470), and the femur has a ball-shaped head on a distinct neck and lacks a
third trochanter. The resemblance to Toxodon (e.g., Cope, 1897; Hopwood,
1928; Paula Couto, 1979, fig. 454) is striking, but it is most likely these
bones belong to the common Central American Pleistocene toxodont
Mixotoxodon, for which the innominate and femur are unknown (Van
Frank, 1957).
The only previously documented record of M. larensis from Nica-
ragua is the teeth Leidy (1886, figs. 2-3) illustrated from Jalapa and
assigned by him to Toxodon burmeisteri. These teeth are catalogued as
ANSP 12110 and are tooth fragments, an incisor fragment and a left m2
(Fig. 6D-F). Measurements of the m2 are: length = 44.9 mm, trigonid
width = 18.2 mm and talonid width = 15.9 mm. The teeth from Jalapa are
identical in morphology and size to teeth of M. larensis described and
illustrated by Van Frank (1957), Laurito (1993) and Lucas et al. (1997).
Cuvieronius hyodon
Records of the gomphothere Cuvieronius hyodon are surprisingly
rare in Nicaragua, whereas this proboscidean dominates the Pleistocene
mammal record of adjacent Costa Rica (Laurito, 1988; Lucas et al., 1997).
Leidy (1886) reported “Mastodon andium” from Jalapa, which is a taxon
now assigned to C. hyodon. A molar fragment from El Bosque (Fig. 7A)
and the material from Matagalpa (Fig. 8) are readily referred to C. hyodon.
Espinoza’s (1976) reports of Gomphotherium and Stegomastodon from
El Bosque may be based in part on the molar fragment illustrated here;
they are, nevertheless, unlikely, given the late Pleistocene age of the El
Bosque deposit. Thus, only one gomphothere is known from Nicaragua,
C. hyodon.
Mammuthus columbi
The partial mammoth skeleton from El Palmar in the MNN col-
lection (briefly mentioned by Laurito and Aguilar, 2007, p. 79) consists
of the lower jaw (Fig. 7E-G), the glenoid portions of both scapulae, both
humeral heads, the proximal ends of both ulnae, several vertebral centra,
many rib fragments, some carpals and an incomplete femur. The lower
jaw contains the left and right m3s. The rounded and vertical symphysis,
relatively short and thick horizontal ramus and short, vertical ascending
ramus with its coronoid process well above the tooth are characteristic of
Mammuthus columbi. Dental measurements of the left m3 are: length =
300+ mm, width = 95 mm, number of plates = 21+, plate ratio (number
of plates/100 mm) = 6 and average enamel thickness = 2.8. These mea-
surements fall in the range of variation of North American M. columbi,
but they also overlap the measurements of advanced M. imperator (Mad-
den, 1981; Agenbroad, 1994). However, given the advanced dentary
structure, relatively short and broad m3 and relatively thin m3 enamel,
the El Palmar mammoth jaw is best assigned to M. columbi.
A second Nicaraguan record of Mammuthus columbi is of fossils
from Masachapa. These are a tusk fragment (Fig. 7B) and incomplete
M3 (Figs. 7C-D). The M3 is 103 mm wide, has 5-6 plates per 100 mm
and an enamel thickness of ~ 3.8 mm.
Mammuthus columbi (the Columbian mammoth) was the com-
mon late Pleistocene mammoth in North America (Agenbroad, 1994). It
stood 3.6-4.0 meters at the shoulder, and was a grazer that lived in and
around grasslands and savannahs (Kurtén and Anderson, 1980). All
Mammuthus records in Central America are of this species, and are from
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FIGURE 5. Sloth, toxodont and horse fossils from Nicaragua. A-C, Eremotherium sp. from the El Bosque locality, right femur in anterior (A) and posterior
(B) views and right radius (C). D, Eremotherium sp. from the Masachapa locality, distal end of left humerus. E-G, Toxodont from the El Bosque locality,
incomplete right femur in anterior (E) and posterior (F) views and innominate (G) in lateral view. H-J, ANSP 11497, Equus cf. E. occidentalis from the
Jalapa locality, left M1-2 in occlusal (H), anterior (I) and posterior (J) views. Scale bars = 2 cm.

FIGURE 6. Capybara and toxodont fossils from Nicaragua. A-C, ANSP 12109, holotype of Hydrochoerus robustus Leidy, 1886, from Jalapa, left dentary
fragment with incomplete p4 in occlusal (A), labial (B) and lingual (C) views. D-F, ANSP 12110, left m2 of Mixotoxodon larensis. Scale bars = 1 cm.
Guatemala, Honduras, El Salvador, Nicaragua and Costa Rica (Lucas et
al., 1997, 2007). Clearly, M. columbi was a taxon that originated in North
America and immigrated into Central America during the late Pleistocene.
The El Palmar and Masachapa mammoths thus suggest a late Pleistocene
age for the sediments that contained them and the presence of nearby
open country habitats, either grasslands or savannas. Indeed, the relative
rarity of the common Central American gomphothere Cuvieronius and
relative abundance of Mammuthus may suggest a greater prevalence of
savanna in western Nicaragua during the late Pleistocene than in other
Central American countries.
Equus cf. E. occidentalis
Leidy (1886) first reported two upper molars of Equus from
Jalapa (Fig. 5H-J). The teeth are M1-2 (ANSP 11497) and have rela-
tively simple fossettes. Measurements (in mm) are M1 length = 24,
width = 28.1, M2 length = 26.7, width = 26.9. These molars compare
well in morphology and size to the large Rancholabrean horse Equus
occidentalis (e.g., Merriam, 1913, figs. 7-8; Harris and Porter, 1980).
However, the two molars from Jalapa are not sufficient to determine a
species-level identification with certainty, so we only identify them as E.
cf. E. occidentalis.
Espinoza (1976) reported Equus and Amerihippus stantahelenae
from El Bosque. However, no horse fossils are available to us from El
Bosque. Equus is also known from the Río Viejo locality (LACM collec-
tion).
423
Artiodactyla
Leidy (1886) reported horn cores of an “ox” from Jalapa, but the
specimens were never catalogued into the ANSP collection and cannot be
located. Espinoza (1976) reported Odocoileus from El Bosque, and
Reshetov (1982) reported bison from a locality he termed El Horno, but
no specimens can be located to document these reports. Howell and
Macdonald (1969) reported Bison sp. from Rio Viejo, for which diagnos-
tic material exists in the LACM collection (see above). It seems highly
likely that fossils of cervids should be found in the Nicaraguan Pleis-
tocene, but no extant specimens document the published reports.
Cetacea
The only available cetacean fossil from Nicaragua is an incomplete
skeleton (Fig. 9) from Mine K-11. It consists of 14 vertebral centra,
parts of 10 ribs and a right humerus, all on display in the Museo
Comunitario de San Rafael del Sur. According to Larry Barnes (written
commun., 2007) this is the partial skeleton of a mysticete. Measure-
ments of the specimen are in Table 1.
PLEISTOCENE MAMMAL-HUMAN
ASSOCIATION AT EL BOSQUE
The El Bosque locality (Figs. 3, 10-11) received much attention in
the mid-1970s because of the proposed association of the Pleistocene
animal bones at the locality with human artifacts (stone tools) (Espinoza,
1976). Given recent evidence of Pleistocene (pre-Clovis = pre-11,500
424

FIGURE 7. Selected proboscidean fossils from Nicaragaua. A, Occlusal view of incomplete molar (now lost) of Cuvieronius hyodon from El Bosque
(courtesy of Wade Miller). B-D, Mammuthus teeth from Masachapa in the MNN collection, tusk fragment (B) and incomplete M3 in lateral (C) and
occlusal (D) views. E-G, Lower jaw of Mammuthus columbi from El Palmar in the MNN collection, occlusal view of left m3 (E), antero-occlusal view of
entire jaw (F) and lateral view of jaw (G). Scale bars = 2 cm, except for E-F, which have a scale in cm.

FIGURE 8. Photograph in the MNN archive taken 27 February 1940 by
Omas Medrano of some fossils from the Matagalapa locality. Note
proboscidean fossils (now lost) on floor that include a complete m2 and
other molar fragments of Cuvieronius hyodon (arrow).
TABLE 1. Measurements of vertebrae of the Mine K-11 cetacean, based on
their alignment in the exhibit (in mm). length = antero-posterior length of
centrum.
yBP) human occupation at Monteverde in Chile and other sites in the
Americas (e.g., Meltzer, 1997, 2004; Marshall, 2001; Sarnthein et al.,
2006), the possibility of a Pleistocene human presence at El Bosque
should be re-evaluated. Also, these recent finds add plausibility to simi-
lar long-claimed associations of artifacts and extinct mammal fossils at
425
FIGURE 9. Fossil of a mysticete cetacean from the Pliocene El Salto
Formation at Mine K-11. A, Vertebrae and ribs on display. B-F, Vertebral
centra. G-H, Humerus, Scale bars = 3 cm.
sites in México, such as Tlapacoya (Mirambell, 1967) and Valsequillo
(Irwin-Williams 1967), among others (cf. MacNeish and Nelken-Turner
1983).
Page (1978) presented a very detailed study of the geology of the
El Bosque locality, documenting that the site is developed on a bedrock
tuff of pink welded rhyolitic ignimbrite andesite of the mid-Cenozoic
(Oligocene-Miocene) Coyol Group. Locally, this andesite has veins of
chert (jasper). At the fossil site a lenticular deposit of green clay (Fig. 11)
contains the bones and can be divided into two layers: (1) a lower, 1-2-m-
thick clay bed that contains most of the bones, some sandstone pebbles
and the chert/jasper “artifacts” (Fig. 12); and (2) an overlying clay bed,
0.25-1 m thick, that is mixed with andesite boulders. The overlying
colluvium consists of a layer of andesitic boulders 0.4-1 m thick (Fig.
10C) capped by up to 0.6 m of dark brown to grayish silty and sandy
clay (Fig. 11) interpreted by Page to be a soil horizon. Radiocarbon dates
on the bone and carbonate nodules in the deposit suggest a minimum age
of 32,000 yBP, and geomorphic considerations are consistent with dat-
ing the El Bosque bonebed as pre-late Wisconsinan (older than 30,000
yBP) (Page, 1978). However, it should be noted that such an age is twice
that of the older dates from Monteverde (~ 15000 BP), which is appar-
ently the oldest archeological site in the New World.
Page (1978) inferred a sequence of Late Pleistocene-Holocene
events (Fig. 13) to develop the deposit at El Bosque: (1) a landslide into
nearby Los Horcones Creek dammed drainage, forming a small pond in
which the green clays were deposited; the bones and supposed artifacts
were buried in the clays; (2) the downcutting of Los Horcones Creek re-
established drainage, and a boulder colluvium buried the lake deposit; and
(3) further downcutting and erosion produced the current topography at
the site.
With regard to the supposed artifacts, Page (1978, p. 257) reached
no definite conclusion as to their origin. In assessing the petrography of
the artifacts, he noted that “the different types of jasper found in the
deposit suggest that early man possibly carried some of the material to
the site.” In contrast, Page (1978, p. 254), noting that the so-called
artifacts are in gravelly beds within the clay layer, stated that “it is also
possible that the gravelly materials found with the bones had their origin
in a sheet wash deposit and that the jasper artifacts are natural in origin.”
Indeed, Gruhn (1978, p. 261) noted that the “flaked” pieces of chert
were found “banked against boulders,” implying that the shape of the
426
FIGURE 10. The El Bosque locality in 2001. A, Overview of the structure
built to protect the bonebed. B, View inside the structure; plastic sheeting
covers the active excavation surface. C, Rhyolitic boulder layer that covers
the bone-bearing clays at the site. Rod is 1 m long.
chert pieces could be due to inorganic processes. It should also be added
that natural sources of jasper similar to those from El Bosque are com-
mon along ravines near the site in the Mesetas de Estelí region. Wherever
a creek washes out these deposits it produces large amounts of naturally-
flaked debris. Given that this material is quite brittle, it tends to chip
easily along the edges. This particular jasper might be useful to produce
cutting tools as long as the user does not require these to perform some
heavy task. Burins and scrapers would be impractical and choppers and
hammers ineffective.
FIGURE 11. Selected cross sections of the main trench of the 1970s
excavation at El Bosque (from Page, 1978, figs. 7A and 7C).
Aside from the radiocarbon dates from El Bosque covering a time
range of 18000-35000 yBP, which largely exceeds the best evidence on
the antiquity of human presence in the Americas, the lithic specimens
recovered by Espinoza and presented as artifacts do not fit, morphologi-
cally or technologically, extant criteria on the manufacture and use of
lithic artifacts pertaining to the earliest stages of New World peopling.
MacNeish and Nelken-Turner (1983, p. 73), while considering the El
Bosque site as a reliable representative of an extremely old “Stage I”,
note that only a chopper was found in association with the remains of
extinct fauna. Indeed, MacNeish had the opportunity to examine a selec-
tion of the lithic specimens taken (by Espinoza) to an international
meeting in 1974, and concluded that none of them appeared to be arti-
facts. Most were pebbles and small nodules with some erratically lo-
cated scars left by detached flakes, the probable result of violent alluvial
transportation. Further, none of the specimens suggested “any given
cultural function.”
Thus, various authors have commented that the “tools” at the El
Bosque site are neither morphologically nor technically similar to human

FIGURE 12. Supposed stone artifacts from the El Bosque excavation (from
Gruhn, 1978).
artifacts (e.g., Hurtado de Mendoza and Alvarado, 1988; Alvarado, 1994).
Therefore, the “tools” at el Bosque are just chert broken by physical
processes, not artifacts manufactured by humans. We thus conclude that
there is no demonstrable association of human artifacts and the Pleis-
tocene megafauna at the El Bosque locality.
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CONCLUSIONS
Nicaragua is the largest country in Central America, yet it has the
least extensive vertebrate fossil record of any Central American country.
Other than the cetacean fossils from mine K-11, all the fossil mammals
from Nicaragua appear to be of late Pleistocene age. Certainly, all post-
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The Nicaraguan fossil record of Pleistocene mammals is meager,
but a few preliminary observations can be offered: (1) like other Central
American Pleistocene mammal records, a microfauna, especially of ro-
dents, is essentially absent and needs to be a goal of further fieldwork; (2)
also like other Central American Pleistocene mammal records, the Pleis-
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common Central American gomphothere Cuvieronius and relative abun-
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western Nicaragua during the Pleistocene than in other Central American
countries.
ACKNOWLEDGMENTS
The work of Lucas, Alvarado and Vega in Nicaragua was made
possible by the generous support of the Museo Nacional de Nicaragua
through then Director Clemente Guido Martínez, and by the assistance
of Frederick Lange and his family. Francisco Arias helped in the field.
Sam McLeod and Wade Miller generously supplied unpublished infor-
mation. Larry Barnes provided comments on the fossil whale from Mine
K-11. Ted Daeschler made access to the ANSP collection possible. Greg
McDonald, Gary Morgan and Richard White provided helpful reviews
of the manuscript.
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FIGURE 13. Inferred sequence of geological events at the El Bosque locality (from Page, 1978, fig. 15).

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