LATE CENOZOIC VERTEBRATE FOSSIL ASSEMBLAGES FROM JALISCO, MEXICO

Abstract

Fossil vertebrates (fishes, amphibians, reptiles, birds and, mostly, mammals) of Plio-Pleistocene age are known from 23 localities/collecting areas in the Mexican state of Jalisco. Almost all of the localities are in alluvial, fluvial or lacustrine strata deposited in the three extensional basins of the Jalisco triple junction, the Tepic-Zacoalco, Colima and Chapala rifts. The majority of the localities (12 of 23) are records of proboscideans, mostly of late Pleistocene Mammuthus (Acatlán de Juarez, Cocula, Ejutla, El Tule, Guadalajara, Palo Dulce [Atotonilco el Alto], Río de Juchipila, San Martin Hidalgo [El Tepehuaje], Santa Catarina, Tamazula, Tapalpa and Tecuitatlán). One site is a fossil fish record (Barranca de Santa Rosa), whereas another (San Gabriel) is a record of the giant ground sloth Eremotherium laurillardi. Two localities yield low diversity Hemphillian mammal assemblages (Los Corrales, Teocaltiche), and two sites yield moderately diverse but largely undocumented late Pleistocene (Rancholabrean) mammal assemblages (Ameca, Atotonilco El Bajo), whereas a third site (San Juan de los Lagos) yields fossil cat (Felipeda) and bird (Avipeda) footprints. Only three areas yield diverse fossil mammal assemblages and a stratigraphic succession of fossils that represent more than one land-mammal " age: " Lago de Chapala, Tecolotlán (including nearby Juchitlán and Colotlán) and Zacoalco. The Chapala succession includes Blancan, Irvingtonian? and Rancholabrean mammal assemblages, whereas in the Tecolotlán area Hemphillian and Blancan assemblages are present; the Hemphillian assemblage is associated with an ~4.9 Ma radioisotopic age. At Zacoalco, Irvingtonian? and Rancholabrean mammal assemblages are present. There is no demonstrable human association with Pleistocene mammals in Jalisco. Much remains to be published on the Plio-Pleistocene vertebrate fossils of Jalisco, particularly with regard to careful documentation of the fossils and their stratigraphic context. However, the assemblages are characteristic of southern North American mammal assemblages both prior to and after the " Great American biotic interchange. " INTRODUCTION

Full text

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Lucas et al., eds., 2008, Neogene Mammals. New Mexico Museum of Natural History and Science Bulletin 44.
LATE CENOZOIC VERTEBRATE FOSSIL ASSEMBLAGES FROM JALISCO, MEXICO
SPENCER G. LUCAS
New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104
Abstract—Fossil vertebrates (fishes, amphibians, reptiles, birds and, mostly, mammals) of Plio-Pleistocene age
are known from 23 localities/collecting areas in the Mexican state of Jalisco. Almost all of the localities are in
alluvial, fluvial or lacustrine strata deposited in the three extensional basins of the Jalisco triple junction, the Tepic-
Zacoalco, Colima and Chapala rifts. The majority of the localities (12 of 23) are records of proboscideans, mostly
of late Pleistocene Mammuthus (Acatlán de Juarez, Cocula, Ejutla, El Tule, Guadalajara, Palo Dulce [Atotonilco el
Alto], Río de Juchipila, San Martin Hidalgo [El Tepehuaje], Santa Catarina, Tamazula, Tapalpa and Tecuitatlán).
One site is a fossil fish record (Barranca de Santa Rosa), whereas another (San Gabriel) is a record of the giant
ground sloth Eremotherium laurillardi. Two localities yield low diversity Hemphillian mammal assemblages (Los
Corrales, Teocaltiche), and two sites yield moderately diverse but largely undocumented late Pleistocene
(Rancholabrean) mammal assemblages (Ameca, Atotonilco El Bajo), whereas a third site (San Juan de los Lagos)
yields fossil cat (Felipeda) and bird (Avipeda) footprints. Only three areas yield diverse fossil mammal assem-
blages and a stratigraphic succession of fossils that represent more than one land-mammal “age:” Lago de Chapala,
Tecolotlán (including nearby Juchitlán and Colotlán) and Zacoalco. The Chapala succession includes Blancan,
Irvingtonian? and Rancholabrean mammal assemblages, whereas in the Tecolotlán area Hemphillian and Blancan
assemblages are present; the Hemphillian assemblage is associated with an ~4.9 Ma radioisotopic age. At Zacoalco,
Irvingtonian? and Rancholabrean mammal assemblages are present. There is no demonstrable human association
with Pleistocene mammals in Jalisco. Much remains to be published on the Plio-Pleistocene vertebrate fossils of
Jalisco, particularly with regard to careful documentation of the fossils and their stratigraphic context. However,
the assemblages are characteristic of southern North American mammal assemblages both prior to and after the
“Great American biotic interchange.”
INTRODUCTION
The Mexican state of Jalisco is at the western end of the Mexican
volcanic belt, a mostly calc-alkaline continental arc associated with a
subduction zone. The western edge of the arc is marked by three linea-
ments. Most of the state west of the capital city of Guadalajara is the
Jalisco block, a distinct crustal unit that has been rifting away from the
Mexican mainland since the Miocene as part of a complex plate-bound-
ary reorganization (e.g., Allan, 1986; Henry, 1989; Allan et al., 1991;
Henry and Aranda-Gomez, 1992; Ferrari, 1995; Rosas-Elguera et al.,
1996; Ferrari and Rosas-Elguera, 2000; Ferrari et al., 2000). The northern
part of Jalisco is the southern margin of the Sierra Madre Occidental
volcanic province and consists of ~1500 m of silicic ash-flow tuffs and
lavas of Oligo-Miocene age.
The boundaries of the Jalisco block are the northwest-trending
Tepic-Zacoalco rift, the east-trending Chapala rift and the north-trending
Colima rift (Allan, 1986). The intersection of these three rifts forms a
triple junction at an elevation of 1350 m about 50 km south-southwest of
Guadalajara. These flat-floored grabens of the Jalisco triple junction have
been the sites of shallow and ephemeral lakes since at least the Pliocene.
The Colima rift is a Plio-Pleistocene rift of the east-trending Mexican
volcanic belt. Its basin fill is largely Pleistocene sediments, and little
effort appears to have made to relate these strata to coeval volcanism.
The Tepic-Zacoalco rift is a series of pull-apart basins and grabens. The
Chapala rift is the site of Mexico’s largest natural freshwater lake, Lago
de Chapala.
This article reviews the late Cenozoic vertebrate fossil record of
Jalisco. In part, it presents records listed by Solorzano (2002), an un-
published report on file with the INAH Museo Regional de Guadalajara.
It is also based on the published literature and on the largely unpublished
vertebrate fossil collections in the INAH Museo Regional de Guadalajara
and the Museo de Paleontologia de Guadalajara. This review identifies
vertebrate fossil assemblages of Hemphillian, Blancan, Irvingtonian? and
Rancholabrean age in Jalisco.
Institutional abbreviations and measurements: AMNH =
American Museum of Natural History, New York; INAH-MRG =
Instituto Nacional de Antropologia e Historia-Museo Regional de
Guadalajara; LACM = Los Angeles County Museum of Natural His-
tory; MPG = Museo de Paleontologia de Guadalajara. All measurements
of fossils in the text are in mm.
LOCALITIES AND VERTEBRATE FOSSIL ASSEMBLAGES
Introduction
I identify 23 localities/collecting areas that yield fossil vertebrates
in Jalisco (Fig. 1). Here, I review the vertebrate fossil taxa reported from
these sites and document some records based on specimens in the INAH-
MRG and MPG collections (Figs. 2-17).
Acatlán
Solorzano (2002) reports both Mammuthus columbi and M.
imperator from this locality. However, I cannot locate publications or
specimens that voucher these reports, so I consider them to indicate the
likely presence of Mammuthus sp.
Ameca
In addition to fish fossils (Cyprinidae and Siluriformes), Solorzano
(2002) reports a variety of mammals from Ameca: Glyptotherium mexi-
canum, G. cylindricum, Geomys sp., Muridae, Sciurus sp., Smilodon
sp., Equus conversidens, E. mexicanus, E. occidentalis, E. sp., Mam-
muthus columbi, M. imperator, M. sp. and Mammutidae. However, most
of these taxa have not been documented.
Brown (1912a) described the collecting of a complete glyptodont
carapace in 1910 from Pleistocene gravels on the San Miguel Ranch ~ 19
km west of Ameca. Brown (1912b) designated the specimen (AMNH
15548) the holotype of Brachyostracon cylindricus. He also reported
(but did not document) the following taxa from the Ameca locality:

52
siluroid and cyprinid fishes, Sciuridae, Geomyidae, Cricetidae, a
machaerodont, Equus and “Elephas” columbi. Gillette and Ray (1981)
reassigned B. cylindricus to Glyptotherium and regarded the species G.
cylindricus as endemic to the Ameca locality.
In the INAH-MRG collection, uncatalogued specimens include a
lower jaw, upper tusk, hyoid and cervical vertebrae of the gomphothere
Cuvieronius hyodon from Ameca (Fig. 2). INAH-MRG records indicate
that these fossils represent a single individual collected at Puerta la Vega.
The tusk has been reconstructed but has a slightly curved enamel band.
The lower jaw lacks tusks and has a spout-shaped symphysis. The m3s
have only four lophids and are metrically well within the range of C.
hyodon (Table 1). The hyoid is a tuning-fork-shaped bone with a basi-
hyal that is cylindrical in cross section and that has a blunt and slightly
rugose median process. The fork into the thyrohyals is asymmetrical,
with one branch being thin, curved and having an oval cross section,
whereas the other (incomplete) branch is thicker, with a nearly rectangu-
lar cross section.
Alberdi and Corona-M. (2005, fig. 2c-d) referred the Ameca tusk
to Rhynchotherium, but it is not diagnostic of the genus and its associa-
tion with the lower jaw suggests it is a poorly-restored upper tusk of
Cuvieronius. Alberdi and Corona-M (2005, fig. 4a) assigned the Ameca
lower jaw to Stegomastodon, but the relatively simple m3 crowns with
only four lophids support assignment to Cuvieronius hyodon.
Atotonilco El Bajo
In addition to a fish (Ictalurus spodius: Smith, 1987), Solorzano
(2002) reports the following mammals from Atotonilco El Bajo: Sigmodon
hispidus, Oryzomys couesi, Ondatra nebracensis, Liomys irroratus,
Cratogeomys gymnurus, Hydrochoeridae, Mammuthus imperator, M.
sp., Mammutidae, Equidae, Cervidae, Bison sp. and Camelidae, appar-
ently based on the published reports of Delgado (1962) and MacDonald
(1962). However, I can only find documentation of the proboscidean
records.
MacDonald (1962) and Delgado (1962) described the excavation
of incomplete mammoth skeletons and other fossils found at Atotonilco
El Bajo near Acatlán. They reported that fossils of capybara, other
rodents, horse, camel, deer, bison, mammoth and mastodon were col-
lected, and Delgado (1962, fig. 7) illustrated a left dentary fragment with
m3 that appears to be Mammut americanum. However, all I could locate
of the collection are two incomplete and not very accurately restored
mounted mammoth skeletons on display in the library of the Universidad
Autonoma de Guadalajara (Fig. 3). I was unable to obtain metric data
from the molars of these specimens, but the relatively large number of
plates, thin enamel and lower jaw morphology (short, vertical symphy-
sis, deep horizontal ramus) support identification as Mammuthus columbi.
Barranca de Santa Rosa
Solorzano (2002) reports the fish Tapatia occidentalis, but no
other data are available.
Chapala
Downs (1958) first listed fossil vertebrates from the Chapala
basin, mostly from the floor of Lake Chapala. Alvarez (1965) and Silva-
Barcenas (1969) repeated this listing. Silva (1969) referred to the
“Zacoalco-Chapala” fauna of 14 genera and 10 species of mammals as
coming from the “Gran Canal Formation” and being of Yarmouthian-
Illinoisan age.
Rufolo (1998) presented a detailed study of the Chapala basin
mammal fossils in the LACM collection. In this volume, Lucas (2008b)
reviews the fossil vertebrate assemblage from the Chapala basin, which,
in addition to fishes, amphibians, turtles, snake, crocodile and birds,
includes a variety of mammals of Blancan, Irvingtonian? and (mostly)
Rancholabrean age.
Cocula
Solorzano (2002) reports Mammuthus columbi and M. imperator
from this locality. However, I cannot locate publications or specimens
that voucher these reports, so I consider them to indicate the likely
presence of Mammuthus sp.
Colotlán
Besides turtle and crocodile, a Hemphillian assemblage of mam-
mals from near Colotlán is listed in a series of articles by Carranza-
Castañeda and Miller (1996, 1998, 2000, 2002, 2004; Miller and Carranza-
Castañeda, 1998, 2001; Carranza-Castañeda, 2006). These are
Neohipparion eurystyle, Astrohippus stockii and “Desmathyus” sp. of
late Hemphillian age.
Ejutla
Solorzano (2002) reports Mammuthus sp. from Ejutla. I cannot
locate publications or specimens that voucher this report, and I consider
it to indicate the likely presence of Mammuthus sp.
El Tule
Solorzano (2002) reports Mammuthus imperator, M. sp. and
Mammutidae from El Tule. However, I cannot locate publications or
specimens that voucher these reports, so I consider them to indicate the
likely presence of Mammuthus sp.
Guadalajara
Osborn (1922, p. 4; 1942, p. 1001) described and illustrated a
mammoth molar from Guadalajara that he designated the neotype of
FIGURE 1. Map of Jalisco showing principal vertebrate fossil localities
discussed in the text. Localities are: 1 = Acatlan, Atotonilco el Bajo, Santa
Catarina and Zacoalco; 2 = Ameca; 3 = Chapala; 4 = Cocula; 5 = Ejutla; 6 =
Guadalajara; 7 = Juchitlan; 8 = Los Corrales; 9 = Mezcala; 10 = San Gabriel;
11 = San Juan de los Lagos; 12 = Tamazula; 13 = Tapalpa; 14 = Tecolotlán;
15 = Teocaltiche. Sites in the text that could not be located precisely are not
indicated here.

53
Archidiskodon imperator. Alvarez (1965), Silva-Barcenas (1969) and
Solorzano (2002) repeated this report as Mammuthus imperator. The
specimen is AMNH 11871, an incomplete right M3 (Fig. 4), here also
assigned to M. imperator.
Juchitlán
Silva-Barcenas (1969) listed Asinus mexicanus, Equus sp. and
Rhinocerotidae from Juchitlán. Hemphillian mammals from near Juchitlán
listed in a series of articles by Carranza-Castañeda and Miller (1996,
1998, 2000, 2002, 2004; Miller and Carranza-Castañeda, 1998, 2001)
are discussed below under Tecolotlán.
Rhinoceros specimens from Juchitlán in the INAH-MRG and
MPG collections mentioned originally by Brunet (1969) are documented
as Teleoceras guymonense by Lucas (2008a) in this volume. Specimens
of Cuvieronius hyodon from Juchitlán (Fig. 5; Table 1) are: INAH-MRG
uncatalogued lower jaw with left and right m2-3, m3s partially erupted
(Fig. 5E); INAH-MRG 10-295048, right dentary with m3 (Fig. 5D);
MPG 3, right dentary fragment with m2; MPG 10, left dentary fragment
with incomplete m3; MPG 709, left dentary fragment with m3; MPG
714, left dentary fragment with very worn m3; MPG 715, right m3;
MPG 721, right dentary fragment with incomplete m3; MPG 722, left
m3; MPG 723, right dentary fragment with m3; MPG 724, left dentary
fragment with m2-3; MPG uncatalogued, right dentary fragment with
m3.
In the MPG collection there are various teeth and an incomplete
foot of the Hemphillian horse Neohipparion eurystyle from Juchitlán:
MPG 167, right dentary fragment with m1-3 (Fig. 6E); MPG 168, middle
digit; MPG 256-259, right P4, M1, M3, left M2 (Fig. 6A-D); MPG 863,
lower jaw with left p2-m3 and right p3-m3 (Fig. 6F). The teeth show
diagnostic features of N. eurystyle as defined by MacFadden (1984)
including: isolated and elongate protocones with angular anterior and
posterior borders, prominent parastyles and mesostyles, pli caballin on
some upper cheek teeth with multiple loops (Fig. 6A, C), well developed
pli caballinids, very shallow ectoflexids, prominent isthmuses, and angu-
lar enamel borders on expanded stylids and conids. Size (e.g., M1 length
= 20.4 mm, width =19.5 mm) is also within the range of measurements of
N. eurystyle. Other measurements include: MPG 258, right P4 l = 22.3, w
= 17.7; 257, left M2, l = 21.4, w =21.6; 256, right M3, l = 19.4, w =15.4;
167, right dentary fragment with m1-3, m1 l = 18.8, w = 13.6, m2 l =
FIGURE 2. Fossils of Cuvieronius hyodon from Ameca in the INAH-MRG collection (uncatalogued). A-B, Lower jaw with left and right m3s. C, Occlusal
view of left m3 in lower jaw in A-B. D-E, Hyoid in two views. Scale bars = 2 cm.
TABLE 1. Measurements (in mm) of cheek teeth of selected specimens of
Cuvieronius hyodon from Jalisco in the INAH-MRG and MPG collections.

54
19.8, w = 12.7, m3 w= 10.4; and 863, lower jaw with left p2-m3 and right
p3-m3, p2 l = 20.0, w = 12.3, p3 l = 17.2, w = 13.5, p4 l = 16.9, w = 13.2,
m1 l = 17.3, 2 = 12.1, m2 l = 19.8, w = 12.1, m3 l = 23.2, w = 11.2.
The MPG collection also has specimens of Mammuthus from
Juchitlán: MPG VF-001, juvenile molar; MPG VF-017, molar fragment;
MPG 295, maxillary with both M3s.
Clearly, the MPG Mammuthus cannot have come from the same
stratigraphic level as the Teleoceras or the Neohipparion. But, the asso-
ciation of Cuvieronius and Mammuthus is possible. Thus, specimens at
the INAH-MRG and MPG from the Juchitlán area are of Hemphillian
and Irvingtonian/Rancholabrean ages.
Los Corrales
Carranza-Castañeda (2006) mentions a locality called Los Corrales
(at 21o57.56’N, 103o18.27’W) that yielded molars of Dinohippus
interpolatus and Callippus cf. C. castilli. He assigned these fossils a late
Hemphillian age, but they have not been documented.
Mezcala
Rodriguez de la Rosa et al. (2004) reported human footprints of
?Pleistocene age from tuffaceous sediments at three sites (El Pando del
Venado, La Cuchilla and La Coatera) near Mezcala. However, no docu-
mentation of the footprints or of their age was presented. Lockley et al.
(2007, p. 83) listed these records as Holocene and well observed that
“various [human footprint] sites are reported from Mexico…though
documentation is sparse and few have been dated reliably.” I regard the
Mezcala footprints as undocumented and not part of the Pleistocene
vertebrate fossil record of Jalisco.
Palo Dulce (Atotonilco el Alto)
Solorzano (2002) reports Mammuthus sp. from this locality. How-
ever, I cannot locate publications or specimens that voucher this report.
Río de Juchipila
Solorzano (2002) reports Proboscidea from this locality. How-
ever, I cannot locate publications or specimens that voucher this report,
so I consider it to indicate the likely presence of Mammuthus sp.
San Gabriel (Venustiano Carranza)
In the INAH-MRG collection, there is an uncatalogued large sloth
skull from San Gabriel assignable to Eremotherium laurillardi (Fig. 7).
This skull has four left molariform teeth (here nominally identified as
M1-M5) and and two right molariform teeth (M4-5). The M1 is triangu-
lar, but the M2 is molariform, and the M5 is a small, round peg-shaped
tooth. The rostrum is long and tubular, and the palate is narrow. The
triangular premaxillae are loosely sutured to the maxillae, and the orbit
and zygomatic arch are relatively ventral, justifying assignment to E.
laurillardi (cf. Cartelle and DeIuliis, 1995). Indeed, the most recent revi-
sion of the genus recognizes only one polymorphic Late Pleistocene
species, E. laurillardi (Cartelle and De Iuliis, 1995). Selected measure-
ments are: skull length = 590+, width across zygomata ~ 380, width
between the orbits = 205, M1 l = 38, w = 43, M2 l = 45, w = 49, M3 l =
42, w = 48, M4 l = 41, w = 44, M5 l = 18, w = 26.
FIGURE 3. Two reconstructed mammoth skeletons (Mammuthus columbi) from Atotonilco el Bajo on display in the library of the Universidad Autonoma
de Guadalajara.

55
San Juan de los Lagos
Dugès (1897, pl. 22) documented footprints of a large felid and of
a bird found on the Rancho de Verdolaga near San Juan de los Lagos.
These tracks were found ~ 1.4 m below the ground surface on top of an
indurated marl below a tuff bed. Dugès suggested that the tracks are of
Pliocene age, but they are more likely Pleistocene. Indeed, he also re-
ported a horn core of Bison latifrons and other bones of Bison (or Bos)
associated with the tracks, which indicate a late Pleistocene age. Based
on Dugès (1897), Silva-Barcenas (1969) listed Felis concolor and Bison
latifrons from San Juan de los Lagos, and Solorzano (2002) reported
Puma concolor, Bison latifrons and Bison.
The track slabs are still present in the INAH-MRG collection
(Fig. 8). Preserved in convex hyporelief., the two cat tracks are 103 x 117
(width x length) and 84 x 113, respectively. They have four oval pads
that form a semicircle in front of the metacarpal/metatarsal pad, and no
claw tip imprints. They are readily assigned to the ichnogenus Felipeda
(cf. Vyalov, 1966; Sarjeant et al., 2002; Lucas and Schultz, 2007).
The bird footprints are tridactyl tracks in which digit III is 20-23
mm long, digit III is ~35% longer than the lateral digits and total digit
divarication is ~ 100-120o. These tracks are readily assigned to the
ichnogenus Avipeda (cf. Vyalov, 1966; Sarjeant and Reynolds, 2001;
Lucas et al., 2007).
San Martin Hidalgo (El Tepehuaje)
Solorzano (2002) reports Mammuthus columbi, M. imperator and
Mammut americanum from San Martin Hidalgo (El Tepehuaje). How-
ever, I cannot locate publications or specimens that voucher these re-
ports, so I consider them to indicate the possible presence of Mammuthus
and Mammut.
Santa Catarina
The mounted mammoth skeleton (Fig. 9) on display in the INAH
Museum in Guadalajara is the “mammut de Catarina” of Larios-Ocampo
(1963). Discovered and collected in 1962, Larios-Ocampo (1963) briefly
described the discovery, excavation and exhibition mounting of the skel-
eton. The specimen was collected on the valley floor at the ranch “El
Gancho” at UTM zone 13Q, 643951E, 2247227N (NAD 27). The bones
were no more than 1 m deep in the playa clays from which they were
excavated. Silva-Barcenas (1969) referred this skeleton from “Santa
Catarina (Municipio de Zacoalco)” to Mammuthus columbi.
This mammoth skeleton is essentially complete (Fig. 9). Dental
metrics (Table 2) support assignment to Mammuthus imperator (cf.
Madden, 1981; Agenbroad, 1994). Selected other measurements (in mm)
are: tusk length = 2710, maximum diameter = 61; humerus length = 1150,
proximal width = 330, distal width = 320; radius length = 880, proximal
width = 170, distal width = 180; ulna length = 980, proximal width =
320, distal width = 200; femur length = 1250, proximal width = 360,
distal width = 250; tibia length = 770, proximal width = 270, distal width
= 220; fibula length = 720, proximal width = 115, distal width = 150;
Tamazula
Solorzano (2002) reports Mammuthus columbi and M. imperator
from Tamazula. However, I cannot locate publications or specimens that
voucher these reports, so I consider them to indicate the likely presence
of Mammuthus sp.
Tapalpa
Solorzano (2002) reports Mammuthus columbi and M. imperator
from Tapalpa. However, I cannot locate publications or specimens that
voucher these reports, so I consider them to indicate the likely presence
of Mammuthus sp.
Tecolotlán
Various articles by Carranza-Castañeda and Miller (1996, 1998,
2000, 2002, 2004; Miller and Carranza-Castañeda, 1998, 2001; Carranza-
Castañeda, 2006) list mammals of Hemphillian, Blancan and “Pleis-
tocene” age from the vicinity of Tecolotlán and Colotlan, all of which
await fuller documentation. These mammals come from two distinct
collecting areas in un-named strata in what is referred to as the “Tecolotlán
basin.” The most recent faunal list, by Carranza-Castañeda (2006, table
1), identifies four mammal assemblages: (1) Hemphillian—Megalonyx
sp., Notolagus velox, Canis ferox, Osteoborus cyonoides, Machairodus
cf. M. coloradensis, Agriotherium schneideri, Rhynchotherium cf. R.
falconeri, Rhynchotherium sp., Gomphotheriidae indet., Teleoceras
fossiger, Callippus castilli, Neohipparion eurystyle, Nannippus minor,
Astrohippus stockii, Dinohippus interpolatus, D. mexicanus, Megatylopus
matthewi, Hemiauchenia sp., Alforjas sp., Camelidae indet. and
Hexabelomeryx fricki; (2) Hemphillian?—Copemys nr. C. valensis,
Hypolagus sp., Calomys sp., Neotoma sp. and “Desmathyus”
brachiodontus; (3) Blancan—Glyptotherium sp., Plaina sp., Neochoerus
cordobai, Platygonus sp., Equus simplicidens and Hemiauchenia sp.;
and (4) Blancan?—Hypolagus mexicanus, Felis? lacustris, Nannippus
peninsulatus and Cuvieronius sp. None of these records have been docu-
FIGURE 4. Osborn’s (1922) designated neotype tooth of the mammoth
Archidiskodon imperator from Guadalajara (after Osborn, 1922, fig. 5).

56
FIGURE 5. Selected specimens of Cuvieronius hyodon from Juchitlán and Zacoalco. A, INAH-MRG 10-295050, reconstructed upper tusk from Zacoalco.
B, MPG 731, right m2-3 from Zacoalco. C, MPG 718, left dentary fragment with m1-2 from Zacoalco. D, INAH-MRG 10-295048, right m3 from
Juchitlán. E, INAH-MRG uncatalogued, lower jaw with left and right m2-3s (m3s erupting) from Juchitlán. Scale bars = 2 cm.
mented, and the faunal list of mammals from the Tecolotlán basin changes
from article to article. The Hemphillian assemblage is associated with an
40Ar/39Ar age of 4.89 ± 0.16 Ma (Kowallis et al., 1998). Steadman and
Carranza-Castañeda (2006) described the grebe Aechmophorus cf. A.
elasson from the Tecolotlán basin.
Teocaltiche
Montellano-Ballesteros (1997) documented an assemblage of late
Hemphillian mammals from near Teocaltiche that includes “Osteoborus”
cyonoides, Astrohippus? stockii, Dinohippus? mexicanus, cf.
Hexabelomeryx fricki and an unidentified gomphothere. Carranza-
Castañeda (2006) also reported Callippus and a bird from this locality.
Teocuitatlán
Solorzano (2002) reports Mammuthus columbi and M. imperator
from Teocuitatlán. However, I cannot locate publications or specimens
that voucher these reports, so I consider them to indicate the likely

57
FIGURE 6. Specimens of Neohipparion eurystyle from Juchitlán. A, MPG 759, right M2. B, MPG 257, left M2. C, MPG 258, right P4. D, MPG 256, right
M3. E, MPG 167, right dentary fragment with m1-3. F, MPG 863, left dentary fragment with p2-m3. Scale bars = 1 cm.
presence of Mammuthus sp.
Zacoalco
Silva-Barcenas (1969) listed “Elephas sp.” from Lago de Zacoalco,
which almost certainly refers to a mammoth. Solorzano (2002) lists the
following taxa (most undocumented) from Zacoalco: the bony fishes
Oncorhynchus sp., Yuriria sp., Ictalurus spodius and I. dugesi (see Alvarez,
1966; Smith, 1987); the frog Rana sp.; a snake, Colubridae indet.; the
birds Phalacrocorax sp., Pelecanus erythrorhynchos, Phoenicopterus
sp., Ardea herodias, Mycteria sp., Aythya affinis, Chen hyperborea,
Buteogallus fragilis, Anas acuta and Mycteria wetmorei ; and the mam-
mals Didelphis marsupialis, Glyptotherium mexicanum, Eremotherium
laurillardi, Glossotherium harlani, Holmesina septentrionalis, Sigmodon
hispidus, Cynomys ludovicianus, Marmosa mexicana, Neochoerus sp.,
Procyon lotor, Arctodus pristinus, Smilodon fatalis, Canis lupus, Canis
latrans, Mammuthus columbi, M. imperator, Mammut americanum, Equus
conversidens, E. mexicanus, E. occidentalis, Equus sp., Platygonus ticuli,
Camelops hesternus, Lama sp., Odocoileus hemionus, O. virginianus,
Cervus canadensis, Tetrameryx sp., Bison latifrons and B. antiquus.
Collections in the INAH-MRG and the MPG document some of
these taxa. The pampathere Holmesina septentrionalis is represented by
a right dentary fragment with five molariform cheek teeth (INAH-MRG
10-294943: Fig. 10A-B). The figure eight cheek tooth cross sections,
size and shallow horizontal ramus justify assignment to H. septentrionalis
(e.g., Edmund, 1985; Hulbert and Morgan, 1993). Measurements are
(the first molariform is called p3): p3 l = 19.8, w = 10.4, p4 l = 21.3, w
= 10.6, m1 l = 20.9, w = 9.2, m2 l = 20.1, w = 8.3, m3 l = 16.9, w = 7.3.
Uncatalogued dermal ossicles in the INAH-MRG collection (Fig.
10C-F) can be assigned to the glyptodont Glyptotherium floridanum.
The dermal ossicles have a central figure surrounded by eight slightly

58
FIGURE 7. INAH uncatalogued skull of Eremotherium laurillardi from San Gabriel in dorsal (A), lateral (B) and ventral (C) views. Scale bar = 3 cm.
smaller figures. The Zacoalco glyptodont armor matches armor of
Glyptotherium floridanum (Gillette and Ray, 1981, figs. 88-91).
A skull and lower jaw of the sloth Megalonyx jeffersonii from
Zacoalco in the INAH-MRG collection (Fig. 11) is edentulous except for
the small caniniforms. It shows clear diagnostic features of M. jeffersoni
(G. McDonald, written commun., 2008; also see Stock, 1925). Selected
measurements are: length occiput to tip of nasals = 290, width occiput =
140, maximum width between orbits = 120, length of lower jaw = 225
and height of coronoid process = 115.
The capybara Neochoerus aesopi is represented in the INAH-
MRG collection by an uncatalogued skull with teeth (Fig. 12), an eden-
tulous skull and various postcrania. In the skull illustrated here, M3 has
16 prisms and is within the size range (M3 l = 51.6, w = 16.7) of N.
aesopi (Mones, 1991, pl. 5). Indeed, the capybara material from Zacoalco
described by Alvarez (1971) was also referred to N. aesopi by Mones
(1991).
Two sizes of Canis are present in the INAH-MRG collection
from Zacoalco. The smaller dog, the coyote C. latrans, is represented by
a left dentary fragment with p4 (Fig. 13A-B). The p4 l = 11.6, w = 5.2.
The relatively small size, very elongate lower jaw and p4 with second
and third cusps and a postero-medial cingulid justify identification as C.
latrans (Nowak, 1979).
Three dentary fragments with teeth (Fig., 13C-E) belong to the
larger dog, here identified as the wolf, Canis lupus. Size, p4 lacking a
third cusp and the relatively broad m1 with a small talonid support
assignment to C. lupus (cf. Nowak, 1979). Measurements are: INAH-
MRG 10-294917 (Fig. 13D) p4l = 15.1, w = 5.5, m1l = 27.2, w = 11.1,
m2l = 10.2, w = 8.4; 10-294918 (Fig. 13C) p3l = 14,5, w = 5.2, p4l =
16.6, w = 6.8, m1l = 27.9, w = 10.5; 10-294919, p2l = 12.7, w = 5.8, p3l
= 14.8, w = 5.9, p4l = 16.4, w = 7.8, m1l = 29.5, w = 10.9, m2l = 12.9,
w = 9.2.
Two fossil cat specimens from Zacoalco in the INAH-MRG col-
lection can be assigned to Smilodon fatalis (cf. Matthew, 1910). They are
an upper canine (Fig. 14B) and a right dentary with p4-m1 (Fig. 14A).
The canine is very curved and has a serrated posterior edge, a minimum
length of 176 and a maximum diameter of 28.7. Lower cheek tooth mea-
surements are: p4l = 10.0, w = 3.8, m1l = 14.6, w = 5.5. The very curved
and well serrated upper canine, lack of p3 and relatively small mandibu-
lar flange are features characteristic of S. fatalis (Kurtén, 1965)
The short-faced bear Arctodus simus from Zacoalco is represented
by a palate with left and right P4-M2 in the INAH-MRG collection (Fig.
14C). The relatively large P4, broad and square M1 and relatively narrow
M2 and size justify assignment to A. simus (cf. Kurtén, 1967). The P4 l
= 19.1, w = 15.1, M1 l = 21.9, w = 19.6, M2 l = 36.9, w = 21.9.
Proboscideans from Zacoalco can be assigned to Cuvieronius (Fig.
5) and to Mammuthus (Fig. 15). Alberdi and Corona M. (2005) identified
several specimens in the MPG and INAH MRG from Zacoalco or Chapala
as Cuvieronius: MPG 712, 713 (Alberdi and Corona-M., 2005, fig. 3a),
718 (Fig. 5C), 722, 723, 724 (Alberdi and Corona-M., 2005, fig. 3b) and
INAH-MRG 10-295052 (Alberdi and Corona-M., 2005, fig. 3e) and 10-
295054. They also assigned a few specimens in the MPG and INAH-
MRG from “Chapala-Zacoalco” to Stegomastodon: MPG 4, right m2;
11-12, tusk fragments (but these bear enamel bands); and INAH-MRG
10-295046, a maxilla with left and right M2-3. However, these speci-
mens are morphologically within the range of variation of Cuvieronius
(see Lucas, 2008b). Also from Zacoalco are MPG 731, a right dentary
fragment with m2-3 (Fig. 5B) and a reconstructed tusk, INAH-MRG 16-
295050 (Fig. 5A).
Only a few mammoth specimens in the INAH-MRG and the
MPG collections can be definitely provenienced as Zacoalco; many mam-
moth specimens in those collections are simply listed as “Chapala-
Zacoalco.” Mammoths certainly from Zacoalco are INAH-MRG
uncatalogued, a right dentary with m3 (Fig. 15A) and INAH-MRG 10-
295058, a maxilla with both M3s (Fig. 15B). These have relatively thick
plates and enamel (Table 2) and thus are assignable to Mammuthus

59
FIGURE 8. Footprints of a cat (A, Felipeda) and birds (B-C, Avipeda) from near San Juan de los Lagos in the INAH-MRG collection.
imperator.
Mones (1973) named the new peccary species Platygonus ticuli
for a skull and lower jaw from Zacoalco. Dental metrics of this taxon
overlap those of P. vetus (cf. Slaughter, 1966), so I consider P. ticuli a
likely junior subjective synonym of P. vetus.
In the INAH-MRG collection there is a skull and a lower jaw of a
peccary from Zacoalco (Fig. 16) that well matches Platygonus ticuli but
that I assign to P. vetus. The skull bears left I1-2, C, P2-M3 and right C,
P2-M3 and the lower jaw has right and left i1-2, c, p2-m3. Measure-
ments are: P2 l = 9.5, w = 11.7; P3 l = 11.5, w = 12.2; P4 l = 11.6, w =
15.1; M1 l = 12.3, w = 14.6; M2 l = 17.5, w = 17.9; M3 l = 19.1, w =
18.3, p2 l = 10.5, w = 8.5, p3 l = 11.1, w = 9.1, p4 l = 11.3, w = 11.5, m1
l = 13.0, w = 12.1, m2 l = 17.1, w = 13.7, m3 l = 24.1, w = 13.9.
Two upper molars from Zacoalco in the INAH-MRG collection
can be assigned to Camelops hesternus (cf. Webb, 1965) based on their
size and morphology. INAH-MRG 10-295018 (Fig. 17A) is a right M2
(l = 46.6, w = 30.6) and 10-295020 (Fig. 17B) is a left M3 (l = 45.2, w =
24.3). In the INAH-MRG collection, an uncatalogued left dentary frag-
ment with m1-3 (Fig. 17C) from Zacoalco can be assigned to Bison but is
not sufficient material for a species-level identification.
DISCUSSION
This review of the fossil vertebrate record of the Mexican state of
Jalisco supports the following observations and conclusions:
1. Fossil vertebrate assemblages of Hemphillian (mostly late
Hemphillian) and Rancholabrean age are common in Jalisco. Less com-
mon are Blancan and Irvingtonian? assemblages, and no assemblages older
than Hemphillian are known.
2. Most of the fossil vertebrate localities in Jalisco are late Pleis-
tocene (Rancholabrean) records of mammoths. Arroyo-Cabrales et al.
(2003) listed 15 mammoth records in Jalisco, whereas I only list 13
mammoth sites. This difference is an outgrowth of my lumping of some
localities (such as Ajijic and Ameca) into a single more areally extensive
locality (such as Chapala). However, although most records of Mammuthus
from Mexico are of the Columbian mammoth M. columbi (Arroyo-Cabrales
et al., 2003), many of the mammoth records in Jalisco are of the imperial
mammoth, M. imperator.
3. Extensive late Hemphillian mammal assemblages in Jalisco, par-
ticularly from the Tecolotlán region in the western part of the state, are
some of the most extensive Hemphillian mammal assemblages in Mexico.
They await further study and published documentation.

60
FIGURE 9. The mounted skeleton of Mammuthus imperator from Santa Catarina in the INAH-MRG display facility.
TABLE 2. Measurements (in mm) of cheek teeth of selected specimens of
Mammuthus imperator from Jalisco in the INAH-MRG and MPG collections.
Abbreviations are: l = length, w = width, h = crown height, pl = plate number,
plr = plate ratio, plt = plate thickness, et = enamel thickness.
4. The Rancholabrean vertebrate assemblages from Jalisco, par-
ticularly from the Lake Chapala and Zacoalco basins, are some of the
most extensive assemblages of this age from Mexico. These assemblages,
too, merit fuller documentation. In particular, micromammals (rodents,
lagomorphs) of Rancholabrean age are known from Jalisco, but little
effort has been expended to collect and document these important ele-
ments of the Jalisco Rancholabrean fauna.
5. Claims of a Pleistocene human presence in Jalisco lack convinc-
ing documentation. Arroyo-Cabrales et al. (2006) reviewed evidence from
Jalisco of human-mammoth associations (Aveleyra-Arroyo de Anda,
1962; Solórzano, 1989) and concluded that they lack sufficiently de-
tailed contextual data to establish such an association. Human footprints
of supposed Pleistocene age have not been convincingly dated.
6. The fossil vertebrates from Jalisco indicate that clastic deposi-
tion in the basins of the Jalisco triple junction began at least during the
late Hemphillian (~ 4.9 Ma) and continued through the Rancholabrean.
The abundance of late Hemphillian and Rancholbrean assemblages may
indicate that sediments of those ages are more abundant in Jalisco and, if
so, this suggests there were relatively high sedimentation rates (pulses of
sedimentation) during the late Hemphillian and Rancholabrean, probably
due to tectoinic events.
7. Fossil mammal assemblages from Jalisco are consistent with
traditional understanding of the great American biotic interchange. Thus,
Hemphillian assemblages in Jalisco are of North American composition,
whereas Rancholabrean assemblages show a typical, post-interchange
mixture of taxa of North and South American origin.
8. Much more needs to be undertaken with regard to establishing
the stratigraphic distribution and taxonomic documentation of fossil ver-
tebrates of Jalisco. Such work will produce a much more useful fossil
vertebrate record for interpreting the climatological, tectonic and biotic
histories of the late Cenozoic basins of the Jalisco triple junction.
ACKNOWLEDGMENTS
This research was supported by the Department of Cultural Af-
fairs and by INAH, as part of a collaborative project to evaluate and
develop the vertebrate fossil collection in the INAH-MRG. Tisa Gabriel
invited me to participate in this project, and assisted my work in many
ways. I am also particularly grateful to Frida Mateos González, Director
of the INAH-MRG, for making much of my work at that museum
possible. At the MRG I am grateful to Javier Juárez Woo, Óscar Rojas
Santana and Ricardo Aguilar Alonso for their generosity and help. Federico
A. Solorzano shared his extensive knowledge of the paleontology of
Jalisco with me, for which I am deeply grateful. Yami Lucas assisted this
project in various ways. Greg McDonald, Gary Morgan and Richard
White provided helpful reviews of the manuscript.

61
FIGURE 10. Holmesina and Glyptotherium from Zacoalco. A-B, INAH-MRG 10-294943, Holmesina septentrionalis, right dentary fragment with five
molariform teeth. C-F, INAH-MRG uncatalogued, Glypotherium floridanum, dermal plates. Scale bars = 1 cm.
FIGURE 11. INAH-MRG 10-294922/294923, skull and lower jaw of
Megalonyx jeffersonii from Zacoalco in dorsal (A), ventral (B) and lateral
(C) views. Scale bar = 3 cm.
FIGURE 12. INAH-MRG uncatalogued, skull of Neochoerus aesopi from
Zacoalco in dorsal (A), lateral (B) and ventral (C) views. Scale bars = 3 cm.

62
FIGURE 13. Canis from Zacoalco. A-B, INAH-MRG 10-294914, C. latrans,
left dentary fragment with p3 in occlusal (A) and lingual (B) views. C-E, C.
lupus, INAH MRG 10-294918, occlusal view of right p3-m1, (C), 294917
occlusal view of right p4-m2 (D) and 294919, occlusal view of left p2-m2
(E). Scale bars = 1 cm.
FIGURE 14. Smilodon fatalis and Arctodus simus from Zacoalco. A, INAH-
MRG 10-294908, S. fatalis, labial view of right dentary fragment with p4-
m1. B, INAH-MRG 294906, S. fatalis, upper canine. C, INAH-MRG
uncatalogued, A. simus, occlusal view of right P4-M2. Scale bars = 1 cm.
FIGURE 15. Mammuthus imperator from Zacoalco. A, INAH-MRG
uncatalogued, right m3. B, INAH-MRG 10-295058, maxilla with both M3s.
Scale bar = 2 cm.
FIGURE 16. The peccary Platygonus vetus from Zacoalco, INAH-MRG
uncatalogued skull and lower jaw in dorsal (A), lateral (B) and ventral (C)
views. Scale bars = 3 cm.
FIGURE 17. Camelops hesternus and Bison sp. from Zacoalco. A-B, C.
hesternus, right M2 (A: INAH-MRG 10-295018) and left M3 (B: INAH-
MRG 10-295020). C, INAH-MRG uncatalogued, Bison sp., left dentary
fragment with m1-3. Scale bars = 1 cm.

63
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