Quaternary International 217 (2010) 53–104

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Quaternary International
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Pleistocene mammals of Mexico: A critical review of regional chronofaunas,

climate change response and biogeographic provinciality
Ismael Ferrusquı́a-Villafranca a, *, Joaquı́n Arroyo-Cabrales b, Enrique Martı́nez-Hernández a,
Jorge Gama-Castro a, José Ruiz-González a, Oscar J. Polaco b,1, Eileen Johnson c
a
Instituto de Geologı́a, Universidad Nacional Autónoma de México, Ciudad Universitaria, Coyoacan, México, DF
b
Laboratorio de Arqueozoologı́a, Instituto Nacional de Antropologı́a e Historia, Moneda 16, Centro Histórico, México, DF
c
Museum of Texas Tech University, Lubbock, TX, USA

a r t i c l e i n f o a b s t r a c t

Article history: Mexico’s Pleistocene terrestrial mammal record includes 13 orders, 44 families, 147 genera and 280
Available online 11 December 2009 species, thus is comparable to the Recent one, but shows greater ordinal and family diversity. Post-
Pleistocene extinction chiefly involved meso- and megabaric species. The mammal, palynologic and
paleosol records are strongly time and space biased in favor of Late Rancholabrean data from a few
morphotectonic provinces; hence, only broad climatic trends could be delineated, which approximately
coincide with those known for the Wisconsinan; they disclose by 25–12 ka, greater moisture and cooler
temperature conditions than at present, coinciding too with a larger mammalian diversity and local
faunas disharmony. Climate fluctuations impacted the fauna, causing species distribution changes and
extinctions. The Recent fauna’s complex biogeographic pattern reflects this; it includes tropical and
temperate species associations outside their respective latitudes. Combining geologic and Recent
mammal distribution data with the Pleistocene record, possible dispersal routes were detected: high and
low land, southward corridors for temperate species, and low land, northward ones for tropical species.
Finally, the existence of a single Mexican Rancholabrean Faunal Province is incompatible with mammal
record’s makeup and distribution, which calls for a multiprovince scheme to better understand Mexico’s
Pleistocene mammal biogeography and faunistics.
Ó 2010 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction Notwithstanding the above, there is a wealth of information

The continental Pleistocene of Mexico is quite extensive. It carries
important resources (e.g. minerals, subsurface water), makes up the
agricultural land-substratum, and receives most of the human-
produced environmental impact, and therefore its scientific and
economic importance is without question. However, for the most
part it remains to be geologically differentiated and characterized.
Pleistocene deposits formed in virtually the whole sedimentary
spectrum make up the bulk of this subsystem, and unconformably
mantles Cenozoic or older rock bodies, which usually are highlighted
in regional studies and maps (Ortega-Gutiérrez et al., 1992; SGM,
2007) at its expense, being minimized or not shown at all.
* Corresponding author. Tel.: þ52 55 5622 4292x197; fax: þ52 55 5622 4281.
E-mail address: ismaelfv@servidor.unam.mx (I. Ferrusquı́a-Villafranca).
1
Lamentably, while the paper was in review, one of the authors, Oscar J. Polaco,
died (October 23, 2009), may he rest in peace. We his colleagues wish to dedicate
this work to his memory.
about several different aspects of Mexico’s Pleistocene, such as
sediments (both lacustrine and fluvial. e.g. Lozano-Garcı́a and
Ortega-Guerrero, 1997; Ortega-Guerrero and Urrutia-Fucugauchi,
1997; Ortega-Ramı́rez et al., 1998; Metcalfe et al., 2000), volcanoes
(e.g. González et al., 2001), geomagnetic signals (e.g. Ortega-
Guerrero and Urrutia-Fucugauchi, 1997; Caballero-Miranda, 1997;
Ruiz-Martı́nez et al., 2000), paleosols (e.g. Gama-Castro, 1996;
Cervantes-Borja et al., 1997; Gama-Castro et al., 2004; Solleiro-
Rebolledo et al., 1995, 2003, 2006), mountain glaciers (e.g. Delgado-
Granados, 1997; Vázquez-Selem, 1997), climate/climate change
(e.g. Bradbury, 1997; Metcalfe, 1997; Ruiz-Martı́nez and Werner,
1997; Metcalfe et ,al., 2000), packrat middens (e.g. Betancourt et al.,
1990; Metcalfe et al., 2000), palynomorphs (e.g. Foreman, 1955;
Clisby and Sears, 1955; González-Quintero, 1986; Lozano-Garcı́a
and Ortega-Guerrero, 1997; Lozano-Garcı́a and Xelhuantzi-López,
1997; Canul-Montañez, 2008), diatoms (e.g. Bradbury, 1971, 1989;
Metcalfe, 1992, 1995; Metcalfe et al., 2000), metaphytes (e.g. Van
Devender and Burgess, 1985; Van Devender, 1990a,b), vertebrates
(e.g. Owen, 1869, 1870; Cope, 1884, Duges, 1896, 1897; Freudenberg,

1040-6182/$ – see front matter Ó 2010 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2009.11.036
54 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
1922; Hibbard, 1955; Downs, 1958; Guenther, 1967, 1968; Mooser,
1958; Mooser and Dalquest, 1975a; Arroyo-Cabrales and Johnson,
1998, 2008; Corona-M., 2002).
Particular data sets, however, vary enormously in quantity,
quality, chronologic positioning, geographic distribution, and
statistical significance; there are important space and time differ-
ences, as well as particular biases, which make their specific
meaning not readily transferable to other data sets (Leyden et al.,
1994; Bradbury, 1997; Cervantes-Borja et al., 1997; Metcalfe et al.,
2000), because of data inconsistency or incongruity (e.g. too few vs.
too many data for a particular space or time interval, or data from
one set contradict data of other sets, etc.). In spite of these facts,
individual data-sets or data-set groups have been widely used to
address general/nationwide questions, such as Quaternary climate
change, faunal evolution (of different taxa at different levels or for
particular time intervals, or specific regions), or biotic reconstruc-
tion, but rarely explicitly pondering in full the dependability and
limitation of the sets used, thus extending their models/conclu-
sions beyond what the data objectively support.
Nevertheless, all are welcome attempts to integrate and make
sense of very diverse, and frequently incongruent data, amenable to
different interpretations, but a systematic, all-encompassing
synthesis remains to be produced. Such a synthesis would require
a long-term, multidisciplinary team effort. The present paper is
a contribution in that direction, combining state of the art infor-
mation on Pleistocene mammals, palynofloras and paleosols, and
parsimoniously integrating them into a coherent, meaningful view
on how the Pleistocene mammals evolved, became biogeo-
graphically differentiated and responded to climate change. The
paper is concerned with these broad issues: (1) review of regional
chronofaunas; (2) climate change response; and (3) biogeographic
provinciality and other relationships.
These subjects are not new, and have been addressed previously,
with different approaches/emphasis on particular aspects. Overall
knowledge on the taxonomic composition of Mexico’s Pleistocene
mammals has steadily improved (Maldonado-Koerdell, 1948;
Alvarez, 1965; Ferrusquı́a-Villafranca, 1978; Barrios-Rivera, 1985;
Arroyo-Cabrales et al., 2002, 2006, 2007a,b). Detailed description of
local faunas is extensive, but quite uneven, so that some are well
known [San Josecito Cave, N.L. (Arroyo-Cabrales and Johnson, 1998,
2008), Tequixquiac, E. Mex. (Hibbard, 1955), Lake Chapala, Jal.
(Downs, 1958), Valsequillo, Pue. (Guenther, 1967, 1968; Guenther
and Bunde, 1973; Guenther et al., 1973; Pichardo, 1997)], while
others are poorly known; sometimes even the precise locality is
unknown. A major effort in compiling and systematizing informa-
tion on Pleistocene mammals is that of Arroyo-Cabrales et al.
(CONABIO Project G-012, Arroyo-Cabrales et al., 2002, 2007a,b).
Such a project, tailored on the database framework from the North
American FAUNMAP (FAUNMAP Working Group, 1994, 1996), has
provided data from which interested scholars have benefited
immensely. Actualistic comparisons of Pleistocene and Recent
mammal faunas concerning diversity and/or species richness
include, among others, those of Barrios-Rivera (1985), Ferrusquı́a-
Villafranca and Barrios-Rivera (1996); Arroyo-Cabrales et al. (2002)
on biogeographic aspects, Mead et al. (2006), Arroyo-Cabrales et al.
(2009); on extinction, Arroyo-Cabrales et al. (2009); on climate
change in general, Urrutia-Fucugauchi et al., (1997); on effects of
climate change on mammals, Arroyo-Cabrales et al. (2009).
Dealing with the same subjects and handling the same infor-
mation sources, it is no surprise that this work overlaps to a certain
extent with some of the studies listed. However, the claim to
originality largely lies on the approach used, whereby rigorous,
explicit space and time frameworks are consistently employed to
locate and chronologically position the local faunas as well as the
palynofloras-bearing and paleosol sites. The latter two are
additional environmental indicators, thus allowing a less subjective
assessment of mammal distribution/occurrence patterns in space
and time, an evaluation of results against ‘‘objective’’ standards,
and to clearly identify avenues of future research.
The analysis of the relevant literature discloses that at least part
of the problem in integrating data sets of different features about the
Pleistocene record (geologic, biotic, climatic etc.) stems from a lax/
loose use of the time and space frameworks, whereby some basic/
‘‘technical’’ terms do not have a precise, unique meaning accepted
for everyone, rather they are used in a particular, often not explicit
way. For instance, northern Mexico or central Mexico do not refer to
an acknowledged-bound segment of the country’s territory, so they
could be interpreted to anyone’s preference. As another example,
late Pleistocene may refer to the whole second half of the Pleistocene
Subsystem/Subperiod, or to any portion of it. In addition, frequently
used terms such as community, biome, fauna and the like are
assemblages of species ‘‘united’’ by diverse relationships (e.g.
ecological without regard to a specific place or with it, space co-
occurrence with or without ecological relations, co-occurrence in
time regardless or place or of ecological relations, or a combination
of the above). To overcome/minimize this problem (and avoid
confusion), this paper uses explicit space and time frames, and the
meaning or sense of important terms is always clarified.
2. Materials and methods
2.1. Geographic framework
Mexico’s extensive territory is geographically and geologically
very complex. To properly describe it, following a western Europe
lead, since late colonial times (Von Humboldt, 1808), it has been
subdivided into ‘‘informal, natural regions’’ or units on the basis of
diverse criteria: geomorphic attributes (largely relief, e.g. sierras,
plateaus or plains), makeup (‘‘Eje Volcánico’’), being a peninsula
(Baja California Peninsula), ‘‘continental’’ setting (Yucatan Plat-
form), complexity (‘‘Nudo Mixteco’’), or a geomorph/geographic
position combinations (e.g. Sierra Madre Oriental; the term Madre
is taken in the sense of importance, i.e., being a major feature). As
geology matured, it became known that chief geomorphic features
the world over resulted from specific events/processes, and that
there was a cause/effect relationship between geologic processes
and geomorphology. Thus it became necessary to provide
a geologic characterization of such regions/features, and eventually
the concept of Morphotectonic Province (i.e. a region of particular
geomorphic features and geologic makeup/history distinctive
enough to distinguish it from neighboring regions) was developed.
Morphotectonic provinces then become practical space frame-
works to objectively and systematically describe large territories,
including their soils, vegetation (cover), fauna, flora, and biota both
Recent or fossil. Doing so has been a common practice in geography
and geology (Murray, 1961; Arbingast et al., 1975; Cook and Bally,
1975; King, 1977; Palmer, 1982; Bally and Palmer, 1989; McKnight,
2004). Thus, it is appropriate to describe and discuss Mexico’s
Pleistocene mammal record using as space framework the mor-
photectonic provinces concept. As any geologic unit, it is as useful
as it is defined. There are several schemes proposed (Guzmán and
de Cserna, 1963; de Cserna, 1989). This paper follows Ferrusquı́a-
Villafranca (1993, 1998), where provinces are delimited and fully
characterized (Table 1). The Morphotectonic Provinces (hitherto
MP) are (Fig. 1): BCP, Baja California Peninsula; NW, Northwestern
Plains and Sierras; SMOc, Sierra Madre Occidental; CH-CO,
Chihuahua-Coahuila Plateaus and Ranges; GCP, Gulf Coastal Plain;
SMOr, Sierra Madre Oriental; CeP, Central Plateau; TMVB, Trans-
Mexican Volcanic Belt; SMS, Sierra Madre del Sur; CHI, Sierra Madre
de Chiapas; and YPL, Yucatan Platform.
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 55

Table 1
Location and basic features of the Morphotectonic Provinces of Mexico. From Ferrusquı́a-Villafranca (1993, 1998); other sources, see footnote.

Provincea Location Surface in km2 and Altitude ranges Climated Chief

percentageb (m)c land
form
1 Northwestern Mexico; 109 300 -117 000 144,000; w7.34% 0–2130; 0–1000 BWh, BShs, Csa Sierras
WL; 23 000 -32 300 NL and
plains
2 Northwestern Mexico; 107 000 -116 000 236,800; w12.02% 0–2200; 200–1000 BWh, BSh Sierras
WL; 23 000 -32 300 NL and
plains
3 Western and northwestern Mexico; 289,000; w14.68% 200–3000; 2000–3000 Cfb, Aw Sierras
102 200 -109 400 WL; 20 300 -31  200 NL and
plateaus
4 Northern Mexico; 101  31 0 -110 31 255,900; w12.52% 200–2000; 800–1200 BShw, BWh, BSk Sierras
0
WL; 26 000 -31 450 NL and
plateaus
5 Northeastern and northcentral Mexico; 145,500; w7.54% 200–3000; 1000–2000 Transverse sector; BWh, BSk; Sierras
Transverse sector; 100 000 -105 000 WL; Eastern sector; Cla, Cwa, BSh
24 300 -26 000 NL; Eastern sector; 97
300 -101  200 WL; 19 400 -26 000 NL
6 Eastern Mexico; Northern sector; 96 170,600; w8.66% 0–200 Northern sector; Aw’, Cw, Plains
300 -100 200 WL; 20 000 -26 000 NL; Cx’w’; Southern sector; Afw’,
Southern sector; 91  150 -96 460 Amw’
WL; 17 100 -19 200 NL
7 Central Mexico; 100 000 -104 000 WL; 85,300; w4.33% 1000–3300; 2000–3000 BSh Plateaus
21 000 -24 000 NL
8 Central Mexico; 96 200 -105 200 WL; 175,700; w9.17% 1000–5000; 1000–2000 Aw’, Cfa, Cwa; BSh, Cw, Cflb, Peaks
17 300 -20 250 NL; Main sector; Aw and
19 000 -21 000 N L plateaus
9 Southern Mexico; 94 450 -104 400 195,700; w9.93% 0–3500; 1200–1800 Aw’, Aw, BShw,; Cwa, Cfa Sierras
WL; 15 400 -19 400 NL and
depressions
10 Southeastern Mexico; 90 300 -95 000 105,400; w5.35% 0–2500; 200–1000 Aw, Cw, Cf Sierras,
WL; 14 300 -17 400 NL depressions,
and plains
11 Eastern Mexico; 87 000 -91 000 167,600; w8.46% 0–200 BShw, Amw Plains and
WL; 17 500 -21  300 NL karst
topography
a
These numbers correspond to those on the map in Fig. 1. Baja California morphotectonic province (mp). 2. Northwestern Plains and Sierras mp. 3. Sierra Madre Occidental
mp. 4. Chihuahuan-Coahuilan Plateaus and Ranges mp. 5. Sierra Madre Oriental mp. 6. Gulf Coast Plain mp. 7. Central Plateau mp. 8. Trans-Mexican Volcanic Belt mp. 9. Sierra
Madre del Sur mp. 10. Sierra Madre de Chiapas mp. 11. Yucatan Platform mp.
b
Percentage ¼ ratio of mp to total surface area of Mexico.
c
First entry ¼ total range, second entry ¼ dominant range.
d
BWh, desert-like, Mat >18  C; BShs, Csa, temperate with dry winter; BSh, dry Mat >18  C; Cfb, temperate humid with no dry season; BShw, steppe-like, winter dry season,
Mat >18  C; BSk, steppe-like, Mat >18  C; Cfa, temperate, no defined dry season; Cwa, temperate with dry winter; Aw’, tropical with dry winter and rainy fall; Cw, temperate
with dry winter; Cx’w’, temperate with little rain throughout the year; Afw’, tropical rainy with no defined dry season; Cf, temperate with no defined dry season. The key to the
letter symbology is: A, warm humid and subhumid Climate Group (lack of a well-defined dry season); m, rainy season restricted to the summer; w, dry winter and warm
season from April to September; w’, less rainy summer with a short dry season. B, warm to cold and very arid to semiarid Climate Group; BS, warm to semicold and arid to
semiarid Climate Subgroup; BW, warm to semicold and very arid Climate Subgroup; h, semiwarm with cool winter; k, temperate with a warm summer; s, rainy winter. C,
temperate to semicold and humid to semihumid Climate Group; a, warm summer; b, cool and long summer; x’, rainy fall. Source: Garcı́a (1988).

(a) To assess the spatial/geographic distribution of the chief
Pleistocene mammal localities and environmental indicator
sites (palynofloras and paleosols), they were plotted on a mor-
photectonic provinces map template. Localities were further
time-discriminated (see Section 2.2). Mammals, palynofloras
and paleosols permit, in principle, a broad discrimination of
past temperatures, humidity and vegetation types; their
accuracy/sensitivity is a function of the quantity and quality of
the record, and it also may be improved as high sensitivity
indicators become available (such as present-day
counterparts).
(b) To assess the environmental significance of the recorded
Pleistocene local mammal faunas, palynofloras and paleosol
sites, their localities were also plotted on a present-day natural
vegetation map template (Rzedowski and Reyna-Trujillo, 1990),
thus allowing detection of congruency/incongruity in the
comparison of data.
(c) To assess the biogeographic significance of the Pleistocene
mammals, the chief localities/local faunas are plotted too on
a present-day biotic provinces map template, revealing
congruency/incongruity among them.
(d) The results of (a)–(c) were compared/contrasted with relevant
published information to validate them or to discuss alterna-
tive interpretations.
2.2. Geochronologic framework
Given that: (a) the Quaternary Period and Pleistocene Epoch
concepts have recently been the subject of a wide and heated
discussion, which also pertains the broader issue of the Paleogene–
Neogene versus the Tertiary concepts (Aubry et al., 1999; Berggren,
1998; Gradstein et al., 2004; Walsh, 2006). (b) The Pleistocene
mammal record of Mexico (and of the whole sedimentary record for
that matter) is chiefly biochronologically dated, because it has very
few radio-isotopic and paleomagnetic calibrated dates (Pichardo,
2000b; Arroyo-Cabrales et al., 2002; Mead et al., 2006). To avoid
misunderstandings, the paper follows the scheme used by Bell et al.
(2004), in which the Pliocene–Pleistocene Boundary is set at 1.8 Ma,
loosely following IGC Project 42, INQUA Subcommission 1d 1984
Report, IUGS Report 1985 (Bassett, 1985), Cande and Kent (1995),
and Berggren et al. (1995). This boundary approximately coincides
56 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Fig. 1. Morphotectonic provinces of Mexico (modified from Ferrusquı́a-Villafranca, 1993, 1998). For province characterization see Table 1.
with Subchron 2n of the Matuyama Chron of the geomagnetic
polarity chronology, as calibrated by the latter authors. The Pleis-
tocene–Holocene (or Recent) Epochal Boundary is set at 10 ka, as
a working compromise of widely diverse views, e.g. INQUA Holo-
cene Commission 1973 Decision (Fairbridge, 1974), Wright and Frey
(1965), Grayson (1989), and Morrison (1991). The Blancan, only its
latest part, Irvingtonian and Rancholabrean North American Land
Mammal Ages (NALMA) are included in the Pleistocene; all are
diachronic. Bell et al. (2004) NALMA definitions/characterizations
are consistently followed, but see Térapa l.f. below.
2.3. Data sets and procedures
The Pleistocene faunal list used (Table 2) is a modified version of
those given in Arroyo-Cabrales et al. (2002, 2007a,b), which heavily
relied on the Quaternary Mexican Mammalian Database (QMMDB),
whose creation and description is presented in Arroyo-Cabrales
et al. (2002). It was compiled largely from Alvarez (1965), Kurtén
and Anderson (1980), and Barrios-Rivera (1985), and supplemented
with many additional sources, thus expanding its primary aim
(records not older than 40 ka). The results of compilations such as
this (e.g. species count) may differ from those of others (Alvarez,
1965; Silva-Bárcenas, 1969; Barrios-Rivera, 1985), because of
various reasons such as taxonomic/nomenclatorial interpretations,
age assignments, and involuntary overlook. What is presented here
is a fair and as complete as possible representation of Mexico’s
Pleistocene mammal fauna, but certainly it can be improved.
Ceballos and Oliva (2005) provide a check list of Recent mammals
of Mexico and their distribution; and this is supplemented by data
from Hall (1981), Ramı́rez-Pulido et al. (2005), and Villa-Ramı́rez
and Cervantes (2003).
Pleistocene mammal body mass was estimated from skeletal
remains (most ‘‘micromammals’’ such as insectivorans, chiropterans
and rodents are less than 1 kg), and supplemented from Kurtén and
Anderson (1980) data. Recent mammal weight data were derived
from Villa-Ramı́rez and Cervantes (2003), and Ceballos and Oliva
(2005). The following general mammal weight categories were used:
microbaric (<9 kg), mesobaric (10–99 kg) and megabaric (>100 kg),
expanding the earlier concept (Ferrusquı́a-Villafranca et al., 2007).
The dietary habits for Recent mammal species were derived from
Villa-Ramı́rez and Cervantes (2003) and Ceballos and Oliva (2005).
Those of Pleistocene species were largely inferred from Webb (1978)
and Kurtén and Anderson (1980). Four broad categories were
recognized: omnivorous, carnivorous (including insectivorous),
herbivorous, and hematophagous. This work benefited from the
Arroyo-Cabrales et al. (2007a,b) compilation.
The possible response of mammals to the Pleistocene environ-
mental change was assessed through actualistic comparisons. The
local faunas plot on the present-day potential vegetation template
allows determination if the local faunal taxonomic makeup is
ecologically congruent with the Recent setting, assuming corre-
spondence between closely related species. An example to illus-
trate this follows: the presence of a species known to live in the
tropical forest (e.g. the didelphid Caluromys derbianus) in the
Central Plateau, where now a xerophilous scrub vegetation occurs,
would indicate that at certain times in the Pleistocene, there was
a tropical forest in the Central Plateau, thus apparently indicating
a major environmental change. To support this contention, inde-
pendent evidence must be sought, including appropriate sedi-
mentary composition of the fossil-bearing strata, co-occurrence in
the locality of ecologically equivalent taxa (mammals or others), or
of demonstrably tropical palynofloras or paleosols. Precise dating
would be a must to evaluate the timing of this change.
The same line of reasoning applies to assess the biogeographic
significance of Pleistocene mammals: the plot of local faunas on
a present-day biogeographic template may disclose congruency/
incongruity of the given fauna (or of selected species) with the
current biogeographic distribution/range now known for the fauna
or selected species therein, which was mainly derived from Hall
(1981), and Ceballos and Oliva (2005). An additional and very
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 57

Table 2 Table 2 (continued)

Pleistocene fossil mammals of Mexico, extant species also included. Chief sources:
Alvarez (1965), Barrios-Rivera, 1985, QMMDB (Arroyo-Cabrales et al., 2002), and Body Diet
Arroyo-Cabrales et al. (2007a,b); (a) extinct taxa regardless of rank are marked by size
the usual y symbol, (b) suprageneric taxa extinct in Mexico, but extant elsewhere are Carollia brevicauda S H
flagged by a diamond (r), (c) species distributed as before are marked by a small, Carollia perspicillata S H
black circle (), and (d) species extinct in a specific morphotectonic province(s) Carollia soweli S H
bearing the fossil locality(ies), but extant in other(s) are flagged by a small open Centurio senex S H
circle (B). Chiroderma villosum S H
Choeronycteris mexicana S H
Body Diet Chrotopterus auritus S C
size Dermanura phaeotis S H
DIDELPHIMORPHIA Desmodus cf. D. draculaey S B
Didelphidae Desmodus rotundus S B
Caluromys derbianus S O Desmodus stockiy S B
Didelphis marsupialis S O Diphylla ecaudata S B
Didelphis virginiana S O Enchisthenes hartii S H
Marmosa canescens S O Glossophaga soricina S H
Marmosa lorenzoiy S O Leptonycteris curasoae S H
Marmosa mexicana S O Leptonycteris nivalis S H
Philander opossum S O Macrotus californicus B S C
XENARTHRA Micronycteris microtis S C
Dasypodidae Mimon bennettii S C
Cabassous centralis B S O Sturnira lilium S H
Dasypus novemcinctus S O Tonatia evotis S C
Holmesina septentrionalisy L O Vespertilionidae
Pampatherium mexicanumy L O Corynorhinus townsendii S C
Glyptodontidaey Eptesicus brasiliensis S C
Glyptotherium cylindricumy L H Eptesicus furinalis S C
Glyptotherium floridanumy L H Eptesicus fuscus S C
Glyptotherium mexicanumy L H Lasionycteris noctivagans S C
Megalonychidaey Lasiurus cinereus S C
Megalonyx jeffersoniy L H Lasiurus ega S C
Megalonyx wheatleyiy L H Lasiurus intermedius S C
Megatheriidaey Myotis californicus S C
Eremotherium laurillardiy L H Myotis keaysi S C
Nothrotheriops mexicanumy L H Myotis thysanodes S C
Nothrotheriops shastensisy L H PRIMATES
Mylodontidaey Cebidae
Glossotherium harlaniy L H Alouatta palliata B S O
Paramylodon harlaniy L H Alouatta pigra S O
Myrmecophagidae Ateles geoffroyi B S O
Myrmecophaga tridactyla B M C CARNIVORA
Tamandua mexicana S C Canidae
INSECTIVORA s.l. Canis cedazoensisy M C
Soricidae Canis dirusy M C
Cryptotis mayensis S C Canis edwardiiy M C
Cryptotis mexicana S C Canis familiaris M C
Cryptotis parva S C Canis latrans M C
Notiosorex crawfordi S C Canis lupus M C
Sorex milleri S C Canis rufus  M C
Sorex oreopolus S C Cuon alpinus  M C
Sorex saussurei S C Urocyon cinereoargenteus S C
CHIROPTERA Felidae
Antrozoidae Herpailurus yagouaroundi S C
Antrozous pallidus B S C Leopardus pardalis M C
Emballonuridae Leopardus wiedii S C
Balantiopteryx io B S C Lynx rufus S C
Peropteryx macrotis S C Panthera atroxy L C
Saccopteryx bilineata S C Panthera onca L C
Molossidae Puma concolor M C
Eumops bonariensis S C Smilodon fatalisy L C
Eumops perotis B S C Smilodon gracilisy L C
Eumops underwoodi B S C Hyaenidae r
Molossus rufus S C Chasmaporthetes johnstoniy M C
Nyctinomops aurispinosus S C Mustelidae
Nyctinomops laticaudatus S C Conepatus leuconotus S O
Promops centralis S C Conepatus mesoleucus S O
Tadarida brasiliensis S C Lontra longicaudis S C
Mormoopidae Mephitis macroura S O
Mormoops megalophylla S C Mephitis mephitis S O
Pteronotus davyi S C Mustela frenata S C
Pteronotus parmellii S C Mustela nigripes B S C
Natalidae Spilogale putorius B S O
Natalus stramineus S C Taxidea taxus S C
Phyllostomidae Procyonidae
Artibeus jamaicensis S H Bassariscus astutus S O
Artibeus lituratus S H Bassariscus ticuliy S O
(continued on next page)
58 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Table 2 (continued) Table 2 (continued)

Body Diet Body Diet

size size
Nasua narica M O Onychomys leucogaster S H
Potos flavus S H Oryzomys alfaroi S H
Procyon lotor M O Oryzomys couesi S H
Procyon pygmaeus M O Oryzomys melanotis S H
Ursidae Otonyctomys hatti S H
Arctodus pristinusy L C Ototylomys phyllotis S H
Arctodus simusy L C Peromyscus boylii S H
Tremarctos floridanusy L O Peromyscus difı́cilis S H
Ursus americanus B L O Peromyscus eremicus S H
RODENTIA Peromyscus leucopus S H
Castoridae Peromyscus levipes S H
Castor cf. C. californicus M H Peromyscus maldonadoiy S H
Cuniculidae Peromyscus maniculatus S H
Cuniculus paca M H Peromyscus melanophrys S H
Dasyproctidae Peromyscus melanotis B S H
Dasyprocta mexicana M H Peromyscus mexicanus S H
Dasyprocta punctata M H Peromyscus ochraventer S H
Erethizontidae Peromyscus pectoralis S H
Erethizon dorsatum B M H Peromyscus truei B S H
Coendou mexicanus M H Peromyscus yucatanicus S H
Geomyidae Reithrodontomys fulvescens S H
Cratogeomys bensoniy S H Reithrodontomys megalotis S H
Cratogeomys castanops S H Reithrodontomys mexicanus S H
Cratogeomys gymnurus S H Reithrodontomys montanus S H
Cratogeomys merriami S H Sigmodon curtisiy S H
Cratogeomys tylorrhinus S H Sigmodon hispidus S H
Orthogeomys grandis S H Synaptomys cooperi B S H
Orthogeomys hispidus S H Tylomys nudicaudus S H
Orthogeomys onerosusy S H Sciuridae
Thomomys bottae S H Ammospermophilus interpres S H
Thomomys umbrinus S H Cynomys ludovicianus B S H
Heteromyidae Glaucomys volans S H
Chaetodipus hispidus S H Marmota flaviventris B S H
Chaetodipus huastecensisy S H Sciurus alleni S H
Chaetodipus nelsoni S H Sciurus aureogaster S H
Chaetodipus penicillatus B S H Sciurus deppei S H
Dipodomys nelsoni S H Sciurus nayaritensis S H
Dipodomys phillipsii S H Sciurus variegatoides B S H
Dipodomys spectabilis S H Sciurus yuctanensis S H
Heteromys desmarestianus S H Spermophilus mexicanus S H
Heteromys gaumeri S H Spermophilus spilosoma S H
Liomys irroratus S H Spermophilus variegatus S H
Perognathus flavus S H LAGOMORPHA
Hydrochaeridae r Leporidae
Neochoerus aesopiy M H Aluralagus sp.y S H
Muridae Aztlanolagus agilisy S H
Baiomys intermediusy S H Lepus alleni S H
Baiomys musculus S H Lepus californicus S H
Baiomys taylori S H Lepus callotis S H
Hodomys alleni B S H Romerolagus diazii S H
Hodomys sp. novy S H Sylvilagus audubonii S H
Microtus californicus B S H Sylvilagus bachmani S H
Microtus guatemalensis S H Sylvilagus brasiliensis S H
Microtus meadensisy S H Sylvilagus cunicularius S H
Microtus mexicanus S H Sylvilagus floridanus S H
Microtus oaxacensis S H Sylvilagus hibbardiy S H
Microtus pennsylvanicus B S H Sylvilagus leonensisy S H
Microtus quasiater S H PERISSODACTYLA
Microtus umbrosus S H Equidae r
Neotoma albigula B S H Equus conversidensy L H
Neotoma angustapalata S H Equus mexicanusy L H
Neotoma anomalay S H Equus pacificusy L H
Neotoma cinerea  S H Equus parastylidensy L H
Neotoma floridana  S H Equus tauy L H
Neotoma lepida S H Tapiridae
Neotoma magnodontay S H Tapirus bairdii B L H
Neotoma mexicana S H Tapirus haysiiy L H
Neotoma micropus S H ARTIODACTYLA
Neotoma phenax B S H Antilocapridae
Neotoma tlapacoyanay S H Antilocapra americana L H
Neotomodon alstoni S H Capromeryx mexicanay L H
Nyctomys sumichrasti S H Capromeryx minory L H
Oligoryzomys fulvescens S H Stockoceros conklingiy L H
Ondatra nebracensis y S H Tetrameryx mooseriy L H
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 59

Table 2 (continued) systematic arrangement is largely that of McKenna and Bell (1997),
Body Diet with some modifications (cf. Wilson and Reeder, 2005; Ceballos
size and Oliva, 2005). Xenarthra was kept at ordinal level, and the work
Tetrameryx shuleriy L H follows Bell et al. (2004) in their preference for Muridae in
Tetrameryx tacubayensisy L H including the ‘‘Cricetidae,’’ because it is extensively used in pale-
Bovidae ontology. However, no judgement on the merits of this scheme
Bison alaskensisy L H
with respect to others (e.g. Carlton and Muser, 2005) is intended.
Bison antiquusy L H
Bison bison B L H
Bison latifronsy L H 2.4. Chief assumptions
Bison priscusy L H
Euceratherium collinumy L H The work takes for granted these premises:
Oreamnos harringtoniy L H
Ovis canadensis B L H
Camelidae r (a) The interpretation of the Pleistocene record follows the
Camelops hesternusy L H actualistic principle.
Camelops mexicanusy L H (b) During the Pleistocene, the geographic position and chief
Camelops minidokaey L H
geomorphic features (mountain ranges, great plains, plateaus,
Camelops traviswhiteiy L H
Eschatius conidensy L H major river systems, etc.) have not significantly changed with
Hemiauchenia blancoensisy L H respect to the present-day ones. The North American Craton
Hemiauchenia macrocephalay L H seems to have been stable, while the Baja California Peninsula/
Hemiauchenia veray L H adjacent California west of the San Andrés Fault appears at
Procamelops minimusy L H
Cervidae
present to have very slowly drifted north-northwest (Busby
Cervus elaphus  L H and Ingersoll, 1995). The rising of the major mountain ranges
Mazama americana M H (e.g. Sierra Madre Oriental, Sierra Madre Occidental) seems to
Navahoceros frickiy L H have been very small or negligible. Volcanic activity only
Odocoileus halliy L H
influenced local climate and biotic distribution during specific,
Odocoileus hemionus L H
Odocoileus virginianus L H geologically instantaneous time intervals (Tarling, 1997).
Tayassuidae (c) Because of (b), it follows that major environmental factors such
Platygonus alemaniiy M H as incoming solar energy, chief atmospheric circulation
Platygonus compressusy M H patterns, and principal oceanic current patterns were broadly
Platygonus ticuliy M H
Tayassu tajacu M H
similar to present-day ones. Further, the Mexican territory was
Tayassu pecari M H not overridden by the Laurentide Glacier, so no direct latitudinal
PROBOSCIDEA glacial advances/retreats actually occurred. Present-day glaciers
Elephantidae r are restricted to the highest picks of the Trans-Mexican Volcanic
Mammuthus columbiy L H
Belt, namely the Pico de Orizaba (¼ Citlaltepétl), Popocatépetl
Mammuthus primigeniusy L H
Gomphotheriidaey and Iztacı́huatl. They are thought to extend back in time to the
Cuvieronius tropicusy L H Pleistocene, however the paucity of research even for current
Stegomastodon mirificusy L H glaciers (Delgado-Granados, 1997), does not permit establish-
Mammutidaey ment of their altitudinal position at that time. The Popocatépetl
Mammut americanumy L H
NOTONGULATAy
glaciers are the best studied. Their lower boundary lies between
Toxodontidaey 4300 and 4760 asl, and during the period 1906–1968, they
Myxotoxodon cf. M. larensisy L H receded at the rate of 7 m/year (Delgado-Granados, 1997). It
?LITOPTERNAy would not be warranted to extrapolate this rate for the Holo-
?Macrauchenidaey
cene, and much less for the Pleistocene.
Gen. et sp. indet.y L H
(d) The taxonomic composition of the mammal fauna for the whole
Notes and abbreviations: Marine taxa excluded. References to taxa identified only at country throughout the Pleistocene, particularly at the species/
generic level are not included, ?Litopterna excepted. y Extinct taxon. r Suprageneric
taxon extinct in Mexico, but extant outside Mexico.  Species extinct in Mexico, but
genus levels still remains imperfectly known. Independent
extant outside this country. B Species extinct in the morphotectonic province(s) dating and/or calibration of faunas are very scarce. However as
bearing the fossil locality(ies), but extant elsewhere in Mexico. Body mass: L, large; a working hypothesis, it is assumed that all Recent species were
M, medium; S, small. Diet: H, herbivore; B, hematophagous; C, carnivore; O, present at least in the Late Pleistocene–Early Holocene.
omnivore. Further information in the text.

valuable source of information on both environmental change and
biogeographic evaluation/assessment is the analysis of extralimital
and disjunct geographic ranges in living species, which may
disclose species area dynamics, such as expansion/contraction,
dispersal, vicariance, refugial isolation and the like (Arroyo-Cab-
rales et al., 2007a,b, 2009). Reliable dating of pertinent geologic or
fossil records would furnish objective support to these inferences.
Problems in the identification and naming of fossil taxa are
inherent and not uncommon in vertebrate paleontologic studies,
given the nature of the fossil record, the methodology used to
describe it, and to the preferences of the scholars making the
studies. To minimize the problem, this paper follows the nomen-
clature used by Arroyo-Cabrales et al. (2002), and by those cited in
Arroyo-Cabrales et al. (2007a, p. 192), modified as needed. The
3. Results
3.1. Mexico’s Pleistocene mammal record: a review
Mexico’s Pleistocene record of terrestrial mammals consists of
12 orders (possibly 13, but see below), 43 families, 146 genera and
278 (named) species (Table 2 and Fig. 2) collected from w800
localities across the territory, mainly during the last 100 years
(Falconer, 1863; Cope, 1884; Freudenberg, 1922; Stock, 1948, 1953;
Arellano and Azcón, 1949; Hibbard, 1955; Downs, 1958; Mooser,
1958; Alvarez, 1969; Dalquest and Roth, 1970; Arroyo-Cabrales and
Johnson, 1998, 2008; Arroyo-Cabrales et al., 2004; McDonald, 2002;
Mead et al., 2006). Marine and largely coastal mammals (i.e.,
Cetacea, Sirenia, Otariidae and Phocidae) are not included, because
their record is scarce, remain little studied, and are thus marginal to
60 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
this work. The extant mammal record (Fig. 3) includes 10 orders, 35
families, 161 genera and 479 species (Ceballos and Oliva, 2005). It is
known that both records are numerically comparable, yet they
differ in structure and distribution (Ceballos et al., 2005a). This
paper addresses more fully some of these differences, and discusses
their probable cause and significance.
The available information on the Pleistocene mammal record is
quite uneven in quality, quantity, depth, and approach, all of which
bear on the results, and in the discussion/ interpretation of its
significance. The record is biased for ‘‘macro-mammals’’ (Alvarez,
1965; Ferrusquı́a-Villafranca, 1978; Barrios-Rivera, 1985), because
screenwashing techniques have been used sparingly. Many local
faunas/single occurrences come from sites that have scarce or
altogether lack adequate geologic/stratigraphic control/informa-
tion (QMMDB, Arroyo-Cabrales et al., 2002), because many collec-
tions were done not in the context of geologic/paleontologic
projects, or were accidental discoveries, thus leading to possible
mixing of chronologically/ecologically different faunal elements.
Less than 22 of the w780 known Pleistocene mammal localities
have geochronometrical dating, some 9 by 14C showing significant
minimal/maximal age differences (Pichardo, 2000b; Arroyo-Cab-
rales et al., 2002, 2007a), the remainder by other methods
including obsidian hydration, K–Ar or Ar–Ar (Ferrusquı́a-Villa-
franca, 1996 and unpublished; Arroyo-Cabrales et al., 2002, 2009;
Mead et al., 2006). The record shows strong space and time biases,
so that ‘‘central Mexico’’ and ‘‘late Pleistocene’’ mammals far
outnumber others (Arroyo-Cabrales et al., 2002). Well-dated envi-
ronmental indicators are known from only a few Pleistocene
mammal localities. Finally, indicators such as palynofloras and
paleosols, which are discussed below, show similar space/time
biases as the mammal record (see Section 3.2).
3.1.1. Geographic distribution
The QMMDB (Arroyo-Cabrales et al., 2002) records some 780
individual Pleistocene mammal-bearing localities across the
country, whose adequate plotting would require a very large scale
map. As a first approximation to assess a possible geographic
distribution pattern for them, the numbers of known localities per
state were grouped in five categories (states with 1–5 localities,
6–15, 16–29, 30–50, and <50). The state areas were plotted on the
morphotectonic province template, thus generating an approxi-
mate state locality density-distribution pattern (Fig. 4). This
portrays the unevenness of the locality density-distribution, and
Fig. 2. Taxonomic composition of Mexico’s Pleistocene terrestrial mammal fauna.
1. Didelphimorphia (1 fam., 4 gen., and 7 sp.). 2. Xenarthra (6 fam., 11 gen., and 15 sp.).
3. Insectivora s.l. (1 fam., 3 gen., and 7 sp.). 4. Chiroptera (7 fam., 35 gen., and 50 sp.).
5. Primates (1 fam., 2 gen., and 3 sp.). 6. Carnivora (6 fam., 23 gen., and 38 sp.).
7. Rodentia (9 fam., 37 gen., and 97 sp.). 8. Lagomorpha (1 fam., 5 gen., and 13 sp.). 9.
Perissodactyla (2 fam., 2 gen., and 7 sp.). 10. Artiodactyla (5 fam., 18 gen., and 35 sp.). 11.
Proboscidea (3 fam., 4 gen., and 5 sp.). 12. Notoungulata (1 fam., 1 gen., and 1 sp.). 13.
?Litopterna (1 fam., 1 gen., and 1 sp.). Total: 13 orders, 43 families, 146 genera and 278
species. Main sources: Table 2, this paper.
Fig. 3. Taxonomic composition of Mexico’s Recent terrestrial mammal fauna. 1.
Didelphimorphia (1 fam., 4 gen., and 8 sp.). 2. Xenarthra (2 fam., 4 gen., and 4 sp.). 3.
Lipotyphla (2 fam., 6 gen., and 32 sp.). 4. Chiroptera (9 fam., 66 gen., and 137 sp.). 5.
Primates (1 fam., 2 gen., and 3 sp.). 6. Carnivora (6 fam., 22 gen., and 34 sp.). 7. Rodentia
(8 fam., 46 gen., and 235 sp.). 8. Lagomorpha (1 fam., 3 gen., and 15 sp.). 9. Peri-
ssodactyla (1 fam., 1 gen., and 1 sp.). 10. Artiodactyla (4 fam., 7 gen., and 10 sp.). Total:
10 orders, 34 families, 161 genera and 479 species. Main source: Ceballos and Oliva,
2005.
singles out the TMVB as the province with the greatest density
(w45% of all localities), distantly followed by the CeP and the NW
(w10% each). It is also apparent, even within a single MP, that the
locality density may vary significantly, because state boundaries are
artificial and may cross MP boundaries, e.g. the western part of the
TMVB has the highest density, because it includes the States of
Mexico and Puebla, where most of the vertebrate collecting effort
has occurred. A portion of Puebla lies in the SMS. However, Fig. 4
better portrays the Pleistocene mammal locality space pattern than
would a mere plot of individual localities on a geopolitical map.
The primary localities were assigned to NALM Ages, and plotted
on the MP template, with the Nearctic Region-Neotropical Realm
added (Fig. 5), and the most significant were identified by name. A
list of these primary localities is provided elsewhere (Table 3). The
resulting pattern resembles the previous one, confirming the
locality distribution unevenness, whereby the TMVB bears six of
the best known localities (and local faunas) namely Tequixquiac
and Tlapacoya, Mex. (Hibbard, 1955; Alvarez, 1969), Valsequillo,
Pue. (Pichardo, 1997), La Goleta, Mich. (Arellano and Azcón, 1949;
Repenning, 1962; Miller and Carranza-Castañeda, 1984), Cuitzeo,
Mich. (Garcı́a-Zepeda, et al., 2008) and Chapala, Jal. (Downs, 1958);
three lie in the SMOr, El Cedral and Mina de San Antonio, SLP.
(Alvarez and Polaco, 1982), and San Josecito, N:L. (Arroyo-Cabrales
and Johnson, 1998, 2008); the NW also includes three, all in Sonora,
El Golfo, Térapa and Santa Brisca (Shaw, 1981; Shaw and McDonald,
1987; Van Devender et al., 1987; Shaw et al., 2005; Mead et al.,
2006); finally the CeP and YPL bear one each, Arroyo El Cedazo and
Loltún respectively (Mooser, 1958; Mooser and Dalquest, 1975a;
Montellano-Ballesteros, 1992; Arroyo-Cabrales and Alvarez, 2003).
It follows that the country’s territory south of the TMVB, largely
corresponding to the Neotropical Realm, contains a significantly
smaller Pleistocene mammal record than that to the north of the
TMVB, which mainly corresponds to the Nearctic Region. These
facts are important to shape and assess the actual/real environ-
mental and biogeographic significance of the record, and determine
the explicit limits on the ability to interpret or modeling the
mammals’ response to Pleistocene environmental change for the
whole epoch and country.
3.1.2. Geochronologic distribution and composition
3.1.2.1. Late Blancan (Blancan V) chronofauna. Fig. 6 shows the
overall chronologic distribution of the Pleistocene mammal record.
Its bias toward the Rancholabrean Age is quite strong, with older
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 61

Fig. 4. Relative state-density distribution of Mexico’s Pleistocene terrestrial mammal-bearing localities set on the morphotectonic provinces template pictured in Fig. 1. Main
source: QMMDB (Arroyo-Cabrales et al., 2002).

Fig. 5. Selected Pleistocene terrestrial mammal localities of Mexico set on the morphotectonic provinces template pictured in Fig. 1. Time frame adapted from Bell et al., 2004. The
geographic position of the localities is approximated. Main sources: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and Arroyo-Cabrales et al.
(2007a,b). For further information see Table 3 and Appendix.
62 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Table 3 Table 3 (continued )

Selected Pleistocene mammal bearing localities of Mexico. The locality names
correspond to the nearest town/village or topographic feature. Chief sources: No. Locality State Morphotectonic Age
Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and province
literature therein. 64 Laguna de las Cruces SLP CeP Pleistocene
65 Rancho la Verdolaga Jal Cep Pleistocene
No. Locality State Morphotectonic Age 66 Guanajuato Gto Cep Pleistocene
province 67 León Gto Cep Pleistocene
1 La Goleta Mich TMBV Late Blancan/ 68 Ameca Jal TMBV Pleistocene
Irvingtonian/ 69 Atotonilco-Zacoalco Jal TMBV Rancholabrean
Rancholabrean 70 Venustiano Carranza Jal TMBV Pleistocene
2 El Cedazo Ags CeP Late Blancan/ 71 Ario de Rayón Mich TMBV Pleistocene
Irvingtonian/ 72 Zacapú Mich TMBV Rancholabrean
Rancholabrean 73 Cuitzeo Mich TMBV Rancholabrean
3 El Golfo Son NW Irvingtonian 74 Palpan Mor TMBV Pleistocene
4 Comondú BCS BCP Rancholabrean 75 Lerma Mex TMBV Rancholabrean
5 La Brisca Son NW Rancholabrean 76 Valle de Toluca Mex TMBV Rancholabrean
6 Terapa Son NW Early 77 Amanalco de Becerra Mex TMBV Pleistocene
Rancholabrean 78 Tacubaya DF TMVB Rancholabrean
7 Cueva Jiménez Chih CH-CO Rancholabrean 79 Amajac Hgo TMBV Pleistocene
8 Cuatro Ciénegas Coah CH-CO Rancholabrean (?Middle-Late)
9 Cerro de la Silla NL SMOr Rancholabrean 80 Real del Monte Hgo TMBV Pleistocene
10 La Boca NL SMOr Rancholabrean 81 Pachuca Hgo TMBV Pleistocene
11 El Cedral SLP SMOr Rancholabrean 82 Atlihuetzia Tlax TMBV Pleistocene
12 Minas NL SMOr Rancholabrean (?Middle-Late)
13 Cueva de Bustamante NL SMOr Rancholabrean 83 Apizaco Tlax TMBV Pleistocene
14 San Josecito NL SMOr Rancholabrean 84 San Pedro Tecamachalco Pue SMS Pleistocene
15 La Presita SLP SMOr Rancholabrean 85 Tehuacan Pue SMS Rancholabrean
16 Mina San Antonio SLP SMOr Rancholabrean 86 Nanacatla Gro SMS Pleistocene
17 Laguna de la Media Luna SLP SMOr Rancholabrean (?Middle)
18 El Abra Tam SMOr Rancholabrean 87 Chichihualco Gro SMS Pleistocene
19 Mixtequilla Ver GCP Rancholabrean 88 Huajuapan de León Oax SMS Pleistocene
20 Teapa Tab GCP Rancholabrean 89 Yolomecatl Oax SMS Late
21 Arperos Gto CeP Rancholabrean Pleistocene
22 Chapala-Zacoalco Jal TMBV Rancholabrean 90 Santa Marta Ejutla Oax SMS Pleistocene
23 Arteaga Mich TMBV Rancholabrean 91 Tehuantepec Oax SMS Pleistocene
24 Moroleón Gto TMBV Rancholabrean 92 Coixtlahuaca Oax SMS Pleistocene
25 Tequixquiac Mex TMBV Rancholabrean 93 Tepelmeme Oax SMS Pleistocene
26 Tlapacoya Mex TMBV Rancholabrean 94 Nochixtlan Oax SMS Pleistocene
27 Hueyatlaco Pue TMBV Rancholabrean 95 Aguacatenango Chis CHI Pleistocene
28 Valsequillo Pue TMBV Rancholabrean 96 Ixtapa Chis CHI Irvingtonian/
29 Cueva Encantada Mor TMBV Rancholabrean Rancholabrean
30 Santa Cruz Pue SMS Rancholabrean 97 Chiapa de Corzo Chis CHI Irvingtonian/
31 Cueva de Monte Flor Oax SMS Rancholabrean Rancholabrean
32 Etla Oax SMS Rancholabrean 98 Villa Corzo Chis CHI Late
33 San Agustı́n Oax SMS Rancholabrean Pleistocene
34 Gruta de Loltún Yuc YPL Rancholabrean 99 Actun Lara Yuc YPL Rancholabrean
35 Actún Spukil Yuc YPL Rancholabrean 100 Cozumel QRoo YPL Pleistocene
36 Actún Coyoc Yuc YPL Rancholabrean
Abbreviations: BCP, Baja California Penı́nsula; NW, Northwestern Plains and Sierras;
37 Punta San José BC BCP Pleistocene
CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; SMOr, Sierra Madre Oriental;
38 Bahı́a Magdalena BCS BCP Late
CeP, Central Plateau; GCP, Gulf Coastal Plain; TMVB, Trans-Mexican Volcanic Belt;
Pleistocene
SMS, Sierra Madre del Sur; YPL, Yucatan Platform.
39 Santa Rita BCS BCP Rancholabrean
40 Santa Anita BCS BCP ?Early
Pleistocene ages barely represented. Mexico’s Blancan faunas include Las
41 El Carrizal BCS BCP Rancholabrean Tunas, B.C., BCP (Miller, 1980), La Concha, Chih., CH-CO (Lindsay,
42 Arizpe Son NW Pleistocene 1984), Rancho Ocote and other San Miguel de Allende Basin local
43 Isla Tiburón Son NW Pleistocene
faunas, Gto., CeP (Flynn et al., 2005; Carranza-Castañeda, 2006),
44 La Guitarra Son NW Pleistocene
45 El Rosario Sin NW Pleistocene Tecolotlán, Jal., Zietla and Amajac, Hgo., TMVB (Carranza-Casta-
46 San Blas Nay NW Pleistocene ñeda, 1998). Appropriate stratigraphic control, adequate paleon-
47 El Pantanal Nay NW Pleistocene tologic studies and radio-isotopic calibration reliably date them as
48 La Erupción Chih CH-CO Pleistocene
Early Blancan. The Rancho Ocote site also bears a large Hemphil-
49 Samalayuca Chih CH-CO Pleistocene
50 La Candela Coah CH-CO Pleistocene
lian fauna. In Tecolotlán, Jal., equid molars collected from gravel
51 Torreón Coah CH-CO Pleistocene and fine grained sand strata radio-isotopically dated as 2.6 Ma
52 Arroyo del Arenal Coah CH-CO Pleistocene (Carranza-Castañeda et al., 2002) allows assignment to Blancan V.
53 Arroyo Ojuelo Coah CH-CO Pleistocene Two other local faunas merit consideration (Fig. 7): La Goleta,
54 Cuevas del Padre NL CH-CO Pleistocene
Mich., TMVB (Arellano and Azcón, 1949; Repenning, 1962; Miller
55 Cuevas de las Iglesias Dgo SMOr Pleistocene
56 Tula Tams SMOr Pleistocene and Carranza-Castañeda, 1984), and El Cedazo, Ags., CeP (Mooser,
57 Mezquital Hgo SMOr Pleistocene 1958; Mooser and Dalquest, 1975a,b), both long-known. The age of
58 El Salitrillo Tams GCP Pleistocene the La Goleta l.f. has been contentious, being assigned both to the
59 El Ocotlán Camp GCP Pleistocene
Hemphillian and Blancan (Repenning, 1962; Ferrusquı́a-Villafranca,
60 Laguna del Salitrillo Zac CeP Pleistocene
61 Zacatecas Zac CeP Pleistocene
1978, Miller and Carranza-Castañeda, 1984), because taxa of
62 Brechas Coloradas SLP CeP Pleistocene Hemphillian, Blancan and Pleistocene age were recovered from the
63 Rancho Peotillos SLP CeP Pleistocene locality and the surrounding area. Additional study of the different
mammal-bearing strata has resulted in their placement in
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Fig. 6. Geochronologic distribution of Mexico’s Pleistocene mammal record. Main sour-
ces: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and
Arroyo-Cabrales et al. (2007a,b). For further information see Table 3 and Appendix.
stratigraphic order, which in turn has allowed researchers to
chronologically separate the taxa, and has identified an unconfor-
mity above which the Pleistocene mammals occur. Radioisotopic
dating at 3.6 Ma (Carranza-Castañeda, 2006, p. 57; strata data and
method not given) indicates most taxa are from the Blancan III, and
are pre-Pleistocene in age.
The El Cedazo l.f. (Table 4) also includes taxa of Blancan-Ran-
cholabrean age (Mooser, 1958; Mooser and Dalquest, 1975a,b;
Montellano-Ballesteros, 1992). The latter author recognized three
informal stratigraphic units, which bear seemingly distinctive
assemblages of mammal taxa. The lower unit was assigned to the
Irvingtonian on the basis of Aluralagus, and the resemblance of
Holmesina to comparable material from an Irvingtonian site in
Florida. Bell et al. (2004) regard this mammal assemblage as latest
Blancan. Further, they establish that the presence of Bison in the
Cedazo l.f. indicates that part of it must be Rancholabrean, and that
inadequate understanding of taxal time relationships makes
Fig. 7. Mexico’s chief Late Blancan and Irvingtonian terrestrial mammal-bearing localities set on the morphotectonic provinces template pictured in Fig. 1. Time frame adapted from
Bell et al., 2004. Sources: See text.
63
unclear the discrimination of an Irvingtonian chronofauna there.
Additional study is necessary to resolve this issue.
3.1.2.2. Irvingtonian chronofauna. Mexico’s Irvingtonian mammal
record is also very scarce, El Golfo l.f., Son., NW (Shaw, 1981; Shaw
and McDonald, 1987; Shaw et al., 2005); remains as the only fauna
confidently assigned to this age (Fig. 7 and Table 5). It is rather
diverse (28 genera, 21 families, and seven orders, but only 11
named species), including largely long lasting taxa, but its overall
co-occurrence is Irvingtonian (Bell et al., 2004). The presence of
Mammuthus imperator is particularly significant. No radio-isotopic
or paleomagnetic calibration is available for this fauna.
Mead et al. (2006) described a large mammal assemblage from
a fluvial and ‘‘marsh,’’ largely medium- to fine-grained sequence in
Térapa, Son, NW, whose stratigraphic relationship with the Toni-
babi 40Ar–39Ar 0.44  0.13 Ma dated basalt flow (Paz-Moreno et al.,
2003) brackets the sequence and assemblage ages between 0.57
and 0.31 Ma. This interval falls within the time span of the Irving-
tonian NALMA (Bell et al., 2004). However, the fauna includes Bison,
which is diagnostic for the Rancholabrean NALMA, and hence Mead
et al. (2006, 2007) assigned it to this age, and discussed the alter-
native of an earlier Bison dispersal event than that accepted by Bell
et al. (2004). They proposed that the earliest occurrence of Bison in
North America took place between 570 and 310 ka, and also
mentioned that an even earlier date was proposed by McDonald
and Morgan (2004). On the other hand, Carranza-Castañeda and
Roldán-Quintana (2007) studied the same area, reporting an
association of Equus excelsus, E. conversidens and Bison from the
same locality, indicating that the fossil-bearing strata overlie
olivine basalt flows 40Ar–39Ar dated between 1.7  0.74 and
0.61  0.08 Ma (Paz-Moreno et al., 2003). Carranza-Castañeda and
Roldán-Quintana (2007, p. 87) assigned the fauna to the Pleisto-
cene, indicating that ‘‘. Bison suggests a post-Rancholabrean age’’;
however, Bison is a Rancholabrean index genus (Lundelius et al.,
Table 4 Table 5
El Cedazo l.f., Late Blancan-Rancholabrean of Aguascalientes, CeP Morphotectonic El Golfo l.f., Irvingtonian of Sonora, NW Morphotectonic Province. Chief sources:
Province. Chief sources: Mooser (1958), Mooser and Dalquest (1975a,b), Mon- Shaw (1981), Shaw and McDonald, 1987, and QMMDB (Arroyo-Cabrales et al., 2002).
tellano-Ballesteros (1992), and QMMDB (Arroyo-Cabrales et al., 2002).
XENARTHRA
DIDELPHIMORPHIA Megalonychidaey
Didelphidae Megalonyx wheatleyiy
Didelphis virginiana Megatheriidaey
XENARTHRA Eremotherium laurillardiy
Dasypodidae Nothrotheriops shastensisy
Holmesina septentrionalisy Mylodontidaey
Glyptodontidaey Glossotherium sp.y
Glyptotherium floridanumy Myrmecophagidae
Glyptotherium mexicanumy Myrmecophaga tridactyla B
Megatheriidaey CARNIVORA
Nothrotheriops shastensisy Canidae
Mylodontidaey Canis rufus B
Glossotherium harlaniy Felidae
CARNIVORA Panthera onca
Canidae Herpestidae r
Canis cedazoensisy Chasmaporthetes johnstoniy
Canis dirusy Ursidae
Canis latrans Tremarctos floridanusy
Urocyon cinereoargenteus RODENTIA
Felidae Castoridae
Lynx rufus Castor sp.
Panthera atroxy Heteromyidae
Panthera onca Liomys sp.
Smilodon fatalisy Hydrochaeridae r
Mustelidae Neochoerus sp.y
Taxidea taxus Muridae
Ursidae Hodomys sp. B
Arctodus pristinusy Neotoma sp.y
Arctodus simusy Sigmodon curtisiy
RODENTIA LAGOMORPHA
Erethizontidae Leporidae
Erethizon dorsatum B Sylvilagus hibbardiy
Geomyidae PERISSODACTYLA
Cratogeomys castanops Equidae r
Pappogeomys sp. Equus complicatusy
Thomomys umbrinus Equus conversidensy
Muridae Tapiridae
Peromyscus sp. Tapirus sp. B
LAGOMORPHA ARTIODACTYLA
Leporidae Antilocapridae
cf. Aluralagus sp.y Stockoceros sp.y
Lepus californicus Tetrameryx sp.y
Sylvilagus audubonii Bovidae
PERISSODACTYLAa Ovis sp.
Equidae r Camelidae r
Equus calobatusy Camelops sp.y
Equus conversidensy Hemiauchenia blancoensisy
Equus excelsusy Paleolama sp.y
Equus mexicanusy Titanotylopus sp.y
Equus pacificusy Cervidae
Equus parastylidensy Odocoileus sp.y
Equus tauy PROBOSCIDEA
ARTIODACTYLA Elephantidae r
Antilocapridae Mammuthus sp.y
Capromeryx mexicanay Gomphotheriidaey
Stockoceros conklingiy Cuvieronius sp.y
Tetrameryx mooseriy
For symbols see Table 2.
Tetrameryx tacubayensisy
Bovidae
Bison alaskensisy 1987; Bell et al., 2004). Additional work on defining and timing the
Camelidae r Rancholabrean NALMA is needed to settle this issue.
Camelops hesternusy Possible Irvingtonian records are those of Cuvieronius sp. from
Camelops traviswhiteiy
Chiapas, CHI. The first is a mandible collected from fluvial, medium-
Hemiauchenia sp.y
Cervidae
to coarse-grained strata in Ixtapa, (Ferrusquı́a-Villafranca, unpub-
Navahoceros frickiy lished), which discordantly underlie the K–Ar 0.56 to 0.37 Ma dated
Odocoileus halliy Punta de Llano Formation, largely a felsic ignimbrite sheet
Tayassuidae (Ferrusquı́a-Villafranca, 1996). The other is a toothed mandible
Platygonus sp.y
fragment collected near Chiapa de Corzo, also from fluvial strata
PROBOSCIDEA
Elephantidae r underlying 0.453 Ma K–Ar dated tuff strata (Ferrusquı́a-Villafranca,
Mammuthus sp.y unpublished). Given that Cuvieronius is not specifically diagnostic
Mammutidaey of the Irvingtonian, but rather a long-lived genus (McKenna and
Mammut americanumy
Bell, 1997), additional study is needed to validate the possible
a
Quotation of the seven Equus nominal species does not mean accepting their Irvingtonian assignment considered here. Finally, whether the
taxonomic validity. For symbols see Table 2. lower unit of the El Cedazo locality, Ags., CeP includes an
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Irvingtonian fauna is open to question (Montellano-Ballesteros,
1992; Bell et al., 2004), as discussed above. Lucas (2008) mentions
two possible Blancan or Irvingtonian records (Nannipus sp. and
Stegomastodon primitivus) from Chapala, Jal., TMBV.
3.1.2.3. Rancholabrean chronofauna. The bulk of Mexico’s mammal
record is included in this chronofauna (Fig. 8), practically
embracing all orders, families and genera listed in Table 2. The
synthetic review nature of this paper only allows a brief discussion
of the criteria on which the Rancholabrean dating is/was given. The
occurrence of the most significant taxa is described in (major) local
faunas across the country.
Didelphimorphia. Didelphis, an extant and common Rancho-
labrean genus (Bell et al., 2004), first appeared in the Irvingtonian
(in Texas, Winkler and Gose, 2003; and in Florida, Martin, 1974),
and is known from the TMVB (Chapala-Zacoalco area, Jal., and
Barranca Seca, Ver. (Alvarez and Ferrusquı́a-Villafranca, 1967), and
Valsequillo, Pue. (QMMDB, Arroyo-Cabrales et al., 2002)). Remains
of Didelphis, Marmosa, and Philander; all extant genera, have been
recovered from Loltún Cave., Yuc., in the YPL (Alvarez and Arroyo-
Cabrales, 1990; Arroyo-Cabrales and Alvarez, 2003). The mammal-
bearing deposits from this cave have been radiocarbon dated
between 30 ka and 0.5 ka (Arroyo-Cabrales and Polaco, 2003).
Xenarthra. Mones (1971) partly reviewed the group, and later
McDonald (2002) comprehensively analyzed Mexico’s xenarthran
record, which is diverse (11 genera) and widespread (in seven of
the 11 MPs). It includes typical Rancholabrean taxa (Table 6). Pilosa:
Megalonyx jeffersoni [SMOr: San Josecito, N:L. (Stock, 1943)],
[TMVB: Valsequillo, Pue. (McDonald, 2002)]. Nothrotheriops shas-
tensis [PBC: Comondú, B.C.S. (McDonald, 2002)], [CeP: El Cedazo,
Ags. (Mooser and Dalquest, 1975a)], [SMOr: San Josecito, N.L. (Stock,
1943), Cueva de Bustamante, N.L. (McDonald, 2002), Cerro de la
Silla, N.L. (Furlong, 1925), Mina, N.L. (Franzen, 1993, 1994). La
Fig. 8. Mexico’s chief Rancholabrean terrestrial mammal-bearing localities set on the morphotectonic provinces template pictured in Fig. 1. Time frame adapted from Bell et al.,
2004. Main sources: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and Arroyo-Cabrales et al. (2007a,b). For further information see Table 3 and
Appendix.
65
Presita, S.L.P. (Polaco and Butron, 1997)]. TMVB: Chapala, Jal.
(Downs, 1958; Lucas, 2008), Las Ollas, Mich. (Silva-Bárcenas, 1969),
Valsequillo, Pue. (Pichardo, 1997), and Mexico Basin (Freudenberg,
1921)]. Eremotherium laurillardi [SMOr: El Cedral, S.L.P. (Polaco,
1981)], [CeP: (Polaco, 1981), Arteaga, Mich. (Polaco, 1981)], [SMS:
Etla, Chapala, Zacoalco and V. Carranza, Jal. Tab]. Paramylodon
harlani [PBC: Cabo Colnett (McDonald, 2002)], [SMOr: Cueva de
Bustamante, N.L. (McDonald, 2002), El Cedral, S.L.P. Oax. (Polaco,
1981)], [CeP: Arperos, Gto. (Duges, 1882; but see Mones, 1973a)],
[TMVB: Tequixquiac, E. Mex. (Freudenberg, 1921), Aquiahua, Tlax.
(Polaco, 1981), Cueva Encantada de Chimalacatlán (Arroyo-Cabrales
et al., 2004), Chapala, Jal. (Lucas, 2008)], and [GCP: Teapa, Tab.
(Polaco, 1981)]. The only fossil record of an anteater is Tamandua
sp., an extant taxon [TMVB: Cueva Encantada de Chimalacatlán
(Arroyo-Cabrales et al., 2004)].
Loricata. Glyptotherium floridanum [CeP: El Cedazo, Ags. (Mooser
and Dalquest, 1975a)]. [TMVB: Barranca Seca, Ver. (Dalquest, 1961)].
G. cylindricum [NW: Térapa, Son. (Mead et al., 2007)], [TMVB:
Ameca, Jal. (Brown, 1912); Valsequillo, Pue. (Pichardo, 1997)].
G. mexicanum [TMVB: Tequixquiac, E. Mex. (Cuatáparo and Ramı́rez,
1875; Hibbard, 1955)], [SMS: Santa Cruz, Pue. Gillette et al., 2000],
and numerous Glyptotherium sp. records from E. Mex., Distrito
Federal, Morelos and Jalisco (McDonald, 2002; Arroyo-Cabrales
et al., 2004; Lucas, 2008)]. Cabassous centralis and Dasypus novem-
cinctus [YPL: Loltún, Yuc. (Arroyo-Cabrales and Alvarez, 2003), an
extant taxa in Mexico]. Holmesina septentrionalis [CeP: El Cedazo,
Ags. (Mooser and Dalquest, 1975a)], and Pampatherium mexicanum
[NW: Térapa, Son. (P. mexicanum, Mead et al., 2007)], [TMVB:
Chapala, Zacoalco and Zula., Jal. (Edmund, 1985a,b, 1987; Hulbert
and Morgan, 1993; McDonald, 2002); Valsequillo, Atenzingo,
Hueyatlaco, and Tenextepe, Pue. (Edmund, 1996; McDonald, 2002);
Tequixquiac, Mex. (Hibbard, 1955), Tecamachalco, D.F. (Edmund,
1996), and an unknown site, Hidalgo (Edmund, 1996)].
66 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Table 6
Selected localities bearing Rancholabrean Xenarthra from Mexico. Sources, see text.

Morphotectonic Provinces: BCP, Baja California Penı́nsula; NW, Northwestern Plains and Sierras; CeP, Central Plateau; SMOr, Sierra Madre Oriental; TMVB, Trans-Mexican
Volcanic Belt; GCP, Gulf Coastal Plain; SMS, Sierra Madre del Sur; YPL, Yucatan Platform. Local faunas: A, Comondú, BCS; B, Terapa, Son; C, El Cedazo, Ags; D, Arperos, Gto; E, San
Josecito, NL; F, La Presita, SLP; G, El Cedral, SLP; H, Minas, NL; I, Cueva de Bustamante, NL; J, Cerro de la Silla, NL; K, Chapala-Zacoalco, Jal; L, Tequixquiac, Mex; M, Valsequillo,
Pue; N, Hueyatlaco, Pue; O, Arteaga, Mich; P, Cueva Encantada, Mor; Q, Teapa, Tab; R, Santa Cruz, Pue; S, Etla, Oax; T, Gruta de Loltún, Yuc.

Insectivora s.s. (¼ Lipotyphla; ¼ Soricomorpha, see Hutterer,
2005). Shrews are common Pleistocene fossils in North America,
but diagnostic of the Rancholabrean NALMA. Mexico’s Rancholab-
rean insectivoran record includes (largely) extant soricids. Cryptotis
mexicana [SMOr: San Josecito, N.L. (Findley, 1953; Esteva et al.,
2005)]. C. parva [CH-CO: Cueva Jiménez, Chih. (Messing, 1986)],
[SMOr: Mina de San Antonio, S.L.P. (Ferrusquı́a-Villafranca and de
Anda-Hurtado, 2008), El Abra, Tams. (Dalquest and Roth, 1970)].
Cryptotis mayensis [YPL: Loltún, Yuc. (Arroyo-Cabrales and Alvarez,
2003)]. Notiosorex crawfordi [CH-CO: Cueva Jiménez (Messing,
1986)], [SMOr: El Abra, Tams. (Dalquest and Roth, 1970); La Calera,
S.L.P., Jau-Mexı́a et al., 2000)]. Sorex cinereus, S. milleri, and
S. sausseuri [SMOr: San Josecito, N.L. (Findley, 1953; Esteva et al.,
2005)].
Chiroptera. Bats are rare in the fossil record because of their
fragility and habits. Not surprisingly, most records of Mexico’s
Rancholabrean chiropterans come from cave deposits, and most lie
in the YPL and SMOr. The record for this age is rather large (Table 7),
including 42 extant and two extinct species (i.e., one-third of those
present-day living in this country), which belong to 30 genera and
six families. Only two of the seven extant families, Noctilionidae
(one genus) and Thyropteridae (one genus), are not represented by
fossils. At present, Phyllostomidae (14 genera, 18 species) and
Vespertilionidae (six genera and 12 species) comprise most records
for the order. Table 6 summarizes the QMMDB (Arroyo-Cabrales
et al., 2002) relevant information, and the following paragraphs
comment on taxa that are biogeographically more significant.
The emballonurids Balantiopteryx io and Peropteryx macrotis are
known from single localities in the SMOr (Dalquest and Roth, 1970;
Arroyo-Cabrales and Johnson, 2008) and the YPL respectively
(Arroyo-Cabrales and Polaco, 2008; Morales-Mejı́a et al., 2009); the
former occurs now in the SMS, CH-CO and YPL. The molossids
include five genera and eight species, nearly all from localities in
the SMOr and YPL (Arroyo-Cabrales and Polaco, 2008; Morales-
Mejı́a et al., 2009). Only Eumops perotis is also known from the CH-
CO. All localities lie within the current geographic range of the
respective species (Ceballos and Oliva, 2005).
The mormoopids (two genera and three species) have been
chiefly recovered from localities in the SMOr (Arroyo-Cabrales and
Johnson, 2008) and YPL (Arroyo-Cabrales and Polaco, 2008;
Morales-Mejı́a et al., 2009); Mormoops megalophylla is also known
from the TMVB (QMMDB, Arroyo-Cabrales et al., 2002). The natalid
Natalus stramineus is present only in the YPL from Loltún Cave, Yuc.,
which lies within this species current range (Arroyo-Cabrales and
Polaco, in press; Morales-Mejı́a et al., 2009).
The phyllostomids are the most diverse group, largely coming
from single localities in the YPL (Arroyo-Cabrales and Polaco, 2008;
Morales-Mejı́a et al., 2009) and the SMOr (Arroyo-Cabrales and
Johnson, 2008). It includes the vampire bats, the extant Desmodus
rotundus, and the fossil species D. draculae and D. stocki, the latter
known from three sites in the SMOr (Arroyo-Cabrales and Ray,
1997; Arroyo-Cabrales and Johnson, 2008) and one in the TMVB
(QMMDB, Arroyo-Cabrales et al., 2002). All extant Rancholabrean
phyllostomid localities lie within the respective species’ current
geographic range. All but one vespertilionid species come from
single localities in the SMOr (Arroyo-Cabrales and Johnson, 2008)
and the YPL (Arroyo-Cabrales and Polaco, 2008; Morales-Mejı́a
et al., 2009), which also lie within the given species current range.
Antrozous pallidus is the only vespertilionid known from outside
the mentioned MP’s, in CH-CO (Messing, 1986).
Primates. The Rancholabrean record of primates of Mexico (and
in fact the whole Pleistocene) is very scarce, consisting of the extant
cebids Alouata palliata, A. pigra, and Ateles geoffroyi, as well as Homo
sapiens. The cebids are recorded from the TMVB (Metropolitan
Table 7
Selected localities bearing Rancholabrean Chiroptera from Mexico. Sources, see text.

Morphotectonic provinces: CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; SMOr, Sierra Madre Oriental; TMVB, Trans-Mexican Volcanic Belt; GCP, Gulf Coastal Plain;
SMS, Sierra Madre del Sur; YPL, Yucatan Platform. Local faunas: A, Cuatro Ciénegas, Coah; B, Cueva Jiménez, Chih; C, El Abra, Tam; D, San Josecito, NL; E, La Boca NL; F, La Presita,
SLP; G, Tlapacoya, Mex; H, Mixtequilla, Ver; I, Cueva de Monte Flor, Oax; J, Gruta de Loltún, Yuc; K, Actún Spukil, Yuc; L, Actún Coyoc, Yuc.
68 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Subway Works in downtown Mexico City, 1967; QMMDB, Arroyo-
Cabrales et al., 2002), and chiefly from some localities in Yucatan
(YPL), including the famous archeological site of Mayapan (Pollock
and Ray, 1957). Confidently dated Rancholabrean records of
H. sapiens come largely from localities in the TMVB (Peñon III, D.F.,
and Tlapacoya, E. Mex., González et al., 2003), 14C dated w12 ka, are
the best known.
Carnivora. The QMMDB (Arroyo-Cabrales et al., 2002) shows
that Mexico’s Rancholabrean carnivore record is extensive, but is
unevenly distributed among the MPs. It includes at least five
families (Canidae, Felidae, Mustelidae, Procyonidae and Ursidae),
21 genera and some 35 species (Arroyo-Cabrales et al., 2002, 2009).
The TMVB and SMOr contain most localities. Procyonids comprise
only a minor fraction of the record, which as a whole requires
a comprehensive and critical review to remove numerous strati-
graphically and taxonomically incorrect taxa. Table 8 summarizes
the QMMDB (Arroyo-Cabrales et al., 2002) relevant information,
stressing the major local faunas. The following paragraphs
comment on the stratigraphically or biogeographically more
significant taxa.
Canidae. The record includes three genera and nine species, and
Canis is the most diverse. C. cedazoensis is only known from the CeP
(El Cedazo, Ags., Mooser and Dalquest, 1975a), C. dirus, a Rancho-
labrean index fossil, is present in the BCP (one site), SMOr (chiefly
in San Josecito, N.L. and El Cedral, S.L.P, Nowak, 1979; Alvarez and
Polaco, 1981; Lorenzo and Mirambell, 1981, 1986), CeP (El Cedazo,
Ags., Mooser, 1958; Mooser and Dalquest, 1975a), TMVB (Chapala-
Zacoalco and Tequixquiac, cf, Aviña, 1969; Nowak, 1979; Berta,
1988), and YPL (Loltún, Yuc., Arroyo-Cabrales and Alvarez, 2003);
and C. edwardii is reported only from the CH-CO (Mina Erupción,
Chih., Nowak, 1979). The Rancholabrean Canis record includes four
extant species. C. familiaris (domestic dog) is recorded from the
TMVB (Chapala-Zacoalco, Jal., Aviña, 1969; QMMDB, Arroyo-Cab-
rales et al. 2002), GCP (Mixtequilla, Ver., Polaco, 1995) and YPL
(Loltún, Yuc., Arroyo-Cabrales and Alvarez, 2003). Domestic dog
remains are also known from many Early Holocene archeological
sites throughout the country. C. latrans (coyote) is the most wide-
spread canid, recorded from localities in these MPs: CH-CO, SMOr
(including San Josecito, N.L. and El Cedral, S.L.P., Alvarez and Polaco,
1981; Nowak, 1979), CeP (El Cedazo, Ags.), TMVB (among others:
Chapala-Zacoalco, Jal., Tequixquiac and Tlapacoya, Mex. and Val-
sequillo, Pue., cf, Aviña, 1969; Kurtén, 1974; Nowak, 1979; Berta,
1988; Lucas, 2008), and YPL (Loltún, Yuc., Arroyo-Cabrales and
Alvarez, 2003). C. lupus (wolf) is present in the SMOr, TMVB and
YPL. It frequently co-occurs with C. latrans, which is also recorded
from the SMOr, La Presita, S.L.P., Polaco and Butron, 1997. C. rufus is
only recorded in El Golfo, Son., NW (Shaw, 1981; C. armbrusteri is
the common Irvingtonian species, whereas C. rufus is little known
in this age), Chapala, Jal. and Tequixquiac, Mex., TMVB (QMMDB,
Arroyo-Cabrales et al., 2002; Lucas, 2008). Cuon alpinus is present
only in the SMOr (San Josecito, N.L., Kurtén and Anderson, 1980).
Urocyon cinereoargenteus is fairly widespread, occurring in the CH-
CO (Cuatro Ciénegas, Coah., Gilmore, 1947), SMOr (San Josecito, N.L
and other sites, Polaco and Butron, 1997), CeP (El Cedazo, Ags. and
another locality), and YPL (Loltún, Yuc., Arroyo-Cabrales and
Alvarez, 2003).
The record of the Felidae is more diverse (six genera and seven
species, three of which are extinct), but less ‘‘abundant’’ than that of
canids. Herpailurus yagouaroundi (¼ Puma yagouaroundi, see
McKenna and Bell, 1997; Wozencraft, 2005) is recorded in the SMOr
and YPL (single sites in each, Arroyo-Cabrales and Alvarez, 2003;
Arroyo-Cabrales and Johnson, 1998). Leopardus pardalis is only
known from the YPL. Lynx rufus, Panthera atrox, P. onca and Puma
concolor (¼ Felis concolor) are fairly widespread, mainly occurring
in the NW, CH-CO, CeP (El Cedazo, Ags., Mooser, 1958; Mooser and
Dalquest, 1975a), SMOr (including San Josecito, N.L., Stock, 1943;
Arroyo-Cabrales and Johnson, 1998; El Cedral, S.L.P., Alvarez and
Polaco, 1981) and TMVB (chiefly Chapala-Zacoalco, Jal. and
Tequixquiac, Mex., Aviña, 1969; Silva-Bárcenas, 1969; QMMDB,
Arroyo-Cabrales et al., 2002; Lucas, 2008); P. atrox is a Rancholab-
rean index, whereas P. onca and P. concolor are common Rancho-
labrean species that survived into the Holocene (Kurtén, 1976; Bell
et al., 2004). Smilodon fatalis is known from few sites in the CeP,
SMOr and TMVB (Hibbard, 1955, Arroyo-Cabrales and Johnson,
1998; Mooser, 1958; Mooser and Dalquest, 1975a), while S. gracilis
is recorded only in the TMVB (Valsequillo, Pue., Kurtén, 1967;
QMMDB, Arroyo-Cabrales et al., 2002). Both species are extinct.
The Mustelidae are as diverse as the Felidae (six genera and eight
species), but somewhat less widespread. Conepatus, a common
Rancholabrean taxon (Bell et al., 2004), includes C. mesoleucus,
known from the CH-CO (Cuatro Ciénegas, Coah., QMMDB, Arroyo-
Cabrales et al., 2002), SMOr (San Josecito, N.L.; Arroyo-Cabrales and
Johnson, 1998) and TMVB (Chapala-Zacoalco, Jal., Aviña, 1969;
QMMDB, Arroyo-Cabrales, et al., 2002), and C. leuconotus, which
occurs only in the GCP (Mixtequilla, Ver.). Lontra longicaudis,
Mephitis macroura and M. mephitis are only recorded in the TMVB
(Freudenberg, 1910; Aviña, 1969; QMMDB, Arroyo-Cabrales et al.,
2002). Mustela frenata and Spilogale putorius are present in the CH-
CO (San Josecito, N.L., Arroyo-Cabrales and Johnson, 1998; La Presita,
S.L.P., Polaco and Butron, 1997). Mustela sp. is known from SMOr (La
Presita, S.L.P., Polaco and Butron, 1997), and in the YPL (Arroyo-
Cabrales and Alvarez, 2003). Taxidea taxus is known from localities in
the CH-CO, SMOr and TMVB (Arroyo-Cabrales and Johnson, 1998).
The Procyonidae are less diverse and widespread. Bassariscus
astutus is known from the CH-CO and SMOr (including San Josecito,
N.L., Arroyo-Cabrales and Johnson, 1998), but B. ticuli, an extinct
species, is only recorded from the latter MP (La Presita, S.L.P., Polaco
and Butron, 1997). Nasua narica and P. lotor are present in the CH-
CO, SMOr (San Josecito, N.L.), TMVB and YPL (Arroyo-Cabrales and
Alvarez, 2003); P. lotor also occurs in the NW (Térapa, Son., Mead
et al., 2006).
The Ursidae (three genera and four species) are largely extinct in
Mexico. Arctodus pristinus, an uncommon Blancan and Irvingtonian
species that survived in the Early Rancholabrean (Dalquest and
Mooser, 1980; Bell et al., 2004) and A. simus from the Rancholab-
rean are known from these MPs: CeP (El Cedazo, Ags., Churcher
et al., 1996; QMMDB, Arroyo-Cabrales et al., 2002), SMOr (San
Josecito, N.L., Stock, 1950; Arroyo-Cabrales and Johnson, 1998), and
the TMVB (Chapala-Zacoalco, Jal., Tequixquiac, Mex., Churcher
et al., 1996); A. simus is also recorded in Valsequillo, Pue., TMVB
(Churcher et al., 1996; QMMDB, Arroyo-Cabrales et al., 2002) and is
the southernmost record of the species. Tremarctos floridanus is
only known from the SMOr (San Josecito, N.L., Stock, 1950; Arroyo-
Cabrales and Johnson, 1998). Ursus americanus, the only extant
ursid in Mexico, is present in the CH-CO, SMOr (San Josecito, N.L.,
Stock, 1950; Arroyo-Cabrales and Johnson, 1998) and TMVB (Tla-
pacoya, Mex., Alvarez, 1969).
Rodentia. Rodents make up most of Mexico’s Pleistocene and
Recent mammal record (Tables 2, 9, Figs. 2 and 3). Systematic use of
micromammal collecting techniques in the last decades signifi-
cantly improved the fossil record, which includes nine families, 37
genera and 96 species; most of them are still extant. Salient features
of the record are briefly presented here. Table 9 summarizes the
major Pleistocene local faunas of Mexico, organized by position in
the morphotectonic provinces. In Fig. 15, the fossiliferous localities
are plotted on a Recent natural vegetation template to readily
compare the Pleistocene environmental settings of the locality area
with the current one there.
The temperate family Castoridae is restricted to the Irvingto-
nian El Golfo l.f. (Shaw, 1981). Rodents derived from South
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 69

Table 8
Selected localities bearing Rancholabrean Carnivora from Mexico. Sources, see text.

Morphotectonic provinces: NW, Northwestern Plains and Sierras; CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; CeP, Central Plateau; SMOr, Sierra Madre Oriental;
TMVB, Trans-Mexican Volcanic Belt; GCP, Gulf Coastal Plain; SMS, Sierra Madre del Sur; YPL, Yucatan Platform. Local faunas: A, Terapa, Son; B, La Brisca, Son; C, Cuatro
Ciénegas, Coah; D, Cueva Jiménez, Chih; E, El Cedazo, Ags; F, San Josecito, NL; G, El Cedral, SLP; H, Mina San Antonio, SLP; I, Laguna de la Media Luna, SLP; J, Chapala-Zacoalco,
Jal; K, Tequixquiac, Mex; L, Tlapacoya, Mex; M, Valsequillo, Pue; N, Mixtequilla, Ver; O, San Agustı́n, Oax; P, Gruta de Loltún, Yuc; Q, Actún Spukil, Yuc.

America include the tropical Cuniculidae, which are limited to the
YPL (QMMDB, Arroyo-Cabrales et al., 2002), along with the
Dasyproctidae, which are also known from the SMS (Cueva de
Monte Flor, Oax.; QMMDB, Arroyo-Cabrales et al., 2002). Rancho-
labrean Erethizontidae includes the extant Erethizon dorsatum and
Coendou mexicanus (Table 9). The former occurred in or north of
the TMVB (Apaxco, E. Mex.; Alvarez, 1983): Cuatro Ciénegas, Coah.
(CH-CO, Gilmore, 1947; Frazier, 1981), CeP (El Cedazo, Ags., Hib-
bard and Mooser, 1963; Frazier, 1981), SMOr (Cueva de San Jose-
cito, N.L., Arroyo-Cabrales and Johnson, 1998) and La Presita, S.L.P.
(Polaco and Butron, 1997). C. mexicanus records are restricted to
the YPL (Arroyo-Cabrales and Alvarez, 2003). Rancholabrean
Hydrochoeridae (Table 9; notice it is not Hydrochaeridae, see
Woods and Kilpatrick, 2005) includes Neochoerus aesopi and
Hydrochoerus sp. The former is largely recorded from the TMVB
(Chapala-Zacoalco Jal., Alvarez, 1971, 1983; Lucas, 2008);
Tlapacoya, E. Mex. (Alvarez, 1969), and the SMOr (Laguna de la
Media Luna, S.L.P., Hernández-Junquera, 1977), and first appears in
Mexico in the Irvingtonian El Golfo locality, Son. (Table 5).
Hydrochoerus sp. is only known from Térapa, Son., NW province
(Mead et al., 2006).
Rancholabrean Geomyidae includes four genera and seven
species, fairly widespread across the country (Table 9 and Fig. 15).
The chief records are: Cratogeomys C. bensoni [NW, Babı́cora, Son.
(Alvarez, 1983)], and Orthogeomys onerosus [SMOr, San Josecito
Cave, N.L (Russel, 1960)] among the extinct species. As for the
extant species, there are abundant records for two of them:
Thomomys umbrinus [CH-CO, Jiménez Cave, Chih. (Messing, 1986),
Cuatro Ciénegas, Coah. (Gilmore, 1947); TMVB, Tequesquinahua,
E. Mexico (Alvarez, 1983); Valsequillo, Pue. (Cruz-Muñoz et al.,
2009); SMOr, San Josecito Cave, N.L (Russel, 1960); CeP, Arroyo El
Cedazo, Ags. (Mooser and Dalquest, 1975)]; Thomomys sp. [TMVB,
Table 9
Selected localities bearing Rancholabrean Rodentia from Mexico. Sources, see text.

Morphotectonic provinces: NW, Northwestern Plains and Sierras; CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; SMOr, Sierra Madre Oriental; CeP, Central Plateau;
TMVB, Trans-Mexican Volcanic Belt; YPL, Yucatan Platform. Local faunas: A, Terapa, Son; B, La Brisca, Son; C, El Golfo, Son; D, Babı́cora, Son; E, Cuatro Ciénegas, Coah; F, Cueva
Jiménez, Chih; G, San Josecito, NL; H, La Presita, SLP; I, El Cedral, SLP; J, Mina San Antonio, SLP; K, El Cedazo, Ags; L, Chapala-Zacoalco, Jal; M, Tequixquiac, Mex; N, Tlapacoya,
Mex; O, Tequesquinahua, Mex; P, Valsequillo, Pue; Q, Santa Cruz Nuevo, Pue; R, Gruta de Loltún, Yuc.
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Tequixquiac (Hibbard, 1955)], and Cratogeomys castanops [CH-CO,
Jiménez Cave, Chih. (Messing, 1986), Cuatro Ciénegas, Coah.
(Gilmore, 1947); SMOr, San Josecito Cave, N.L (Russel, 1960),
Rancho La Amapola, Cedral, S.L.P. (Alvarez and Polaco, 1981);
Mina San Antonio (de Anda-Hurtado, 2009), CeP, Arroyo El
Cedazo, Ags. (Mooser and Dalquest, 1975); TMVB, Tequesquina-
hua, E. México (Alvarez, 1983); Tlapacoya, E. Mex. (Alvarez, 1986);
Valsequillo, Pue. (Cruz-Muñoz et al., 2009)]. Other recorded
species are Cratogeomys tylorrhinus [TMVB, Tequixquiac (Hibbard,
1955)]; Cratogeomys gymnurus [TMVB, Atotonilco El Bajo and
Chapala (Alvarez, 1983)]; Cratogeomys merriami [TMVB, Chi-
malhuacán, E., México (Alvarez, 1983); Valsequillo, Pue. (Cruz-
Muñoz et al., 2009)]; and Orthogeomys hispidus [YPL, Loltún, Yuc.,
(Alvarez, 1982)].
The Heteromyid Rancholabrean record consists of five genera
and seven species largely occurring in the CH-CO and SMOr.
(Table 9, Fig. 15). The chief records include: Chaetodipus nelsoni,
C. penicillatus [CH-CO, Cuatro Ciénegas, Coah. (Gilmore, 1947)]
and C. huastecensis [SMOr, El Abra, Tams., Dalquest and Roth,
1970)]; Perognathus flavus [SMOr, La Presita Cave, S.L.P. (Polaco
and Butron, 1997)]; Perognathus sp. [CH-CO, Jiménez Cave, Chih.
(Messing, 1986); SMOr, La Presita Cave, S.L.P. (Polaco and Butrón-
M., 1997)]; Dipodomys nelsoni [CH-CO, Jiménez Cave, Chih.
(Messing, 1986)]; D. spectabilis [CH-CO, Jiménez Cave, Chih.
(Messing, 1986); Cuatro Ciénegas, Coah. (Gilmore, 1947)];
Dipodomys sp. [NW, Rancho La Brisca, Son. (Van Devender and
Bradley, 1990); SMOr, La Presita Cave, S.L.P. (Polaco and Butrón-
M., 1997)]; Liomys irroratus [TMVB, Atotonilco El Bajo, Jal.
(Alvarez, 1983), Tlapacoya, E. Mex. (Alvarez, 1986); SMOr, San
Josecito Cave, N.L Jackway, 1958)]; Liomys sp. [NW, El Golfo de
San Clara, Son. (Shaw 1981)]; and Heteromys gaumeri [YPL; Lol-
tún, Yuc. (Alvarez, 1982)]. One extinct pocket mouse, Chaetodipus
huastecensis, is known from the Mexican Pleistocene in the SMOr
(El Abra Cave, Tams.).
Rancholabrean murids are by far the most diverse family
(Tables 2 and 9), including 24 genera and 41 species. However,
most are recorded from the TMVB and provinces north of it
(Table 9, Fig. 15). Main records include: Baiomys intermedius
[TMVB, Tequesquinahua, Mex. (Packard and Alvarez, 1965)], Per-
omyscus maldonadoi [TMVB, Tequesquinahua, Mex., (Alvarez,
1966); Valsequillo, Pue., (Cruz-Muñoz et al., 2009)], Microtus
meadensis [TMVB, Tequixquiac, Mex. (formerly Pitimys meadensis
Hibbard, Repenning, 1983)]; and Ondatra nebracense [TMVB,
Tequesquinahua, Mex. (Alvarez, 1983)]. Some arvicolines (strictly
nearctic) are environmentally very sensitive, and thus quite
important, among them are Microtus pennsylvanicus [SMOr, El
Cedral, S.L.P. (SMOr, Alvarez and Polaco, 1981)], Microtus qua-
siater [TMVB, Cuitzeo, Mich. (Garcı́a-Zepeda et al., 2008)],
Micrototus mexicanus [SMOr, San Josecito Cave, N.L (Jackway,
1958)], [TMBV, Tlapacoya, E. Mex. (Alvarez, 1969); Valsequillo,
Pue. (Cruz-Muñoz et al., 2009)]. They all indicate important
climate changes from cool and moist conditions in the Rancho-
labrean, at least locally to drier and warmer conditions in post-
Rancholabrean time.
Conversely, Hodomys alleni [SMOr, Apesco, Hgo. (Alvarez, 1983)
and Mina de San Antonio (H. alleni, de Anda-Hurtado, 2009)] and
[TMVB, Valsequillo, Pue., Cruz-Muñoz et al., 2009]; and Neotoma
magnodonta [TMBV, Tequesquinahua, Mex. (Alvarez, 1966)] and
Neotoma sp. [SMOr, La Presita, S.L.P. (Polaco and Butron, 1997)],
indicate tropical/subtropical conditions (i.e., moist and warm)
prevailing there at least locally during the Rancholabrean, which
contrast with those existing at present. Ototylomys phyllotis, an
arboreal species [YPL, Loltún, Yuc.], Oryzomys [YPL, Loltún, Yuc.
(Alvarez, 1982)], and Neotomodon alstoni [TMVB, Tequesquinahua,
Mex. (Alvarez, 1966), Valsequillo, Pue. (Cruz-Muñoz et al., 2009)] do
71
not record important biogeographic displacements, but significant
deforestation.
Neotoma, another nearctic taxon currently thriving on xeric
areas, is fairly widespread (Table 9, Fig. 15), most Rancholabrean
records though come from the TMVB: N. mexicana from Tlapacoya,
Mex (Alvarez, 1969), Valsequillo, Pue. (Cruz-Muñoz et al., 2009),
Hueyatlaco, Pue. (Alvarez, 1983) N. tlapacoyana and N. anomala
from Tlapacoya, Mex. (Alvarez and Hernández-Chávez, 1994); N.
albigula [TMVB, Zacoalco, Jal. (Alvarez, 1983); La Cinta, Mich.
(Garcı́a-Zepeda et al., 2008); and N. phenax from Chapala-Zacoalco,
Jal. (Alvarez, 1983)]. N. mexicana has also been recorded from the
SMOr [San Josecito, N.L. (Arroyo-Cabrales and Johnson, 2008)] and
the SMS [Santa Cruz Nuevo, Pue. (Tovar-Liceaga et al., 2007)].
Peromyscus, Reithrodontomys, and Sigmodon are quite widespread
(Table 9, Fig. 15), but they have been most frequently recorded
from the SMOr (e.g. La Calera, S.L.P., Jau-Mexı́a et al., 2003), and
the TMVB (Chapala, Jal., Lucas, 2008). The SMS records of P. diffi-
cilis, P. eremicus, P. maniculatus, P. P. pectoralis, and S. hispidus come
from Santa Cruz Nuevo, Pue. (Tovar-Liceaga, 2005; Tovar-Liceaga
et al., 2007), south of the TMVB.
The Rancholabrean sciurid record consists of five genera and
eight species largely occurring in the CH-CO and SMOr (Table 9).
The main records include Cynomys ludovicianus from the NW
(Babı́cora, Son., Alvarez, 1983), SMOr (Apaxco, Hgo.), and the TMVB
(Valsequillo. Pue., Cruz-Muñoz et al., 2009). Cynomys sp. is known
from the CH-CO, Jiménez Cave, Chih. (Messing, 1986), and the SMOr,
La Presita Cave, S.L.P. (Polaco and Butrón-M., 1997). Other records
are Ammospermophilus interpres [CH-CO, Cuatro Ciénegas, Coah.
(Gilmore, 1947)]; Sciurus alleni [SMOr, San Josecito Cave (Jackway,
1958)]; Sciurus yucatanensis [YPL, Loltún, Yuc. (Arroyo-Cabrales and
Alvarez, 2003)]; Spermophilus mexicanus [CH-CO, Cuatro Ciénegas,
Coah. (Gilmore, 1947)]; S. spilosoma [CH-CO, Jiménez Cave, Chih.
(Messing, 1986); Cuatro Ciénegas, Coah. (Gilmore, 1947); SMOr, San
Josecito Cave (Jackway, 1958); CeP, La Presita Cave, S.L.P. (Polaco and
Butrón-M., 1997)], and Spermophilus variegatus [CH-CO, Jiménez
Cave, Chih. (Messing, 1986); Cuatro Ciénegas, Coah. (Gilmore,
1947)]. The rockchuck Marmota flaviventris was recorded in the
SMOr, San Josecito, N.L. (Arroyo-Cabrales and Johnson, 2008),
considerably south from the southern edge of its range in New
Mexico.
Lagomorpha. Mexico’s Rancholabrean lagomorph record
includes one extinct (Aztlanolagus) and two extant (Lepus and Syl-
vilagus) leporid genera. Aztlanolagus is represented by a single
species, A. agilis [CH-CO: Cueva Jiménez (Messing, 1986; Russel and
Harris, 1986)]; SMOr: or CeP: La Calera, Zac. (Jau-Mexı́a et al.,
2000)].
Sylvilagus, a common Rancholabrean genus, is represented by
five species: S. leonensis, extinct [SMOr: San Josecito, N.L. (Cushing,
1945; Jackway, 1958; Arroyo-Cabrales and Johnson, 1995); Mina de
San Antonio, S.L.P. (Ferrusquı́a-Villafranca and de Anda-Hurtado,
2008), La Presita, S.L.P. (Arroyo-Cabrales et al., 2004)]. S. floridanus
[SMOr: San Josecito, N.L (Arroyo-Cabrales and Johnson, 1995); El
Cedral, S.L.P. (Lorenzo and Mirambell, 1981; Alvarez and Polaco,
1981); La Calera, Zac. (Jau-Mexı́a et al., 2003)]; [TMVB: Tlapacoya, E.
Mex. (Alvarez, 1969, 1986), Actopan, Hgo. (S. S. floridanus, Castillo-
Cerón et al., 1996), Barranca Seca, Ver. (S. cf. S. floridanus, Dalquest,
1961)]; [YPL: Loltún, Yuc. (Alvarez, 1983)]. S. cunicularius [SMOr: La
Calera, Zac. (Jau-Mexı́a et al., 2000), La Presita, S.L.P.] (S. S. cunicu-
larius, Polaco and Butron, 1997); [TMVB: Tocuila, E. Mex. (Dalquest,
1961)]. S. brasiliensis [YPL: Loltún, Yuc. (Alvarez, 1983)]. S. cf. S.
bachmani [PBC: El Carrizal, B.C.S. (Ferrusquı́a-Villafranca and
Torres-Roldán, 1980)]; S. audubonii [SMOr: El Cedral, S.L.P. (Polaco
and Butron, 1997); SMOr or CeP: La Calera, Zac. (Jau-Mexı́a et al.,
2000); CeP: El Cedazo, Ags. (S. audubonii, Mooser and Dalquest,
1975a)].
72 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Lepus is much less common in the record than Sylvilagus. It
includes Lepus californicus [SMOr: La Calera, Zac (Jau-Mexı́a et al.,
2000); CeP: El Cedazo, Ags. (L. L. californicus, Mooser and
Dalquest, 1975a)]. Lepus callotis [TMVB: Cueva La Nopalera y
Tepetitlán, Hgo (QMMDB, Arroyo-Cabrales et al., 2002)]; Lepus
alleni [NW: Juana Gómez, near Escuinapa, Sin. (QMMDB, Arroyo-
Cabrales et al., 2002)]. Lepus sp. [SMOr: El Cedral, S.L.P. (Alvarez
and Polaco, 1981); Mina de San Antonio, S.L.P. (Torres-Martı́nez,
1995; Ferrusquı́a-Villafranca and de Anda-Hurtado, 2008); TMVB:
Chapala, Jal. (Lucas, 2008); SMS: Santa Cruz, Pue. (Gillette et al.,
2000)].
Perissodactyla. Horses (Equus) and tapirs (Tapirus) make up
Mexico’s Rancholabrean record. Equus remains are common
throughout the country, and have been extensively collected and
studied (Reynoso-Rosales and Montellano-Ballesteros, 1994), so
that numerous nominal species have been proposed (more than 20,
Maldonado-Koerdell, 1948; Alvarez, 1965; Ferrusquı́a-Villafranca,
1978; Barrios-Rivera, 1985; Alvarez and Ocaña, 1987), but no
comprehensive evaluation of them all has been attempted. The
better-described species are described here. E. conversidens [SMOr,
San Josecito, N.L. (Arroyo-Cabrales and Johnson, 1998); El Cedral, S.
L.P. (Alberdi et al., 2003); CeP, El Cedazo, Ags. (Reynoso-Rosales and
Montellano-Ballesteros, 1994); TMVB, numerous localities in
Jalisco, Michoacán, Edo. Mexico, Puebla, Morelos and Hidalgo;
major sites occur in Mexico’s Basin, E. Mex.-D.F. (Owen, 1869;
Hibbard, 1955); Chapala-Zacoalco, Jal. (Downs, 1958; Lucas, 2008);
YPL, Yucatán (Ray, 1957); Loltún, Yuc. (Arroyo-Cabrales and Alvarez,
2003); E. conversidens was the first horse species described from
Mexico (Owen, 1869)]. E. excelsus [NW, Térapa, Son. (Carranza-
Castañeda and Roldán-Quintana, 2007)]. E. mexicanus [SMOr: El
Cedral, S. L.P. (Alberdi et al., 2003); CeP: El Cedazo, Ags. (Mooser,
1958); Laguna de la Media Luna, S.L.P. (Hernández-Junquera, 1977);
TMVB, largely from Chapala-Zacoalco, Jal. (Downs, 1958; Lucas,
2008), Mexico’s Basin, E. Mex.-D.F. (Hibbard, 1955), and Valle de
Puebla, Pue. (Freudenberg, 1922; Silva-Bárcenas, 1993; Pichardo,
2000a, 2004); QMMDB (Arroyo-Cabrales et al., 2002)]. E. pacificus
[CeP: El Cedazo, Ags. (Mooser, 1958); QMMDB (Arroyo-Cabrales
et al., 2002); TMVB: Mexico’s Basin, E. Mex.-D.F. (Hibbard, 1955)]. E.
parastylidens and E. tau [CeP: El Cedazo, Ags. (Mooser, 1958)]. The
latter three species merit further consideration. E. pacificum was
previously known from North America’s Pacific Coast, hence its
presence in the CeP and TMVB would greatly extend its geographic
range southward. E. parastylidens and E. tau are only known from El
Cedazo, Ags., thus bringing to at least five the number of Equus
sympatric species, which if not impossible seems unlikely. As
a working solution, all nominal species from El Cedazo could be
regarded now as an Equus species complex, until they are formally
revised.
The Rancholabrean record of tapirs includes the extant species
Tapirus bairdii [YPL: Actún Lara, Yuc (Hatt, 1953)] the extinct Tapirus
haysii [SMOr: Mina, N.L. (Franzen, 1993, 1994; Arroyo-Cabrales
et al., 1996); San Josecito, N.L. (Arroyo-Cabrales et al., 1996); El
Cedral, S.L.P. (Alvarez and Polaco, 1981; Arroyo-Cabrales et al.,
1996)], and Tapirus sp. [NW: Térapa, Son. (Mead et al., 2006); TMVB:
Chapala, Jal. (Lucas, 2008)]. Although T. californicus is known from
California’s Irvingtonian, and thus is geographically close to the El
Golfo l.f. (Jefferson, 1989) in Sonora, the geologic setting and dating
of the Térapa locality (Mead et al., 2006), also from Sonora, suggests
that the Rancholabrean tapir is a southern immigrant rather than
a survivor species.
Artiodactyla. Mexico’s Rancholabrean artiodactyl record is
large, unevenly scattered occurring in nearly all MPs. It includes five
families (Antilocapridae, Bovidae, Camelidae, Cervidae and Tayas-
suidae), 18 genera and some 32 species (Arroyo-Cabrales et al.,
2002, 2009). The TMVB and SMOr have most of the localities with
these groups. The record as a whole requires a comprehensive
review to eliminate taxa that have been inappropriately placed
stratigraphicaly or taxonomicaly and to identify others that have
been overlooked. Table 10 summarizes the QMMDB (Arroyo-Cab-
rales et al., 2002) relevant information, stressing the major local
faunas. The following comments focus on taxa that are strati-
graphically or biogeographically more significant.
The Antilocapridae (pronghorns) were diverse (four genera and
seven species), and widely distributed during the Rancholabrean,
yet at present, only a single species remains, Antilocapra americana,
which is recorded in the CH-CO (three sites, Lundelius, 1975, 1980;
Russel and Harris, 1986) and SMOr (two localities, Kurtén, 1975;
Hernández-Junquera, 1977). Capromeryx and Stockoceros are more
widespread occurring in the NW (two localities, Rea, 1980; Van
Devender et al., 1985; Mead et al., 2006), CeP (El Cedazo, Ags.,
Mooser, 1958; Mooser and Dalquest, 1975a), SMOr (six localities
including San Josecito and Minas, N.L., Furlong, 1925; Arroyo-Cab-
rales and Johnson, 1998), TMVB (several sites including Tequix-
quiac, Mex. and Valsequillo, Pue., Freudenberg, 1921; Guenther and
Bunde, 1973), and the GCP Mixtequilla, Ver., Polaco, 1995). Tetra-
meryx is less common occurring in the CeP (El Cedazo, Ags., Mooser,
1958; Dalquest, 1974; Mooser and Dalquest, 1975a), TMVB
(including Chapala-Zacoalco, Jal., Downs, 1958; Lucas, 2008), and
the GCP (Polaco, 1995).
The Bovidae record is diverse and abundant. Bison, the index
genus used to indicate the beginning of the Rancholabrean (Bell
et al., 2004) includes five named species (B. alaskensis, B. antiquus, B.
bison, B. latifrons and B. priscus), which largely come from localities in
the CeP (El Cedazo, Ags., Mooser, 1958; Mooser and Dalquest, 1975a),
SMOr (including Minas., N.L. and Laguna de la Media Luna, S.L.P.,
Hernández-Junquera, 1977), TMVB (many localities including Cha-
pala-Zacoalco, Jal., Tequixquiac and Tlapacoya, Mex., and Valsequillo,
Pue., Downs, 1958; Hibbard and Villa-Ramı́rez, 1950; Hibbard, 1955;
Pichardo, 1999; Lucas, 2008). Bison is much less common in the PBC,
NW and CH-CO (Skinner and Kaiser, 1947; Ferrusquı́a-Villafranca and
Torres-Roldán, 1980). B. alskensis and B. bison are the most wide-
spread. In addition to the above records, B. bison is also known from
the YPL (Loltún, Yuc., Arroyo-Cabrales and Alvarez, 2003). Bison sp.
(probably B. latifrons) is recorded from La Mixteca and Valles Cen-
trales, Oax. (Ferrusquı́a-Villafranca, 1976 and unpublished data).
Euceratherium collinum is recorded from sites in the NW (La Guitarra,
Son., Hibbard, 1955), SMOr (including San Josecito, N.L., Cushing,
1945; Arroyo-Cabrales and Johnson, 1998) and TMVB (Tequixquiac,
Mex., Hibbard, 1955). While Oreamnos is a common Rancholabrean
taxon north of Mexico (Bell et al., 2004), the extinct species,
Oreamnos harringtoni is only present in the SMOr (San Josecito, NL.,
Arroyo-Cabrales and Johnson, 2008). Ovis canadensis occurs in the
CH-CO [Cuatro Ciénegas, Coah. (Gilmore, 1947; Lundelius, 1980); La
Candelaria, Coah. (Aveleyra-Arroyo de Anda, 1956)] and TMVB [Tula,
Hgo. (Valadez and Paredes, 1988)]; this record might be Early
Holocene, because there is no compelling evidence of its Rancho-
labrean age). O. canadensis, like Bison, first appears in the Rancho-
labrean in North America (Bell et al., 2004).
The Rancholabrean Camelidae record is also diverse (four
genera and seven species), but less widespread than that of the
previous families, being rare south of the TMVB. Four species
belong to Camelops, the most diverse genus. C. hesternus had the
greatest range. Occurring in the NW (Mead et al., 2006), CeP (El
Cedazo, Ags., Mooser, 1958; Mooser and Dalquets, 1975b), SMOr
(San Josecito, Stock, 1943; Arroyo-Cabrales and Johnson, 1998) and
TMVB (including Chapala-Zacoalco, Jal., Tequixquiac, Mex., and
Valsequillo. Pue., Downs, 1958; Furlong, 1925; Hibbard, 1955;
Guenther, 1968; Lucas, 2008). By contrast, C. minidokae, C. mex-
icanus, and C. traviswhitei are known from single/few localities each
in the TMVB (Puebla localities, Guenther and Bunde, 1973), SMOr
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 73

Table 10
Selected localities bearing Rancholabrean Artiodactyla from Mexico. Sources, see text.

Morphotectonic provinces: NW, Northwestern Plains and Sierras; CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; CeP, Central Plateau; SMOr, Sierra Madre Oriental;
TMVB, Trans-Mexican Volcanic Belt; GCP, Gulf Coastal Plain; SMS, Sierra Madre del Sur; YPL, Yucatan Platform; Local faunas: A, Terapa, Son; B, La Brisca, Son; C, Cuatro
Ciénegas, Coah; D, Cueva Jiménez, Chih; E, El Cedazo, Ags; F, San Josecito, NL; G, Minas, NL; H, Mina San Antonio, SLP; I, Laguna de la Media Luna, SLP; J, La Presita, SLP; K,
Chapala-Zacoalco, Jal; L, Tequixquiac, Mex; M, Tlapacoya, Mex; N, Valsequillo, Pue; O, Hueyatlaco, Pue; P, Moroleón, Gto; Q, Mixtequilla, Ver; R, San Agustı́n, Oax; S, Gruta de
Loltún, Yuc; T, Actún Spukil, Yuc. * Extinct species.

(San Josecito, N.L., Arroyo-Cabrales and Johnson, 1998), and CeP
respectively (El Cedazo, Ags., Mooser and Dalquest, 1975a,b). C.
conidens is recorded only in the TMVB (Tequixquiac, Mex., Cope,
1884). Hemiauchenia macrocephala is known from the PBC (El
Carrizal, B.C.S., Ferrusquı́a-Villafranca and Torres-Roldán, 1980),
SMOr (Minas, N.L.) and TMVB [Tequixquiac, Mex. and Valsequillo,
Pue. (Hibbard, 1955; Guenther and Bunde, 1973)]. At present, no
members of the Camelidae live in Mexico.
The Cervidae (four genera and six species) are as diverse as
camelids, but more widespread. Cervus elaphus and Mazama
americana are known from single localities in the CH-CO (Cuatro
Ciénegas, Coah., Gilmore, 1947) and GCP (Mixtequilla, Ver., Polaco,
1995). Navahoceros fricki is recorded from the CeP (El Cedazo, Ags.,
Mooser and Dalquest, 1975a), SMOr (San Josecito, N.L, Kurtén, 1975),
and SMS (San Agustı́n, Oax., Kurtén, 1975). Odocoileus includes three
species: O. halli, O. hemionus and O. virginianus. The first is extinct,
occurring in the CeP (El Cedazo, Ags., Mooser, 1958; Mooser and
Dalquest, 1975a), and TMVB (Tlapacoya, Mex., Alvarez, 1986). O.
hemionus (mule deer) is present in these MPs: CeP (El Cedazo, Ags.,
Mooser and Dalquest, 1975a), SMOr, and TMVB (Chapala-Zacoalco,
74 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Jal., Downs, 1958), whereas O. virginianus is more widespread, being
recorded from CH-CO, SMOr (two sites in San Luis Potosı́, Hernán-
dez-Junquera, 1977; Polaco and Butron, 1997), TMVB (Chapala-
Zacoalco, Jal. and Tlapacoya, Mex., Downs, 1958; Alvarez, 1986), GCP,
and YPL (Loltún and Actún Spukil, Yuc., Arroyo-Cabrales and Alvarez,
2003).
The Tayassuidae are the least diverse family, including two
genera and four species. The extant Pecari tajacu is recorded from
the YPL (Loltún and Actún Spukil, Yuc., Arroyo-Cabrales and
Alvarez, 2003). Platygonus, an extinct genus includes Platygonus
‘‘alemanii,’’ P. compressus and P. ticuli. The first is known from the
TMVB (Moroleón, Gto., Duges, 1891; but see Cope, 1884). P. com-
pressus, a Rancholabrean index (Bell et al., 2004), is present in the
CeP (El Cedazo, Ags., Mooser, 1958; Mooser and Dalquest, 1975a)
and TMVB (Tequixquiac, Mex. and Valsequillo, Pue., von Thenius,
1970; Guenther, 1968; Guenther and Bunde, 1973). P. ticuli is only
recorded from the TMVB (Chapala-Zacoalco, Jal., Downs, 1958;
Mones, 1973b; Lucas, 2008). The systematics and nomenclature of
Platygonus species are still unsettled (Barrios-Rivera, 1985).
Notoungulata and ?Litopterna. These are the taxa least rep-
resented in Mexico’s Pleistocene record. The notoungulate Mix-
otoxodon sp. is known from Hihuitlán, Mich., TMVB, and La
Estribera, Ver, GCP (Polaco et al., 2004). Both extend northward of
the range of notoungulates from La Pasión, Guatemala, the previous
northernmost record (Woodburne, 1969). In eastern Mexico City
[TMVB], while building a large market, workers dug out mammal
remains, but unfortunately very little material was salvaged (a
single tooth and very fragmentary post-cranial bones). The highly
hypsodont tooth is a molar with an occlusal pattern that resembles
that of macrauchenid litopterns. On this basis, the material was
tentatively and questionably identified as a litoptern (Ferrusquı́a-
Villafranca and Malvido-Arriaga, 2006).
Proboscidea. The Rancholabrean record of proboscideans in
Mexico is abundant and widespread, occurring in all MPs. Remains
of Mammuthus, a common Rancholabrean genus (Agenbroad,
1989), have been recovered from most of the states in this country
(Arroyo-Cabrales et al., 2003, 2007a,b; Polaco, 2003), with a large
concentration in the TMVB, CeP and SMOr, whereas in the PBC and
YPL the number of localities is very small. Precise taxonomic
identification and nomenclature is a major problem, because this
vast record has been studied by numerous authors holding
different views (Aveleyra-Arroyo de Anda, 1955; Pichardo, 1960,
2001; Arroyo-Cabrales et al., 2007b). M. columbi hayi is known from
El Mezquital, B.C.S. (Arroyo-Cabrales et al., 2007b) (PBC) and Arizpe,
Son. (Lucas and González-León, 1996) (NW), however, Arroyo-
Cabrales et al. (2007a,b) question these identifications. M. columbi
is present in most MPs clustering in the TMVB and CeP (Polaco,
1982; Arroyo-Cabrales et al., 2003, 2007a,b; Polaco, 2003). M.
imperator has recently been reported from Chapala, Jal. by Lucas
(2008), but this requires confirmation. Stegomastodon S. primitivus
is reported by Alberdi et al. (2002, 2004) from Chapala, Jal., TMVB,
while Stegomastodon sp. occurs in Valsequillo, Pue. (Alberdi and
Corona-M. 2005) (TMBV). The presence of Stegomastodon in the
Rancholabrean Chapala and Valsequillo local faunas is noteworthy,
because its temporal range is mainly Blancan-Irvingtonian. Lucas
(2008) argues that this record should be assigned to the Blancan,
and clearly additional work is needed to settle this issue. Remains
of Cuvieronius, a genus with a long temporal range (McKenna and
Bell, 1997) have been recorded from nearly 30 localities, which
largely lie in the TMVB (eastern part), and much less so in the SMS
and CHI (Corona-M. and Alberdi, 2006; Arroyo-Cabrales et al.,
2007a,b; Lucas 2008). C. tropicum seems to be present there
(Montellano-Ballesteros, 2002). Cuvieronius sp. is also known from
the Irvingtonian of Chiapas, as was already discussed. Mead et al.
(2006) report Cuvieronius sp. from Térapa, Son. (NW). Mammut
americanum, a common Rancholabrean species throughout North
America (Bell et al., 2004), is recorded from some 15 localities,
which chiefly lie in the TMVB (eastern part, about eight), SMOr and
CeP (Polaco et al., 2001; Arroyo-Cabrales et al., 2007a,b).
3.2. Mexico’s Pleistocene environment and faunal response to
climate change: a first approach
The information on Mexico’s Pleistocene terrestrial environ-
ment, both physical and biotic is truly enormous and highly scat-
tered; it mainly resulted from research conducted during the last
decades on very diverse subjects, with little or no coordination
among the different participant working groups; further, the
explicit aim of any particular research may not necessarily be
contributing knowledge on the environment per se. The chief
topics include lake sediments (Leyden et al., 1994; Lozano-Garcı́a
and Ortega-Guerrero, 1997; Ortega-Ramı́rez et al., 1998; Caballero-
Miranda et al. 1999, 2002; Ortega-Guerrero and Urrutia-Fucu-
gauchi, 1997; Metcalfe et al., 2000; Ortega-Guerrero et al., 2000);
other depositional settings (White and Valastro, 1984; Ortega-
Ramı́rez et al., 2004); palynomorphs (Foreman, 1955; Clisby and
Sears, 1955; Sears and Clisby, 1955; Bradbury, 1971, 1989; Meyer,
1973; Ohngemach, 1973, 1977; Brown, 1984; González-Quintero,
1986; Straka and Ohngemach, 1989; Metcalfe, 1992; Lozano-Garcı́a
and Ortega-Guerrero, 1994; Anderson and Van Devender, 1995;
Leyden et al., 1996; Ortega-Ramı́rez et al., 1998; Caballero-Miranda
et al., 1999; Canul-Montañez, 2008); diatoms (Bradbury, 1971, 1989;
Caballero-Miranda et al., 1999, 2002; Metcalfe, 1992; Metcalfe et al.,
1989, 1994; Rico et al., 1995, 1998); mammals (Alvarez, 1983;
Montellano-Ballesteros, 1992; Arroyo-Cabrales and Johnson, 1998;
McDonald, 2002; Gillette et al., 2000; Mead et al., 2006); other
vertebrates (Corona-M., 2002; Castillo-Cerón et al., 1996); paleosols
(Cervantes-Borja et al., 1997; Gama-Castro et al., 2004; Ortega-
Guerrero et al., 2004; Solleiro-Rebolledo et al., 2006); and
archeological studies pertaining to the early presence of humans
(Lorenzo and Mirambell, 1986).
To even summarily review this vast amount of information would
require a multidisciplinary team, and is clearly out of place here.
Moreover, given the differences in objectives, scope and method-
ologies, the results of particular research may not readily compare or
be congruent with those of others, even when working in the same
areas (Leyden et al., 1996; Metcalfe, 1997; Arroyo-Cabrales and
Alvarez, 2003 for paleoclimate in Yucatán). Under these circum-
stances, the review of Pleistocene environmental information is
restricted to that provided by the palynomorphs and paleosols, as
they provide a means of independent references to compare with
the data derived from the mammals themselves.
3.2.1. Pleistocene environmental indicators (a), the palynologic
record: a review
The results of the review (Foreman, 1955; Clisby and Sears, 1955;
Sears and Clisby, 1955; Ohngemach, 1973, 1977; Bradbury, 1971,
1989; Meyer, 1973; Brown, 1984, 1985; González-Quintero, 1986;
Straka and Ohngemach, 1989; Metcalfe, 1992; Leyden et al., 1994,
1996; Lozano-Garcı́a and Ortega-Guerrero, 1994; Anderson and Van
Devender, 1995; Lozano-Garcı́a and Ortega-Guerrero, 1997; Loz-
ano-Garcı́a and Xelhuantzi-López, 1997; Ortega-Ramı́rez et al.,
1998; Caballero-Miranda et al., 1999; Lozano-Garcı́a et al., 2005;
Canul-Montañez, 2008) are presented in Fig. 11, where the major
palynofloral localities are set on a morphotectonic template, with
the current Nearctic Region/Neotropical Realm boundary also
plotted. The geographic distribution of the localities is very uneven,
as about one-third lies in the TMVB, and the remaining 15 are
dispersed in the other MPs. Thus, most localities (about 2/3) lie in
the TMVB or in the morphotectonic provinces north of it.
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
The dating of the localities varies considerably. Only a few have
independent radio-isotopic (14C) or paleomagnetic dating (Brown,
1984; Bradbury, 1989; Metcalfe, 1992; Lozano-Garcı́a and Ortega-
Guerrero, 1994; Urrutia-Fucugauchi et al., 1994; Caballero-Miranda,
1997; Lozano-Garcı́a and Ortega-Guerrero, 1997; Ruiz-Martı́nez
et al., 2000; Lozano-Garcı́a et al., 2005). Other localities are dated
on the basis of associated available stratigraphic volcanic bodies
amenable to radio-isotopic dating, or paleontologic (largely fossil
mammals) information (Alvarez, 1986). Most known Quaternary
palynofloras are of very Late Pleistocene age, rarely exceeding
40 ka. The majority are 20 ka old or less. Few palynological studies
are from deposits older than the Rancholabrean NALMA.
Fossil pollen assemblages are samples of past vegetation
communities (or in general of biomes) that lived in the vicinity of
the fossil-bearing locality area at the time of deposition. Thus
pollen assemblages readily indicate broad humidity and tempera-
ture conditions of the community’s physical setting at the time of
deposition. To avoid misinterpretation, due allowance must be
made to discriminate between distant (allochthonous) and local/
indigenous (autochthonous) pollen source components.
Changes in temperature and humidity may occur seasonably,
and are affected by latitude, altitude or local topographic features.
In turn, such changes could also be short-term or long-term, and
may occur gradually or suddenly. These environmental conditions,
their change or permanency, as well as their possible significance
could only be established through systematic studies of long
sections rather than of short stratal intervals. Unfortunately not
many studies (those whose results are plotted in Fig. 11 and a few
others) meet these requirements.
The most common and abundant taxa occurring in Mexico’s
Pleistocene sediments are mainly wind pollinated trees including
Abies, Pinus, Picea, Podocarpus, Cupressus, Juniperus, Taxodium,
Tsuga, Quercus, Alnus, Carya, Celtis, Fagus, Fraxinus, Juglans, Liquid-
ambar, Engelhardtia, Ilex, Myrica, Populus, Salix, Moraceae, Tiliaceae,
and Ulmaceae, as well as wind pollinated herbs such as Poaceae,
Amaranthaceae-Chenopodiaceae, Plantaginaceae and Asteraceae.
There are however, other taxa restricted to certain depositional
environments: such taxa include Arenaria, Bursera, Eriocaulon,
Asplenium, Selaginella, Isoetes, Malvaceae, Cyperaceae, Juncaceae,
Eleocharis, Piscidia, Prosopis, Luzula, Polygonum, Ranunculus, Berula,
Datura, Hedyosmun, Ruppia, Heteranthera, Hydrocotyle, Ludwigia,
Myriophyllum, Nymphaea, Nuphar, Potamogeton, Roripa, Sagittaria,
Sphagnum, Schoenoplectus, Thalictrum, Typha, and Utricularia.
3.2.1.1. The Late Pleistocene Trans-Mexican Volcanic Belt palynofloral
record as an environmental indicator. The palynofloral record of this
morphotectonic province is far better than that of the others (Fig. 9),
coming from the fine-grained clastic deposits laid down in this
region’s largely endorreic basins in the States of Jalisco (e.g. Lake
Chapala), Michoacán (e.g. Lake Pátzcuaro), Mexico (e.g. Lakes Tex-
coco and Chalco), Puebla (Valsequillo area), Tlaxcala and Hidalgo
(Foreman, 1955; Clisby and Sears, 1955; Sears and Clisby, 1955;
Ohngemach, 1973, 1977; Bradbury, 1971, 1989; Meyer, 1973; Ohn-
gemach, 1973, 1977; Brown, 1984; González-Quintero, 1986; Straka
and Ohngemach, 1989; Metcalfe, 1992; Leyden et al., 1994, 1996;
Lozano-Garcı́a et al., 1993; Lozano-Garcı́a and Ortega-Guerrero,
1994; Anderson and Van Devender, 1995; Lozano-Garcı́a and
Ortega-Guerrero, 1997; Lozano-Garcı́a and Xelhuantzi-López, 1997;
Ortega-Ramı́rez. et al., 1998; Caballero-Miranda et al., 1999; Canul-
Montañez, 2008). No major lake deposits are present in the TMVB’s
eastern third. The TMVB palynological record is summarized in Table
11, including five divisions, seven classes, 42 orders, 58 families, and
54 named genera, and thus the fossil recorded diversity is compa-
rable to the Recent pollen rain. The record indicates compositional
fluctuations through time that largely involve greater or lesser
75
percentages of pine to pine-oak forests and deciduous forest, as well
as of other vegetation type components, seemingly reflecting
changes in altitude of the timber line, in turn associated with
temperature and humidity changes, which most probably were
driven by glaciation (Clisby and Sears,1955; Bradbury,1997; Lozano-
Garcı́a and Ortega-Guerrero, 1997; Metcalfe et al., 2000).
However, there is not a strict correlation between the Pleisto-
cene advances/retreats of the Laurentide Glacier and advances/
retreats of coeval climate belts in southern subtropical/tropical
latitudes, because other factors like relief, atmospheric and oceanic
circulation particular patterns at specific times strongly influence
this general trend at the local/regional level. Mexico lies at such
latitudes, thus discriminating the impact of temperature and
humidity changes in the vegetation, lake sediment and lake biota is
complicated and may lead to different conclusions (Bradbury, 1989;
Leyden et al., 1994; Metcalfe et al., 1997, 2000). Under these
circumstances, it appears that minimal temperatures have been
recorded across the TMVB at 35 ka, 23 ka and 16 ka, which correlate
well with a significant increase of the pine and pine-oak forest
components corresponding to these ages, and with thermal and
salinity changes in the lake sediments, largely disclosed by the
diatom assemblages and by the sediment chemistry (Brown, 1984;
Steininger, 1988; Metcalfe et al., 1989, 1991, 1994). Such vegetation
changes largely involve changes in the position of timber line
altitudinal position, with very little (if any) latitudinal vegetation
community displacements (González-Quintero, 1986; Rzedowski,
1993). The intricate topography of this morphotectonic province,
and the different sampling density throughout the TMVB do not
allow us to pursue this matter further at present. Nonetheless, the
pollen record seems to indicate that during the Rancholabrean, the
TMVB had a cooler, more humid climate regime, which allowed the
development of numerous freshwater lakes in the low-lying basins;
at present, such lakes are severely reduced or altogether absent.
A recent palynological study in Tehuacán, Pue (Canul-Mon-
tañez, 2008), which lies in the SMS, just south of the TMVB
(eastern sector), indicates that the xeric flora now prevailing there
may be Recent, perhaps not older than 12–10 ka, thus lending
support to the above contention. Further, one of the authors (E.
Martı́nez-Hernández) thinks that the palynological evidence
discloses that at least for the 25–12 ka interval, there was greater
humidity than present in the TMVB, as indicated by the wide
extent of cloud forest, as well as cooler temperatures, as shown by
the presence of Picea.
At any rate, this pattern of major altitudinal vegetation shifts
associated with minor latitudinal displacements seems to be also
recognizable in other morphotectonic provinces. Fig. 12 is a plot of
the palynofloral localities on a present-day natural vegetation
template, with the Nearctic Region/Neotropical Domain boundary
also marked. Comparing the vegetation type inferred from the
palynologic assemblage(s) of a particular locality with that of the
vegetation living at present in the same area/region, little lat-
itudinal displacement is apparent. On the other hand, longitudinal
(E–W) expansion of grassland and shrub forest at the expense of
pine and pine–oak forest seems to have occurred in the CH-CO, CeP
and northern GCP. However, the scarcity of sampling for the whole
country prevents us from positively proposing this conclusion.
Even in regions where a large, multidisciplinary database is
available, such as the Chihuahuan Desert (i.e. the CH-CO and
adjacent Texas, New Mexico and Arizona) for the last 40 ka
(Betancourt et al., 1990 and literature therein; Metcalfe et al., 2002),
data have been interpreted differently by diverse authors, which is
not surprising given this region’s large size (w2200 km long and
800 km wide at the widest sector), geographic position (spanning
w10 latitude degrees), and varied geomorphic setting (200–
2000 m asl altitudinal range), so that latitude, altitude, and age of
76 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Fig. 9. Mexico’s chief Pleistocene palynofloral localities set on the morphotectonic provinces template pictured in Fig. 1. Time frame adapted from Bell et al., 2004. Chief sources:
Foreman, 1955; Clisby and Sears, 1955; Sears and Clisby, 1955; Ohngemach, 1973, 1977; Bradbury, 1971, 1989; Meyer, 1973; Brown, 1984; González-Quintero, 1986; Straka and
Ohngemach, 1989; Metcalfe, 1992; Leyden et al., 1994, 1996; Lozano-Garcı́a, and Ortega-Guerrero, 1994; Anderson and Van Devender, 1995; Lozano-Garcı́a and Ortega-Guerrero,
1997; Lozano-Garcı́a and Xelhuantzi-López, 1997; Ortega-Ramı́rez et al., 1998; Caballero-Miranda et al., 1999; Canul-Montañez, 2008; Martı́nez-Hernández, E., unpublished data.

the material sampled, as well as its particular environmental
sensitivity influence the interpretation results. Extensive studies of
pack rat (Neotoma ssp.) middens by Van Devender and associates
(Van Devender et al., 1987; Van Devender, 1990a,b; Van Devender
and Bradley, 1990) disclose vegetation composition changes for this
time interval, seemingly due to both latitudinal and altitudinal
displacements related to temperature and humidity shifts, ulti-
mately driven by advance/retreat of the Laurentide Glaciar during
the Late Pleistocene (Wisconsinan).
The net result was an increase/reduction of the woodland s.l.
communities, in close, but opposite lockstep fashion with an
increase/reduction of the xeric s.l. communities. However, the
vegetation changes were not uniform across the region, so that the
northern and southern sectors responded differently (latitude
factor), as did the lower and higher elevation areas (altitude factor).
Both temperature and humidity paleoclimate factors influenced the
vegetation at any particular place and time, but to discern their
particular role is not straightforward. It appears that in the Late
Wisconsinan (w22–11 ka), in general the Chihuahuan Desert (i.e.,
chiefly the CH-CO morphotectonic province) had a more equable,
more humid climate regime, with cooler summers and mild
winters, that allowed an extensive biotic mixing, generating an
enormous diversity, unparalleled at present, whereby species of
contrasting ecological requirements (thermophilous and
temperate-prone species, or humidity-prone and xeric species)
could occur in proximity. This phenomenon had already been
detected (Bell et al., 2004).
In summary, the composition of the vegetation during the Late
Pleistocene is broadly similar to the present one (Fig.10). Differ-
ences largely pertain to the forest extension, because its altitudinal
limits were modified according to climate changes. The TMVB
Rancholabrean cloud forest (mountain forest) is a possible excep-
tion to this model.
3.2.1.2. Lake sedimentary information complementary to the paly-
nologic record. Most palynologic studies performed in the TMVB
and elsewhere are directly or indirectly related to paleolimnologic
research (Bradbury, 1989; Urrutia-Fucugauchi et al., 1995; Lozano-
Garcı́a and Ortega-Guerrero, 1997; Lozano-Garcı́a and Xelhuantzi-
López, 1997; Ortega-Guerrero and Urrutia-Fucugauchi, 1997;
Caballero-Miranda and Ortega-Guerrero, 1998; Caballero-Miranda
et al., 1999; Ortega-Guerrero et al., 2000). The goal is to be able to
characterize the Pleistocene climate regime and change, given that
lake level, depth, water chemistry, sedimentary structure and
architecture, as well as autochthonous or allochthonous fossil
content are strongly influenced by the climatic regime prevailing at
the time of deposition. Further, lake sediments frequently contain
volcanic ash, which is amenable to radio-isotopic dating (e.g.,
Ortega-Guerrero and Urrutia-Fucugauchi, 1997). The signature of
paleomagnetic secular variation calibrated with radio-isotopic
dating in one site, could be recognized elsewhere, thus providing
means to indirectly date other sequences (e.g. Lake Chalco, Ortega-
Guerrero and Urrutia-Fucugauchi, 1997; Caballero-Miranda and
Ortega-Guerrero, 1998). However, the interpretation of the record
is not straightforward, because lake features may also reflect
different factors, e.g. lake level rise may be caused by higher
humidity and pluviosity than usual, temperature decrease, and
increased water supply caused by river piracy. Lake shallowing may
indicate higher evaporation rate driven by higher temperatures,
less humidity and pluviosity, higher sedimentary supply than usual,
volcanic ash input, reduced water supply by river channel
obstruction or diversion, or even drainage caused by fracturing and/
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 77

Table 11 Table 11 (continued)

Late Pleistocene (Rancholabrean) Trans-Mexican Volcanic Belt flora identified from
its palynological record. Taxonomical arrangement sensu Cronquist (1981); ranks Gentianales
employed (from higher to lower): Division, Class, Order, Family and Genus. Chief Loganiaceae
sources: Foreman, 1955; Clisby and Sears, 1955; Sears and Clisby, 1955; Bradbury, Buddleja
1971, 1989; Ohngemach, 1973, 1977; Meyer, 1973; Brown, 1984; González-Quintero, Hamamelidales
1986; Straka and Ohngemach, 1989; Metcalfe, 1992; Lozano-Garcı́a et al., 1993; Hamamelidaceae
Lozano-Garcı́a and Ortega-Guerrero, 1994; Caballero-Miranda et al. 1999; Canul- Liquidambar
Montañez, 2008; Martı́nez-Hernández, E., unpublished data. Juglandales
Juglandaceae
Bryophyta Carya
Sphagnopsida Engelhardtia
Sphagnales Juglans
Sphagnaceae Lamiales
Sphagnum Lamiaceae
Lycopodiophyta Magnoliales
Isoetopsida Magnoliaceae
Isoetales Drymys
Isoetaceae Malpighiales
Isoetes Hypericaceae
Selaginellopsida Hypericum
Selaginellales Malvales
Selaginellaceae Malvaceae
Selaginella Tiliaceae
Pteridophyta Myricales
Polypodiopsida Myricaceae
Filicales Myrica
Aspleniaceae Myrtales
Asplenium Melastomataceae
PinoPhyta Myrtaceae
Coniferopsida Onagraceae
Coniferales Ludwigia
Cupressaceae Nymphaeales
Juniperus Nymphaeaceae
Taxodium Nuphar
Cupressus Nymphaea
Pinaceae Piperales
Abies Chloranthaceae
Picea Hedyosmun
Pinus Plantaginales
Tsuga Plantaginaceae
Podocarpaceae Polygonales
Podocarpus Polygonaceae
Magnoliophyta Polygonum
Magnoliopsida Ranunculales
Apiales Ranunculaceae
Araliaceae Ranunculus
Hydrocotyle Thalictrum
Asterales Rosales
Asteraceae Rosaceae
Capparales Holodiscus
Brassicaceae Salicales
Rorippa Salicaceae
Caryophyllales Populus
Amaranthaceae – Chenopodiaceae Salix
Caryophyllaceae Santalales
Celastrales Santalaceae
Aquifoliaceae Arceutholobium
Ilex Sapindales
Dipsacales Burseraceae
Adoxaceae Bursera
Viburnum Saxifragales
Caprifoliaceae Haloragidaceae
Valerianaceae Scrophulariales
Valeriana Lentibulariaceae
Ericales Utricularia
Ericaceae Oleaceae
Fabales Fraxinus
Fabaceae Solanales
Piscidia Solanaceae
Prosopis Datura
Fagales Urticales
Betulaceae Moraceae
Alnus Ulmaceae
Betula Celtis
Fagaceae Urticaceae
Fagus (continued on next page)
78 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Table 11 (continued) Ohngemach, 1989; Metcalfe, 1992; Lozano-Garcı́a et al., 1993;

LILIOPSIDA Lozano-Garcı́a and Ortega-Guerrero, 1994; Caballero-Miranda and
Alismatales Ortega-Guerrero, 1998; Caballero-Miranda et al., 1999; Canul-
Alismataceae Montañez, 2008). Their sediments carry palynonomorph assem-
Sagittaria blages (Fig. 9), and in most, there are associated important Pleis-
Poramogetonaceae
Potamogeton
tocene mammal-bearing localities (Figs. 5 and 8). Outside the
Ruppiaceae TMVB, very few places have been studied. One exception is El
Ruppia Fresnal, Chih., Babı́cora, Chih., CH-CO morphotectonic province
Cyperales (Ortega-Ramı́rez et al., 1998, 2004).
Cyperaceae
Besides the geographic bias, there is also a strong time bias, so
Eleocharis
Schoenoplectus that most lake sediment cores span only the latest Pleistocene–
Poaceae Early Holocene (Bradbury, 1997; Ortega-Ramı́rez et al., 1998; Met-
Eriocalulales calfe et al., 2000), rarely reaching beyond 35 ka. Lakes Texcoco and
Eriocaulaceae Chalco are among the most studied. Radioisotopic dating (14C
Eriocaulon
Juncales
method applied to organic matter-rich strata; Lozano-Garcı́a et al.,
Juncaceae 1993; Caballero-Miranda, 1997) brackets recorded events between
Luzula 34 and 14 ka, and supplementary data from other lake sites in the
Liliales Mexican bridge the 12–2 ka span, thus furnishing glimpses of the
Pontederiaceae
lake history from the Middle Wisconsinan to the Holocene. Making
Heteranthera
Typhales due allowance for methodological differences, studies of diatoms
Typhaceae and of other evidence (water chemistry and sediment input among
Typha others) permit the recognition of a largely moister and cooler
climate regime in late Middle Wisconsinan–Late Wisconsinan time
(ca. 24–12 ka) than at present, followed by warmer and somewhat
or faulting. Not surprisingly, the evidence from a given site may be drier conditions prevailing in Mexico’s basin, and elsewhere across
interpreted differently. the TMVB. However, this climate trend is far from regular, showing
The TMVB lacustrine basins that have been the most intensively alternations between higher humidity/pluviosity and cooler pre-
studied include Chapala-Zacoalco, Jal., Zacapu, Cuitzeo, Patzcuaro, vailing temperatures, as reflected in higher lake levels and expan-
Mich., Texcoco, Chalco, Mexico, Hueyatlaco, Pue. and Central Tlax- sion of forests between 17 ka and 11 ka (Lozano-Garcı́a and Ortega-
cala (Foreman, 1955; Clisby and Sears, 1955; Sears and Clisby, 1955; Guerrero, 1997; Caballero-Miranda et al., 1999, 2001, 2002),
Bradbury, 1971, 1989; Ohngemach, 1973; Meyer, 1973; Ohngemach, preceded by a climate regime of lesser humidity and warmer
1977; Brown, 1984; González-Quintero, 1986; Straka and temperature between 20 ka and 17 ka. The glacial maximum effect

Fig. 10. Mexico’s chief Pleistocene palynofloral localities set on a Recent natural vegetation template (adapted from Rzedowski and Reyna-Trujillo, 1990; Challenger, 1998). Chief
sources: Foreman, 1955; Clisby and Sears, 1955; Sears and Clisby, 1955; Ohngemach, 1973, 1977; Bradbury, 1971, 1989; Meyer, 1973; Brown, 1984; González-Quintero, 1986; Straka
and Ohngemach, 1989; Metcalfe, 1992; Leyden et al., 1994, 1996; Lozano-Garcı́a and Ortega-Guerrero, 1994; Anderson and Van Devender, 1995; Lozano-Garcı́a and Ortega-Guerrero,
1997; Lozano-Garcı́a and Xelhuantzi-López, 1997; Ortega-Ramı́rez et al., 1998; Caballero-Miranda et al., 1999; Canul-Montañez, 2008; Martı́nez-Hernández, E., unpublished data.
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
is estimated to have occurred about 18 ka. A similar story can be
deduced from Chalco, Mex. (Lozano-Garcı́a and Ortega-Guerrero,
1994, 1997), and Zacapu, Mich. (Lozano-Garcı́a and Xelhuantzi-
López, 1997). The diatoms from the lake sequence of Tlaxcala,
although not specifically studied for detecting climate change
(Dı́az-Lozano, 1917; Rico et al., 1995, 1998), are consistent with
these findings. Although the lake records of the TMVB show
frequent volcanic activity, which indicates that at times, at least
locally, temperature and humidity were influenced by such activity.
Both terrestrial and lake biotas would have been strongly and
adversely affected by ash flow and ash fall emplacements, so
interpreting diminished abundance and/or diversity of the pollen
and diatom records must discriminate between climatic-induced
and volcanic-induced impacts (Lozano-Garcı́a and Ortega-Guer-
rero, 1997; Lozano-Garcı́a and Xelhuantzi-López, 1997; Martin-del
Pozo et al., 1997; Caballero-Miranda et al., 1999; Caballero-Miranda
et al., 2002).
The Fresnal and Babı́cora lake studies (Ortega-Ramı́rez et al.,
1998, 2004) from Chih. also record a moister and cooler interval
from 26 to 14 ka that is congruent with the results from the
TMVB lake sites studied. Bradbury’s (1997) conclusion about
a cooler and moister climate regime during the Wisconsinan in
regions lying west of 95 W, including the TMVB, CeP CH-CO, and
NW in Mexico, as well as Arizona and Texas in the United States,
is proving to be right. He related this fact to the pattern of global
atmospheric circulation driven by glaciation, which quite
unevenly and differently from present-day conditions transferred
moisture from the oceans to the continents, at least during this
time period.
3.2.2. Pleistocene environmental indicators (b), the paleosol record:
a review
Paleosol studies in Mexico were pioneered by Bryan (1948) in
the TMVB. Subsequent research chiefly focused on detecting
reliable markers to work out the Mexican Basin stratigraphy
(Arellano, 1953), or to understand sediment source and transport
(Heine and Shonhals, 1973). This delayed development of major
efforts on paleosols as environmental indicators chiefly until the
last decade (Cervantes-Borja et al., 1997; Sedov et al., 2001, 2008;
Gama-Castro et al., 2004, 2005; Ortega-Guerrero et al., 2004;
Solleiro-Rebolledo et al., 2003, 2004, 2006; McClung et al., 2005;
Jasso-Castañeda et al., 2006). Paleosols are dependable paleo-
climate indicators, because of their ability to record as soil prop-
erties or features, the effect of the particular set of conditions
under which a given paleosol is developed (Johnson and Watson,
1987; Birkeland, 1992). Such a property set is the soil climate
‘‘memory’’ (Targulian and Sokolova, 1996). In order to obtain
readily repeatable results decoding such memory, the field and
laboratory methodology, as well as the taxonomy used, are those
of USDA (1988, 1996).
The study of Mexico’s paleosols is hampered by the limited
number of specialists, and by the lack of precise knowledge on
their actual spatial distribution. The information is scattered in
numerous bibliographic and cartographic works mainly related to
geology, geomorphology, topography and morphometrics. A
major source is the INEGI (2006) Soil Chart, which indirectly
discriminates exposed or relict paleosols. The detection and
cartography of buried paleosols requires available maps of diverse
kinds, and particularly the extensive use of satellite imagery,
digitized air photos, as well as sophisticated computer equipment
and software.
3.2.2.1. Mexico’s Late Pleistocene paleosol record as an environmental
indicator. Fig. 11, a plot of the main paleosol localities on a mor-
photectonic template, is largely derived from INEGI (2006) and
79
from direct analysis by J. Gama of satellite imagery kindly furnished
by this institution with the Nearctic Region/Neotropical Domain
boundary also marked. The contrasting paleosol types, Luvisol and
Acrisol, are discriminated. In order to facilitate comparing the
environmental significance of the paleosol record, including the
vegetation it supported, the paleosol sites were also plotted on
a Recent natural vegetation template (Fig. 12).
Luvisols are most common in the TMVB and in the SMOc, largely
developed on silicic pyroclastics (volcanic ash, tuffs and lapilli tuffs,
surge deposits, lahars), less so on mafic to intermediate flows, and
even occur on alluvial deposits derived from mafic rock material.
Weathering occurs in a climate regime of humid to subhumid
conditions and of moderate temperature. The best developed
Luvisols seem to have been generated during the last glacial
maximum, circa 18–20 ka (Solleiro-Rebolledo et al., 1995, 2003,
2006; Cabadas-Báez, 2007). An actualistic analogy suggests the
following stages of plant succession. At an early stage, lichens,
ferns, shrubs and Gramineae made the vegetation cover; later
(juvenile stage) it was replaced by a pine forest; during the mature
stage, oak forest and grass land became associated with the pine
forest; finally, once Luvisols were fully developed, pine forest and
pine–oak forest became dominant. Phytoliths found in the luvisols
seem to bear out this interpretation (Sedov et al., 2003).
Standing in strong contrast, acrisols developed on heteroge-
neous material, and weathering took place under hot and humid
conditions, i.e., a tropical climate regime. The vegetation cover was
a tropical rain forest. Acrisol profiles disclose a high degree of
weathering, and morphogenetic development indicative of long-
term landscape/climatic stability (periods necessarily >10 ka).
Numerical dating of Acrisols in Mexico is still lacking, but associ-
ated and intrinsic soil evidence from the SMS probably place them
in the Middle Pleistocene. However, a cautious proposal is that the
acrisols formed between 25 and 50 ka (Figs. 11 and 12).
The term Acrisol in Mexico also refers to the ‘‘red soils’’ developed
on the YPL (Isphording, 1974; Bautista et al., 2003). Although the
substratum is calcareous, some red soils show Luvisol characteristics
(Sedov et al., 2003), although the significance of this observation is
not yet clear. Also regarded as acrisols are the paleosols of eastern-
most Chiapas, close to the Guatemala boundary, which seem to be
older and have developed under conditions of greater temperature
and humidity than those leading to ‘‘true’’ acrisols: because of this,
they are dubbed Ferralsols (Gama-Castro, 1996).
3.2.3. Pleistocene environmental indicators (c), the mammal record:
a review
The preceding review (Section 3.1) shows that there are only
three Pleistocene pre-Rancholabrean faunas (Fig. 7): La Goleta,
Mich., El Cedazo, Ags., and El Golfo, Son. Strata at La Goleta locality
include an unconformity, below which pre-Pleistocene Blancan III
mammals occur (Carranza-Castañeda, 2006). Above, the assem-
blage consists of undoubtedly Rancholabrean mammals coming
from the upper strata, and probably latest Blancan–Irvingtonian
ones from below those. However, given that most fossil mammals
were collected in various sites before the unconformity was
recognized, and without adequate stratigraphic control, the avail-
able collection is mixed (Arellano and Azcón, 1949; Repenning,
1962 (and R. Tedford and R. Shultz, unpublished data, quoted
therein); Miller and Carranza-Castañeda, 1984). Under these
circumstances, this fauna merits no further discussion.
Similarly, the El Cedazo fauna includes Rancholabrean and
earlier mammals. Most fossils were collected some 50 years ago
(Mooser, 1958), from different localities, without adequate
stratigraphic control, and hence the resulting collection is mixed.
Later, Montellano-Ballesteros (1992) revisited the area, and
recognized three informal stratigraphic units, each of which
80 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Fig. 11. Mexico’s chief Pleistocene paleosol sites set on the morphotectonic provinces template pictured in Fig. 1. Time frame adapted from Bell et al. (2004). Chief source: INEGI
(2006).

Fig. 12. Mexico’s chief Pleistocene paleosol sites set on a Recent natural vegetation template (adapted from Rzedowski and Reyna-Trujillo, 1990; Challenger, 1998). Chief source:
INEGI (2006).
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
contains different taxa. The lower unit, with Ailuralagus sp. and
Holmesina sp., resembles material from an Irvingtonian site in
Florida, thus allowing this level to be dated as Irvingtonian.
However, this assignment was disputed by Bell et al. (2004), who
contended that El Cedazo assemblage is best regarded as Latest
Blancan, that the presence of Bison indicates that part of the
assemblage is Rancholabrean, and that inadequate understanding
of the time relationships of the individual taxa makes unclear the
discrimination of an Irvingtonian chronofauna at El Cedazo.
Again, under these circumstances, this fauna is not further
considered here. This leaves El Golfo l.f., Irvingtonian of Sonora,
NW MP, as Mexico’s only undisputed pre-Rancholabrean Pleis-
tocene fauna.
The composition of the El Golfo fauna is provided in Table 5. It
includes seven orders, 21 families, 28 genera, and at least 29
species, of which only 12 are named. Based on the taxonomic
diversity, the amount of mammalian biomass would have required
a much higher productivity than that afforded by the present-day
(xeric shrub-grassland) vegetal community, and would also have
needed a much moister environment. However, the available
information does not permit quantification of climate parameters,
and hence the following environmental reconstruction is inferred
only from the mammalian fauna (Shaw, 1981; Shaw and McDonald,
1987). The presence of large herbivores, both grazers and mixed-
feeders including antilocaprids, bovids, and camelids, and probos-
cideans suggests a subtropical, seasonal savanna, located close to
a forested area, where the xenarthrans would have thrived. The
presence of castorids and hydrochoerids indicate the existence of
permanent bodies of water (perhaps a shallow lake, or a river pond)
in the area or nearby. Such a setting could have been developed in
the Early–Middle Pleistocene Colorado River Delta, where the El
Golfo area lies.
The Rancholabrean Pleistocene chronofauna is Mexico’s largest
(Fig. 6), based on many localities unevenly distributed across the
country (Figs. 5, 7, 8). Most of these faunas lack isotopic or paleo-
magnetic calibration, and thus dating of most local faunas is strictly
biochronological, which only allows a broad positioning within this
time span (roughly between 200–150 ka and 10 ka (Bell et al., 2004).
However, even with this chronologic restriction, most major faunas
from the TMVB, SMOr, CeP, and YPL are undoubtedly Rancholabrean.
Those few faunas with an independent means of assigned age belong
to this category (Arroyo-Cabrales et al., 2002, 2007a,b).
In order to facilitate environmental comparison/discussion of the
Rancholabrean local faunas, analysis is restricted to the major ones,
and those listed in Fig. 8 are regarded as of Late Rancholabrean age
(about 40–10 ka). This approach permits coherent examination of
the country’s Pleistocene mammal record, dealing with high rank
taxonomic categories (orders and families) rather than with local
faunas and/or low rank groups (i.e., genera and species), which
would require more space than is available in this study. Where
needed, specific references to local faunas and/or lower ranked taxa
are made. This procedure enhances detection of patterns (Graham,
1986). Additionally, the major local faunas were plotted on a present-
day natural vegetation template (Fig. 13) to ease comparison of
whether the Pleistocene mammal species sampled there would be
congruent with the biome or ecological community defined by the
natural vegetation now living in the area. Among faunas located in
and north of the TMVB, this quite frequently is not the case, as is
discussed below.
The Early Rancholabrean record is restricted to the Térapa l.f.,
Son., NW MP, which as discussed earlier, is the oldest known
Rancholabrean fauna of this country (Mead et al., 2006; Carranza-
Castañeda and Roldán-Quintana, 2007). Its mammalian faunal
composition, accompanying biota and sedimentary evidence indi-
cates the fauna lived on a semitropical savanna with a localized
81
marsh environment that supported an aquatic biota. The presence
of Crocodylus C. acutus in far inland Sonora, which today is
restricted to western and southern Pacific Mexico, is best explained
by the existence of estuarine/coastal northward migration from the
south, and immigration inland following the Rı́o Moctezuma to
Térapa, during a much more humid time interval than present. This
northward and inland dispersal was perhaps made possible by
climatic conditions during the glacial maximum 445–430 ka
(Winograd et al., 1992, 1997; Mead et al., 2006). The presence of
Tapirus sp., now living in tropical Mexico (eastern Tabasco–
southern Campeche and adjacent Chiapas), lends additional
support to this contention. This taxon is also known from the Late
Blancan and/or Irvingtonian of Florida (McKenna and Bell, 1997;
Bell et al., 2004). Currently, there are no Middle Rancholabrean
faunas known from Mexico.
The Pleistocene Rancholabrean mammal record of Mexico is
quite extensive (Table 12; Figs. 5, 8, 13). The table is a taxonomic
summary of local faunas arranged by morphotectonic provinces. It
is designed to ease comparison among faunas, and to provide an
indication of the geographic spread of particular taxa. In addition,
the figures plot the major fossiliferous localities on the morpho-
tectonic and the Recent natural vegetation templates, to collate past
environmental conditions (inferred from the environmental indi-
cators in each locality) with current ones, as reflected by the
vegetation occurring in the sites where the localities lie.
Didelphimorphia and Xenarthra. The didelphid Rancholabrean
record (Tables 2, 12) provides little environmental information.
Didelphis, which is known from the TMVB, currently lives in tropical
to temperate forested areas, and today is found in this MP. All of the
other didelphids known from fossils are found only in the YPL, and
today now occur in the tropics, the type of habitat found in
southern YPL at present.
The Rancholabrean xenarthran record is moderately large
(Tables 6 and 12), and widespread (Fig. 12). Living pilosans in
Mexico are limited to myrmecophagids, a strictly tropical group
(Ceballos and Oliva, 2005). Megalonychids browsed in forest and
woodland in temperate North America (Kurtén and Anderson,
1980; Anderson, 1989). In Mexico, they occur chiefly within and
north of the TMVB. Nothrotheres are regarded as savanna/wood-
land dwellers, but at least the Shasta ground sloth (Nothrotheriops
shastensis) during the latest Rancholabrean thrived on more xeric
vegetation, in Arizona, under a mild climate (Phillips, 1984), and
perhaps other megatheriids were able to do the same. Mylodonts
fed on shrubs and grass (Anderson, 1989), and their record in the
TMVB (Table 12, Chapala-Zacoalco, Jal. and the Basin of Mexico)
together with palynological record of grasses and shrubs (Table 11,
Fig. 9) is consistent with this interpretation. Summing up, it
appears that the record of pilosan xenarthrans within and north of
the TMVB indicates a more humid, and perhaps warmer climate
regime in this vast territory (circa 1.38  106 km2) than at present,
at least during the Late Rancholabrean. The glyptodonts and
pampatheres are regarded as cold-intolerant, but their humidity
preferences are still unknown (Gillette and Ray, 1981; McDonald,
personal communication, August, 2009), so their record might be
largely congruent with that of loricates. However, extant dasypo-
dids are able to stand more xeric settings (Honacki et al., 1982;
Wilson and Reeder, 2005).
Insectivora s.s. (¼ Lipotyphla; ¼ Soricomorpha). The Rancho-
labrean record of insectivores is limited (Tables 2 and 12), chiefly
coming from the SMOr, with minor contributions from the CH-CO
and YPL (one each). At present, insectivorans live in forested and
savanna areas, as well as in prairie areas with rivers or standing
water bodies nearby (Hall, 1981; Honacki et al., 1982; Wilson and
Reeder, 2005). Most fossil records lie within in the respective extant
species’ range (Ceballos and Oliva, 2005). Two exceptions are
82 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Fig. 13. Mexico’s chief Pleistocene terrestrial mammal-bearing localities set on a Recent natural vegetation template (adapted from Rzedowski and Reyna-Trujillo, 1990; Challenger,
1998); the Nearctic Region/Neotropical Realm boundary is also plotted. Main sources: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and Arroyo-
Cabrales et al. (2007a,b). For further information see Table 3 and Appendix.
Cryptotis parva and Notiosorex sp., with records outside their
current distribution, the latter barely. The environmental signifi-
cance of this record is unclear.
Chiroptera. The Rancholabrean chiropteran record is extensive
(Tables 2 and 12), but is strongly biased toward cave deposits (Table
7). Hence, most comes from the YPL and the SMOr, and thus its
environmental representation is limited. The molossids, phyllos-
tomids and vespertilionids, which form most of the record, largely
include tropical and cosmopolitan species (Ceballos and Oliva,
2005). Seven of the 44 fossil species records lie within areas with
a different biome/vegetation type than that in which the corre-
sponding extant species are known to thrive (Fig. 13). The tropical
species Balantiopteryx io (Emballonuridae) and Desmodus spp.
(Phyllostomidae; Ceballos and Oliva, 2005) are recorded as fossils
in the SMOr and the TMVB (Table 7), now chiefly covered by pine
forest. Macrotus californicus (Phyllostomidae) currently occupies
xeric habitat (Ceballos and Oliva, 2005), is also recorded from the
SMOr. Eptesicus brasiliensis (Vespertilionidae) at present lives in
savanna and mesophyllus forest settings (Ceballos and Oliva, 2005),
but it is recorded from the largely tropical YPL. Lasionycteris noc-
tivagans (Vespertilionidae), currently known to occupy riparian to
temperate deciduous forest communities (Ceballos and Oliva,
2005), is recorded from the SMOr (Table 7), south of their present
range. The molossids Eumops underwoodi and Nyctinomops auri-
spinosus are recorded as fossils east of their extant range. The N.
aurispinosus record also indicates different habitat conditions than
the current ones (Ceballos and Oliva, 2005).
Carnivora. Mexico’s Rancholabrean Carnivore record is large (at
least 37 species, Tables 2, 8 and 12). However, only in six instances
do the localities of the fossil forms lie outside the respective species
current range and/or preferred community (Fig. 13). These include
the felids Herpailurus yagouaroundi and Panthera onca, the
mustelids Lontra longicaudis, Mephitis mephitis, and Spilogale
putorius, and the ursid Ursus americanus. The first currently is
a tropical-mesophyllus-thorn forest dweller in the SMOc western
slope, SMS southern GCP, CHI, and YPL (Ceballos and Oliva, 2005),
and hence its record from the SMOr at San Josecito indicates
warmer conditions, at least locally.
The extant P. onca largely lives in the same communities utilized
by H. yagouaroundi, which is a tropical-mesophyllus-thorn forest
dweller from the NW, western SMOc, westernmost TMVB, western
SMS, CHI, YPL GCP (Ceballos and Oliva, 2005). Therefore, its fossil
records in the CH-CO, SMOr and TMVB also indicate a warmer
moister climate regime then presently exists in these areas. Living L.
longicauda occupies a riparian habitat adjacent to tropical/
subtropical, mesophyllus and/or thorn forests in the SMOc, western
TMVB, SMS, CHI, YPL and southern GCP (Ceballos and Oliva, 2005).
Its fossil record from Tequixquiac, Mex. in central TMVB suggests
warmer and moister conditions in this region during the Rancho-
labrean. The extant M. mephitis is a grassland, thorn and oak forest
dweller in the NW, northern SMOc and CH-CO, as well as north-
ernmost GCP. Therefore, its fossil record in the SMOr at San Josecito
lies somewhat south of its current range, although not as far from
the habitat where it thrives.
Living S. putorius resides in shrub, thorn and pine-oak forest
from the northernmost GCP and adjacent SMOr (Ceballos and Oliva,
2005). During the Rancholabrean it was more widely distributed
and is found in the CH-CO, SMOr and YPL (Tables 8 and 12; Fig. 13).
There are at least three alternative interpretations for this wider
distribution: (a) assuming strict contemporaneity of the fossil
records involved, more xeric conditions extensively prevailed in
Mexico at that time, at least in the north (CH-CO) and east (YPL),
although this contention is not supported by other taxa; (b) the
extensively prevailing xeric conditions in Mexico during the
Table 12
Selected main Pleistocene local mammal faunas of Mexico. Chief sources: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and Arroyo-Cabrales
et al. (2007a,b); for additional sources see text. y, Extinct taxon everywhere in the world; , species extinct in Mexico, but still extant outside this country; B, species extinct in
the morphotectonic province(s) bearing the fossil locality(ies), but extant elsewhere in Mexico.

(continued on next page)

84 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Table 12 (continued)
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 85

For symbols, see Table 2. Morphotectonic provinces: NW, Northwestern Plains and Sierras; CH-CO, Chihuahuan-Coahuilan Plateaus and Ranges; SMOr, Sierra Madre Oriental;
CeP, Central Plateau; GCP, Gulf Coast Plain; TMVB, Trans-Mexican Volcanic Belt; SMS, Sierra Madre del Sur; YPL, Yucatan Platform. Local faunas: A, Terapa, Son; B, La Brisca,
Son; C, El Golfo, Son; D, Cuatro Ciénegas, Coah; E, Cueva Jiménez, Chih; F, San Josecito, NL; G, La Presita, SLP; H, El Cedral, SLP; I, Minas, NL; J, Mina San Antonio, SLP; K, El Cedazo,
Ags; L, Mixtequilla, Ver; M, Chapala-Zacoalco, Jal; N, Tequixquiac, Mex; O, Tlapacoya, Mex; P, Valsequillo, Pue; Q, Hueyatlaco, Pue; R, San Agustı́n, Oax; S, Gruta de Loltún, Yuc;
T, Actún Spukil, Yuc.
86 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
Rancholabrean, is an artificial climate pattern, resulting from
inaccurate fossil dating; or (c) S. putorius might have become less
competitive, being now restricted to a fraction of its former range.
Clearly, additional study is needed to resolve this issue.
Extant U. americanus is a pine, pine–oak, shrub forest and
occasional grass land dweller in the SMOcc, CH-CO and northern
SMOr (Hall, 1981; Ceballos and Oliva, 2005). Its Rancholabrean
record from the TMVB at Valsequillo, Pue. may indicate cooler
conditions during the Rancholabrean, at least locally. Other carni-
vore taxa also merit consideration. Canis dirus was probably
ecologically like C. lupus (Kurtén and Anderson, 1980). Arctodus and
Tremarctos were somewhat similar in habitat requirements to
Ursus, but with different food habits (Kurtén and Anderson, 1980;
Arroyo-Cabrales et al., 2010). Smilodon on the other hand, has no
Recent counterpart. The record of Cuon alpinus, a canid now living
in central Asia (Honacki et al., 1982; Wilson and Reeder, 2005) from
the SMOr at San Josecito, N.L., needs further corroboration.
Rodentia. Rodent species seem to be more narrowly adapted to
particular environmental conditions than other mammal species,
thus making them important environmental indicators. Given that
nearly a third of the recorded Pleistocene mammal species belong
to this order, and that most are still extant (hence amenable to
actualistic comparisons), they could afford a reliable environmental
reconstruction of the Pleistocene climate and subsequent change
for the entire country. However, this theoretical expectation is
actually hampered by several factors, such as (a) the strong time
and space biases of the record; (b) the taphonomic component of its
genesis (largely raptor predation); and (c) the multiplicity of
temperature/moisture and related conditions that may produce
largely similar habitats, as discussed elsewhere (Sections 3.2.1 and
3.2.2). With this in mind, the Pleistocene rodent record as an
environmental indicator is reviewed.
Castoridae and Hydrochoeridae (Table 9; Fig. 13) suggest
wetter environments in the NW and TMVB during part of the
Pleistocene. However, the former family is found at mid and high
latitudes in temperate North America, and may indicate relatively
cooler conditions in the NW (Sonora) during the Irvingtonian
than at present, whereas the tropical hydrochoerids require
warmer conditions in the TMVB at least locally in the Rancho-
labrean than at present. Members of the Cuniculidae and Dasy-
proctidae were restricted to tropical areas during the
Rancholabrean. On the other hand, the erethizontid record
(Table 9; Fig. 15) includes Erethizon, currently confined to
temperate habitats. Coendou at present occupies warm and
humid habitats, thus suggesting climatic amelioration, at least
along the paths that allow their displacement northward from
southern tropical latitudes.
Sciurids, although able to thrive in a wide variety of habitats/
biomes (McLaughlin, 1984), include taxa with strong habitat pref-
erences, e.g. Cynomys is best adapted to grassland, Sciurus prefers
forest and woodland, and Spermophilus dwells best in sparsely
vegetated rocky settings (Ceballos and Oliva, 2005). Given that
a combination of humidity and temperature conditions could
accommodate these biomes and habitats, the climate signal affor-
ded by the sciurids is inconclusive. Geomyids (Table 9; Fig. 13)
usually thrive in grasslands and open country (Ceballos and Oliva,
2005).
Heteromyids (Table 9; Fig. 13) are eurytopic at the family level,
and include species that live in arid/semiarid environments (e.g.
Chaetodipus and Dipodomys), and others that prefer deciduous
tropical forest, like Liomys and Heteromys (Ceballos and Oliva,
2005). The presence of Liomys in the Irvingtonian El Golfo l.f.
indicates moister conditions at that time.
Murids also include temperate species able to thrive in cool and
moist settings, such as forest, woodland and even grassland, e.g. the
arvicolines such as Microtus. Their Rancholabrean records from
currently warmer, less humid areas (Table 9; Fig. 13) indicate
a post-Rancholabrean climate change. On the other hand, genera
like Hodomys, Ototylomys and even Oryzomys thrive in warmer/
moister settings of tropical to subtropical environments (Hall, 1981;
Ceballos and Oliva, 2005). Their presence in places now having
different environmental conditions (Table 9; Fig. 13) also supports
a post-Rancholabrean climate change.
Neotoma is best adapted to xeric (dry, temperate to cool) areas
(Hall, 1981; Ceballos and Oliva, 2005), whereas Peromyscus, Rei-
throdontomys, and Sigmodon are eurytopic at the generic level,
including species able to live in a wide variety of habitats/biomes
(Hall, 1981; Ceballos and Oliva, 2005). Therefore, their Rancholab-
rean records (Table 9) must be environmentally evaluated on
a species by species basis.
Lagomorpha. Leporids are present in Mexico’s modern and
Rancholabrean fauna but are scarce as fossils (Table 2). The fossil
record is biased toward cave deposits, and thus most of the fossil
record comes primarily from the SMOr and the YPL (Table 12;
Fig. 13). Extant species of Lepus mainly live in xeric communities
(Ceballos and Oliva, 2005), but it also occurs in the surrounding
woodlands. Thus, its fossil record from the SMOr, covered at
present by pine and pine–oak forest, is not unexpected. The extinct
genus Aztlanolagus also from the SMOr, probably ecologically was
closer to Lepus than to Sylvilagus (Arroyo-Cabrales et al., 2004). The
latter genus lives in a wide biomic spectrum from tropical to pine
and pine-oak forest, to xeric communities (Ceballos and Oliva,
2005). The fossil record of Sylvilagus includes the PBC, SMOr, TMVB
and the YPL. Only the record of S. cunicularius in the SMOr lies
outside the extant species range and the expected community.
Perissodactyla. Extant equids roam widely in savanna and
grassland communities from tropical to temperate climate regimes
(Honacki et al., 1982; Wilson and Reeder, 2005), and their fossil
counterparts are considered to have similar preferences (Kurtén
and Anderson, 1980). As discussed elsewhere, Rancholabrean
equids are recorded from all MPs, but are more common in the CeP,
SMOr and TMVB (Table 12; Fig. 13). Their abundance suggests that,
at least during the Late Rancholabrean, probably these provinces
had a greater organic productivity than at present, which sup-
ported large populations of these big herbivores. In turn, an overall
greater humidity across the country during this time span, as well
as a somewhat different climate regime might have made such
productivity possible; yet no quantitative studies to test these
contentions have been undertaken. However, in the recent past the
North American Great Plains supported large populations of feral
horses and bison, which indicates that even under current climate
conditions, this territory’s organic productivity suffices for this.
Extant tapirs are tropical mammals living associated with
permanent water bodies (Hall and Kelson, 1959; Honacki et al.,
1982; Wilson and Reeder, 2005), and so are interpreted as Ran-
cholabrean. Thus, their record from the NW, SMOr and TMVB is
noteworthy, indicating for that time span a rather different envi-
ronmental setting (Arroyo-Cabrales et al., 2010), at least locally.
However, in Mexico and Central America extant tapirs could also
thrive at high elevation (w2500 masl; Ceballos and Oliva, 2005) in
mesophyllous and cloud forests, withstanding relatively cool
conditions, and thus the Rancholabrean record of tapirs in the
SMOr and the TMVB is not so unexpected. The Tapirus sp. record
from Térapa, Son. (NW MP) has already been discussed.
Artiodactyla. The Rancholabrean artiodactyl record of Mexico is
large (Tables 2, 10 and 12) and widespread (Figs. 5, 8 and 13).
However, only seven of the 34 recorded species remain extant, thus
limiting its actualistic assessment. Currently, Antilocapra americana
thrives in xeric communities in the PBC, NW, CH-CO and SMOr
(Ceballos and Oliva, 2005), as a fossil it is known from the CH-CO. A
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
similar ecological setting might be hypothesized for the other
Rancholabrean antilocaprids (Tables 2 and 12; Fig. 13). None are
recorded south of the TMVB. Extant Bison bison grazes on grass-
lands and adjacent temperate forests in a reduced area in the CH-
CO MP (Ceballos and Oliva, 2005). During the Rancholabrean, Bison
roamed nearly all over Mexico (QMMDB, Arroyo-Cabrales et al.,
2002). At present, Ovis canadensis largely shares the same envi-
ronment and area as Bison, and as fossils it is recorded from the
same MP. The other bovid records lie in or north of the TMVB
(Tables 2 and 12), which might suggest that they were ecologically
closer to Ovis than to Bison.
Cervids fared better than other artiodactyls, and three of the six
Rancholabrean species survive to the present: Mazama americana,
Odocoileus virginianus, and O. hemionus. The first two are eurytopic,
but preferred forested areas. M. americana’s fossil record in Mexico
lies outside its current range. O. hemionus is a dry community
dweller, including some xeric conditions, and its fossil record partly
lies outside its present-day range. Extant Cervus elaphus dwells in
pine forest and moorlands from British Columbia to southern Cal-
ifornia, Arizona, and New Mexico (Hall, 1981; Honacki et al., 1982;
Wilson and Reeder, 2005). Its fossil record from the CH-CO may
indicate prevailing cooler and moister conditions there during Late
Rancholabrean time. The other fossil cervids (Table 12; Fig. 13) are
recorded from the CeP, SMOr and TMVB, but only the extinct genus
Navahoceros is present in the SMS. Their ecological requirements
were probably similar to that of Mazama and O. virginianus. Tayassu
tajacu is eurytopic (Ceballos and Oliva, 2005). However, the extinct
genus Platygonus, the other tayassuid, is unrecorded south of the
TMVB, so it probably was a pine-thorn-forest-savanna dweller.
Camelids are now extinct in Mexico, Central and North America
(Table 12) (QMMDB, Arroyo-Cabrales et al., 2002). They probably
were savanna-grassland inhabitants, perhaps able to thrive as well in
more xeric conditions. Their Rancholabrean record from Mexico is
quite extensive north of the TMVB, whereas it is very meager south
of it. Hemiauchenia sp. from the YPL (Table 12; Fig. 13), seems to be
the only southern record (QMMDB, Arroyo-Cabrales et al., 2002).
Proboscidea. Mexico’s Rancholabrean proboscidean record is
abundant (Table 12) and widespread (Fig. 13). Mammuthus is known
from all MPs but the YPL, and also in Central America, and thus it
must have been fairly eurytropic. In and north of the TMVB, it is
commonly associated with Camelops hesternus, Bison, Equus, Pan-
thera atrox and Arctodus, whereas in the SMS, CHI and southern GCP
it is associated with Megalonyx, Eremotherium, Mixotoxodon and
other tropical taxa (Arroyo-Cabrales et al., 2010). Gomphotheriids
are chiefly recorded from localities in and south of the TMVB (Table
12). They were mixed-feeders living in savanna and associated
woodland habitat (Alberdi and Corona-M., 2005; Alberdi et al.,
2009). Cuvieronius remains have been recorded in Ixtapa, Chiapas
from fluvial strata underlying a K–Ar 400 ka ignimbrite sheet
(Ferrusquı́a-Villafranca, 1996, and unpublished data), thus adding to
the meager Mexican Irvingtonian mammal record. Mammut occurs
in the CeP, SMOr, TMVB, and YPL (Table 12) (Arroyo-Cabrales et al.,
2007a,b). It is largely regarded as a pine and pine-oak forest dweller,
feeding on herbs and shrubs (Arroyo-Cabrales et al., 2007a,b), but
also it was able to thrive in grassland/savanna and tropical forest, as
records of the genus from the CeP and YPL seem to indicate.
Notoungulata and ?Litopterna. The record of these two groups
in Mexico is scarce (Table 2), and limited to the TMVB and GCP. Both
groups are regarded as large tropical herbivores (Scott, 1937; Pat-
terson and Pascual, 1963, 1968, 1972) that arrived in Mexico from
South America via Central America, where the former are already
known late in the Pleistocene (Woodburne, 1969; MacFadden,
2005). The TMVB notoungulate record indicates either that at least
locally, this MP had tropical patches during the Rancholabrean,
because of their complex topography, or that notoungulates could
87
thrive in subtropical to temperate savannas. Diagnostic records of
Litopterns are known only from South America (Patterson and
Pascual, 1963, 1968, 1972), and their presence in Mexico is still
doubtful (Ferrusquı́a-Villafranca and Malvido-Arriaga, 2006).
4. Discussion
4.1. Geochronological assessment and overall composition of
Mexico’s Pleistocene mammal record
Section 3.1 examined the local faunas within a wide time and
space perspective in order to disclose broader patterns about the
chronologic spread and geographic distribution of Mexico’s Pleis-
tocene mammals. Fig. 14 depicts the biogeochronologically
assessed range of selected local faunas, which are arranged by
morphotectonic province provenance. The expressed local faunas
ranges are approximate, and were estimated combining the bio-
chronologic information afforded by the mammal taxa with rele-
vant geologic information, and with radioisotopic and
paleomagnetic dates when available. The emerging pattern is
consistent with the observed strong time and space bias of the
mammal record, whereby the Late Rancholabrean is the best rep-
resented, whereas pre-Rancholabrean faunas and isolated occur-
rences are few and far apart: TMVB (La Goleta l.f., Mich., Blancan,
?Irvingtonian-Rancholabrean), CeP (El Cedazo l.f., Late Blancan-
Rancholabrean), NW (El Golfo, Son., Irvingtonian), CHI (Chiapa de
Corzo and Ixtapa, Chis., Irvingtonian Cuvieronius remains coming
from strata underlying w400 ka K–Ar dated ignimbrite sheets).
The Rancholabrean local faunas comprise most of Mexico’s
Pleistocene mammal record. Figs. 14 and 15 depict the estimated
geochronologic range of selected major local faunas. Comple-
mentary 14C for the late Rancholabrean and paleomagnetic dating
for the Irvingtonian and Blancan provide an age constraint for
some faunas, largely from the TMVB and a single fauna from the
SMOr (San Josecito, N.L., e.g. Arroyo-Cabrales et al., 1995), but
most faunas are based on biochronology. Both figures also show
that the geographic distribution of mammal bearing localities
across the country is quite uneven, with most concentrated in
a few provinces (TMVB and SMOr), although some from other
provinces (e.g. CeP and YPL) show a large diversity, the latter
particularly in micromammals (see Sections 3.1. and 3.2.4). Nearly
as important as what is recorded is what remains unrecorded,
namely the enormous information gap resulting from the scarcity
of pre-Late Rancholabrean mammal records across the country,
which severely hampers generation of a truly dependable model
of climate change for Mexico’s Quaternary, a record whose
understanding is a vital issue to contemporaneous civilization.
On the other hand, the review of Mexico’s Pleistocene mammal
record does allow some general remarks about several aspects of its
structure. Taxonomically, Table 2 shows that the record includes 13
orders (if Litopterna are present), 43 families, 146 genera and 278
species. However, of these Orders, Chiroptera (50 sp.ecies, 17.85%),
Carnivora (38 sp.ecies, 13.57%), Rodentia (97 sp.ecies, 34.64%) and
Artiodactyla (35 species, 12.50%) dominate and represent 78.57% of
all the species, whereas the other nine orders combined only
contribute 21.43%. The number of Pleistocene species is about 60%
of the Recent ones (about 479 species) (Hall, 1981; Ramı́rez-Pulido
et al., 1996; Villa-Ramı́rez and Cervantes, 2003; Ceballos and Oliva,
2005; Ceballos et al., 2005b), but includes greater diversity at the
ordinal and family levels.
With respect to survivorship of Pleistocene species in the
Recent, 195 (71.16%) are currently extant, and 79 (28.83%) became
extinct (Table 2); the corresponding ratio is 2.46 to 1.0. At higher
hierarchical levels, extinction affected 26 genera (17.80% of all,
a little over 40% of this figure in the Artiodactyla alone), six families
88 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

Fig. 14. Estimated biogeochronologic position of Mexico’s chief Pleistocene terrestrial mammal faunas, which are arranged according to the morphotectonic provinces scheme
pictured in Fig. 1. Time frame adapted from Bell et al. (2004_. Numbers within province columns refer to the localities/local faunas plotted in Fig. 4. Bars in NW and CHI columns
denote geochronologic range of 40A–39A and K–Ar dating respectively. Local faunas 11, 21, 26 and 34 have 14C data. For sources and further information, see text. Abbreviations (from
left to right): LMA, North American Land Mammal Ages; BCP, Baja California Peninsula; NW, Northwestern Plains and Sierras; SMOc, Sierra Madre Occidental; CH-CO, Chihuahuan-
Coahuilan Plateaus and Ranges; GCP, Gulf Coastal Plain; SMOr, Sierra Madre Oriental; CeP, Central Plateau; TMVB, Trans-Mexican Volcanic Belt; SMS, Sierra Madre del Sur; CHI,
Sierra Madre de Chiapas; YPL, Yucatan Platform. Local faunas. BCP, 4, Comundú l.f., BCN; 41, El Carrizal l.f., BCS; NW, 3, El Golfo l.f., Son.; 5, La Brisca l.f., Son.; 6, Térapa l.f., Son. CH-CO,
7, Cueva Jiménez l.f., Chih.; 8, Cuatro Ciénegas l.f., Coah. GCP, 19, Mixtequilla l.f., Ver. SMOr, 11, El Cedral l.f., SLP.; 12, Minas l.f., NL.; 14, San Josecito l.f., NL; 15, La Presita l.f., SLP; 16,
Mina San Antonio l.f., SLP. CeP, 2, El Cedazo l.f., Ags. (a, seemingly Blancan portion; b, seemingly Irvingtonian portion; c, Rancholabrean portion); 21, Arperos l.f. Gto. TMVB, 1, La
Goleta l.f., Mich. (a, seemingly Irvingtonian portion; b, Rancholabrean portion); 22, Chapala-Zacoalco l.f., Jal.; 25, Tequixquiac l.f., Mex.; 26, Tlapacoya l.f., Mex.; 27, Hueyatlaco l.f.,
Pue.; 28, Valsequillo l.f., Pue. SMS, 33, San Agustı́n l.f., Oax. CHI, 96, Ixtapa l.f., Chis (a, seemingly Irvingtonian portion; b, Rancholabrean portion); 97, Chiapa de Corzo l.f. Chis (a,
seemingly Irvingtonian portion; b, Rancholabrean portion); 98, Villa Corzo l.f., Chis. YPL, 34, Gruta de Loltún l.f., Yuc.; 35, Actún Spukil l.f., Yuc.

(4 xenarthran and 2 proboscidean ones), and two orders
(Notoungulata and Litopterna). Thus extinction was important, very
uneven across taxonomic boundaries, but not overwhelming,
diminishing the diversity by 20% at the order and family levels.
Regarding body mass, the Pleistocene mammals could be
grouped in three broad categories as follows: 187 (64.24%) species
are microbaric, i.e., less than 10 kg, 27 (9.85%) are mesobaric
(between 10 and 100 kg, and 59 (21.89%) are megabaric, i.e., more
than 100 kg. Thus, microbaric taxa outnumber megabaric ones,
3.16:1.0, and also the mesobaric ones 6.9:1.0. Microbaric to meso-
baric and mesobaric to megabaric ratios are 6.9:1 and 0.45:1,
respectively. In Recent mammals (479 sp.ecies), given the paucity of
meso- and megabaric species with respect to the microbaric ones,
the ratios are different: microbaric to megabaric is 31.14:1.0, and
microbaric to mesobaric is 15.03:1.0 (species data from Cervantes
and Oliva, 2005).
In reference to dietary habits, Mexico’s Pleistocene mammals can
be arranged in four, highly uneven categories: 179 (65.32%) species
are herbivorous, 66 (24.08%) species are carnivorous, 25 (9.12%)
species are omnivorous, and only four (1.45) are hematophagous.
The ratio between herbivores and carnivores is 2.68. In the current
fauna consisting of 479 species, 306 (63.88%) are herbivorous, 148
(30.89%) are carnivorous, 23 (4.80%) are omnivorous, and 2 (0.41%)
are hematophagous (species data from Cervantes and Oliva, 2005).
The ratio of herbivore species to carnivores is 2.06:1.0, which is not
that different from the Pleistocene one, as might be expected given
the large compositional difference between these two chro-
nofaunas. In turn, this suggests that the overall ecological (predator/
prey) equilibrium of the mammal fauna was somehow maintained,
at least from the Late Pleistocene to the Recent, although the
mechanisms remain to be worked out.
4.2. Mexico’s Pleistocene environment and faunal response to
climate change: a review by MT Provinces
Section 3.2 also shows that individual species and groups (e.g.,
communities, biomes, faunas, floras, biotas) have definitive envi-
ronmental preferences, which are of course related to the particular
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 89

Fig. 15. Estimated biogeochronologic position of Mexico’s chief Late Pleistocene terrestrial mammal faunas, which are arranged according to the morphotectonic provinces scheme
pictured in Fig. 1. Time frame adapted from Bell et al. (2004). Numbers within province columns refer to the localities/local faunas plotted in Fig. 4. Local faunas 11, 21, 26 and 34
have 14C data. Discontinuous line indicates local fauna’s possible temporal extent. For sources and further information, see text. Abbreviations (from left to right): LMA, North
American Land Mammal Ages; BCP, Baja California Peninsula; NW, Northwestern Plains and Sierras; SMOc, Sierra Madre Occidental; CH-CO, Chihuahuan-Coahuilan Plateaus and
Ranges; GCP, Gulf Coastal Plain; SMOr, Sierra Madre Oriental; CeP, Central Plateau; TMVB, Trans-Mexican Volcanic Belt; SMS, Sierra Madre del Sur; CHI, Sierra Madre de Chiapas;
YPL, Yucatan Platform. Local faunas. BCP, 4, Comundú l.f., BCN; 41, El Carrizal l.f., BCS; NW, 5, La Brisca l.f., Son.; 6, Térapa l.f., Son. CH-CO, 7, Cueva Jiménez l.f., Chih.; 8, Cuatro
Ciénegas l.f., Coah. GCP, 19, Mixtequilla l.f., Ver. SMOr, 11, El Cedral l.f., SLP; 12, Minas l.f., NL; 14, San Josecito l.f., NL; 15, La Presita l.f., SLP; 16, Mina San Antonio l.f., SLP. CeP, 2, El
Cedazo l.f., Ags. (c, Rancholabrean portion); 21, Arperos l.f. Gto. TMVB, 1, La Goleta l.f., Mich. (b, Rancholabrean portion); 22, Chapala-Zacoalco l.f., Jal.; 25, Tequixquiac l.f., Mex.; 26,
Tlapacoya l.f., Mex.; 27, Hueyatlaco l.f., Pue.; 28, Valsequillo l.f., Pue. SMS, 33, San Agustı́n l.f., Oax. CHI, 96, Ixtapa l.f., Chis (b, Rancholabrean portion); 98, Villa Corzo l.f., Chis. YPL, 34,
Gruta de Loltún l.f., Yuc.; 35, Actún Spukil l.f., Yuc.

environment where they lived. However, to retrieve the environ-
mental information stored in the fossil record is not a straightfor-
ward process. Consider the temperature and moisture climate
factors and their evaluation through an actualistic comparison
between the past (Pleistocene) and present (Recent) distribution
area of an extant species recorded in the Pleistocene, which indi-
cates different past and present environmental (climatic) condi-
tions. The difference may involve at least these variables: (a)
moisture/pluviosity (dry-humid spectrum), (b) temperature (cold-
hot thermal spectrum), (c–f) any of the four possible moisture–
temperature combinations. Further, considering that climate is
a function of solar energy input, latitude and altitude influence
prevailing climate conditions in a particular place.
The biotic response to these and other environmental condi-
tions occurs at two levels. (a) At the species level, a species even-
tually becomes adapted to a set of environmental conditions,
occupying a certain habitat. However, should such conditions
change, the species may expand/contract its distribution area in
precise lockstep fashion with habitat expansion/contraction, or it
may migrate or move as the environmental conditions displace; or
it may become extinct. (b) The aggregate results of such individual
species responses give a biota its physiognomy or aspect, which
then reflects the accumulated environmental response of all the
species involved through time. It follows that the physiognomy of
a biota occupying a particular area, e.g. a morphotectonic province
may change in time, as the review of the Pleistocene mammal
record discloses, but the available information only permits broad
discrimination of a few individual environmental factors involved
in this complex environment/biota interplay. Therefore, inferring
Pleistocene climate from bioevents alone (i.e., the fossil record)
and actualistic comparisons thereof, is rather limited, allowing at
best to trace broad qualitative patterns. Further, the available
information utilizing only the fossil and environmental record
reviewed here discloses not only biases, but points to important
taxonomic and geographic gaps that must be filled before a more
complete model of the Quaternary climate and climate evolution
for the whole country could be prepared.
The review of environmental indicators presented indicates that:
(1) All data sets show strong time and space biases. In all three sets
the time bias favors the Late Pleistocene. However, the space
bias, although broadly coinciding (Figs. 5, 8, 9 and 11), displays
some differences. The palynofloral set is far denser in the TMVB,
with much less coverage in the other morphotectonic provinces
(Fig. 9). The paleosol set is also denser in the TMVB, and less so in
the SMOc and the YPL, whereas the other provinces have very
few localities if any (Fig. 11). The fossil mammal set has more
localities in the TMVB, but the SMOr, CH-CO and YPL have
yielded important local faunas (Figs. 5 and 8; Table 12).
(2) Each set includes data that may and have been interpreted
differently (e.g. pollen, soil and mammal records at the TMVB).
For instance, in the fossil mammal set, the environmental
sensitivity of the taxa varies a great deal, so that mammals from
the same locality may indicate different conditions (e.g. YPL at
Loltún: temperate ursid associated with tropical dasyproctid
and xenarthran records).
(3) Not surprisingly, the environmental information obtained from
one set only broadly coincides with that of the other. Further-
more, the precise localities of different data sets only broadly
coincide, even in well-sampled provinces like the TMVB,
because they resulted from independent, non-coordinated
research projects; the Chihuahuan Desert case is a notable
exception, but different interpretations of related data sets are
not uncommon.
(4) The available Pleistocene mammal record, because of its strong
time, space and taxonomic biases across the country, does not
90 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

allow mapping the actual biogeographic dynamics of a species
throughout this epoch, such as migration, range expansion/
contraction or disjunction (either vicariant or not). The paucity
of detailed studies aimed at describing and mapping the
paleoclimate regime across the country for the entire Pleisto-
cene further hinders advancement of knowledge on mamma-
lian biogeographic dynamics.
(5) For most morphotectonic provinces, the available information
is restricted to the Late Rancholabrean (approximately not
older than 40 ka). Even within this interval, only broad quali-
tative climate change trends could be delineated (Fig. 16).
Fig. 16 is the result of the parsimonious integration of data from
the different sets for each province, using the available time
framework (chiefly biochronologic), and resolving inconsistencies,
favoring the most congruent or less conflicting interpretation.
Rather than identifying particular paleoclimates (such as tropical,
steppe, temperate), which might not be warranted under these
circumstances, a more cautious approach was followed, discrimi-
nating (qualitatively) separately temperature and humidity/plu-
viosity trends, using as a reference the Recent average climate
conditions for each province (Garcı́a, 1990; Hernández, 1990; Vidal-
Zepeda, 1990a,b). Pleistocene climate change trends and fluctua-
tions are long known (pioneering work of Agassiz, 1840). The
Wisconsinan fluctuating climate pattern and timing was broadly
worked out in the mid-20th century (Flint, 1947), and it was
recognized for Mexico shortly after (Foreman, 1955; Clisby and
Sears, 1955; Sears and Clisby, 1955; Heine, 1984). However, the
environmental information generated afterwards, although quite
abundant and enlightening (e.g. Habib et al., 1970), still does not
allow establishment of detailed climate changes for long time spans
across the country, but only portrayal of trends.
Nonetheless, it is hoped that portraying such trends may help to
better understand the biotic response to climate change. In turn,
this response at various ‘‘ecologic’’ levels (species, community, and/
or biome), may explain the expansion/reduction of current
biogeographic range of particular mammal species with respect to
its Pleistocene one, as well as its present-day permanency or
displacement/dispersal from its Pleistocene range. The other
response outcome is extinction. Eventually, the combined species
response is reflected in the biotic physiognomy of a region, and
ultimately may lead to biogeographic provincialism (Table 12). Brief
discussions of the faunal response in each province and/or province
groups follow, stressing the most significant mammal taxa.
4.2.1. Northern provinces: PBC, NW, SMOc and CH-CO
The NW and CH-CO have some major local faunas, palynofloras
and paleosol sites. The first includes Mexico’s only proven Irvingto-
nian fauna, El Golfo l.f., as well as the earliest Rancholabrean fauna,
Térapa l.f., both in Sonora. They disclose a more humid climate regime
then, which allowed the presence of a subtropical biota, at least
during the Irvingtonian and Early Rancholabrean. The SMOc paleosol
record (Fig. 11), and the CH-CO paleo-vegetation record (Betancourt
et al., 1990) indicate a cooler and moister climate regime for the latest
Rancholabrean circa 25–10 ka), which made possible a biotic diver-
sity unparallelled anywhere at present in Mexico. This meant that
within short distances, mammals with diverse ecological require-
ments coexisted (Table 12; Bell et al., 2004).
The biota became strongly decimated and compositionally
modified as the climate regime changed in the Holocene. With
increasing aridity, xerophilous species largely replaced the former
inhabitants. Many became extinct, mostly the meso- to megabaric
taxa, including most xenarthrans (Pampatherium mexicanum,
Glyptotherium cylindricum), the carnivorans (Canis dirus, Panthera
atrox, Chasmaporthetes johnstoni, Tremarctos floridanus), the rodent
Neochoerus sp., the lagomorph Aztlanolagus agilis, the Equidae, the
artiodactyls (e.g., Capromeryx minor, C. mexicana, Stockoceros sp.,
Camelops hesternus, Hemiauchenia blancoensis, Platygonus sp.), and
the probocideans. Others depending on their climatic tolerances,
experienced a major range contraction either to the north (such as

Fig. 16. Late Pleistocene probable climate trends in Mexico’s morphotectonic provinces (sensu Ferrusquı́a-Villafranca, 1993, 1998). Time frame adapted from Bell et al. (2004). The
reviewed palynologic, paleosol, lake sediment and mammal records were used to assess the trends. For sources and further information, see text. Abbreviations (from left to right):
LMA, North American Land Mammal Ages; BCP, Baja California Peninsula; NW, Northwestern Plains and Sierras; SMOc, Sierra Madre Occidental; CH-CO, Chihuahuan-Coahuilan
Plateaus and Ranges; GCP, Gulf Coastal Plain; SMOr, Sierra Madre Oriental; CeP, Central Plateau; TMVB, Trans-Mexican Volcanic Belt; SMS, Sierra Madre del Sur; CHI, Sierra Madre de
Chiapas; YPL, Yucatan Platform.
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
the temperate rodents Castor and Marmota, and the carnivoran
Canis rufus) or south (the tropical capybara Hydrochoerus), whereby
they currently live outside Mexico. In other instances, some taxa
became extirpated from their Rancholabrean morphotectonic
province, but persisted to the present in the other one (e.g., the
xenarthran Myrmecophaga tridactyla, the rodent Hodomys sp., and
the perissodactyl Tapirus sp., currently living in the tropics).
The palynological and vegetation fossil record, particularly for the
Chihuahuan Desert (which includes most of the CH-CO), the climate
change and biotic response for the last 40 ka were complex, so that
the vegetation distribution changed over shorter time spans, both
latitudinally and altitudinally (Van Devender, 1990a,b). Certainly the
same situation is expected over a larger territory (i.e., the whole
country), and over a much longer time interval (i.e., the entire
Pleistocene, Bell et al., 2004), although the available information only
allows the recognition of broad trends of climate change.
4.2.2. Central and Eastern Provinces: CeP, SMOr and GCP
The palynological and paleosol record of these provinces is very
scarce (Figs. 9 and 11), whereas several major mammal local faunas
come from the CeP and SMOr (Figs. 5 and 8; Table 12). The CeP
yielded a biochronologically mixed mammal assemblage including
taxa of apparently Late Blancan to Rancholabrean age (Montellano-
Ballesteros, 1992; Bell et al., 2004), whose time relations are not
well understood. At any rate, the Late Blancan record is very scarce,
meriting no further consideration, since the better part of the
assemblage, El Cedazo l.f., is Rancholabrean.
By and large, what was inferred from the northern provinces is also
applicable here. The SMOr and CeP faunas include tropical/subtrop-
ical, temperate and even xeric species, which suggest a variety of
ecological settings not possible in the present climate regime. This in
turn indicates different climate conditions during the Rancholabrean,
seemingly ultimately driven by advances/retreats of the Laurentide
Glacier as it responded to global climate pattern changes. Climate
thermal (warm/cold) and humidity/pluviosity (moist/dry) oscilla-
tions occurred, not necessarily univocally related (i.e. warm with
either moist or dry conditions, and cold with moist or dry conditions),
which were also affected by altitude and latitude. The net result was
a complex shifting of species distribution within relatively short time
intervals during the Pleistocene, expressed as biogeographic range
expansion/contraction, or displacement and colonization of ‘‘new’’
habitats, or of course extinction. A latest Rancholabrean interval
(w25–10 ka) of cooler and moister climate in the northern provinces
seems to be also recognizable, based on the palynological and
mammal records. The complex climatic and biotic history preceding
this climate regime in the (Late) Rancholabrean resulted in a mammal
fauna much more diverse than exists at present. By this interval’s end,
the climate had become in general warmer and drier.
Meso- to megabaric mammals fared the worst (Table 12).
Many became extinct, including the xenarthrans (Glyptotherium
mexicanum, Holmesina septentrionalis, Megalonyx jeffersoni,
Nothrotheriops mexicanum, Nothrotheriops shastensis, and Para-
mylodon harlani); the chiropterans (Desmodus stocki); the carni-
vorans (Canis cedazoensis, C. dirus, Panthera atrox, Smilodon fatalis,
Bassariscus ticuli, Arctodus pristinus, A. simus, Tremarctos floridanus);
the Equidae; the artiodactyls (e.g., Capromeryx mexicana, Stock-
oceros conklingi, Tetrameryx mooseri, T. shuleri, T. tacubayensis, Bison
alaskensis, B. antiquus, B. priscus, Camelops hesternus, C. traviswhitei,
Euceratherium collinum, Hemiauchenia macrocephala, H. vera, Nav-
ahoceros fricki, Odocoileus halli, Oreamnos harringtoni, and Platygo-
nus ticuli); and the proboscideans.
Other species reduced their biogeographic range, currently
living outside Mexico (e.g., the carnivore Cuon alpinus, and the
rodent Marmota flaviventris). Others became extinct in their Ran-
cholabrean morphotectonic province, but survived elsewhere
91
within Mexico including the rodent Hodomys alleni, which
currently thrives in the tropics. Other rodents such as Erethizon
dorsatum, Microtus pennsylvanicus, Neotoma albigula and Synapt-
omys cooperi now live in the northern provinces.
4.2.3. Trans-Mexican Volcanic Belt
This province has a greater number of palynofloral loca1ities
(Figs. 9 and 10), paleosol sites (Figs. 11 and 12), and fossil mammal
localities (Figs. 5, 7 and 8) than any of the other provinces. However,
the sampling is strongly biased toward the latest Rancholabrean
(w40–10 ka). The Goleta area, near Morelia, Mich., has yielded
a largely pre-Pleistocene (Blancan III) mammal assemblage, La
Goleta l.f. (discussed elsewhere) from a fine-grained, tuffaceous
fluvio-lacustrine sequence. The strata include an unconformity
(Carranza-Castañeda, 2006), above which Pleistocene taxa such as
Equus and Mammuthus occur, but whether they are Early or later
Rancholabrean is unknown.
For the most part, the palynofloral-paleosol-mammal record is
concordant, allowing the recognition of the 22–10 ka moister/
colder climate regime episode mentioned above. The enormous
diversity of the TMVB (Late) Rancholabrean mammal fauna, far
greater than the current one, includes taxa of different ecological
requirements living close by (e.g. Neochoerus, Platygonus, Odocoi-
leus hemionus and Bison) and indicates a correspondingly diverse
setting, where different vegetation types were either intermixed or
changed over short distances. Pleistocene climatic fluctuations
acting on a complexly rugged territory, such as the TMVB’s could
have promoted the large vegetation diversity implicitly required by
the diverse mammal record. The pollen record seems to support
this contention, as discussed elsewhere.
The climate regime changed by the Early Holocene, becoming
warmer and drier. As in other provinces, meso- to megabaric
species became extinct, including the xenarthrans (Holmesina
septentrionalis, Pampatherium mexicanum, Glyptotherium cylin-
dricum, G. floridanum, G. mexicanum, Megalonyx jeffersoni, M.
wheatleyi, Eremotherium laurillardi, Nothrotheriops mexicanum, N.
shastensis, and Paramylodon harlani); the chiropteran (Desmodus
stocki); the carnivorans (Canis dirus, Panthera atrox, Smilodon fatalis,
S. gracilis, Arctodus pristinus, and A. simus); the rodents (Neochoerus
aesopi, Baiomys intermedius, Microtus meadensis, Neotoma anomala,
N. magnodonta, N. tlapacoyana, and Peromyscus maldonadoi); the
Equidae; the artiodactyls (Capromeryx mexicana, Stockoceros con-
klingi, Tetrameryx shuleri, Bison alaskensis, B. antiquus, B. latifrons,
Euceratherium collinum, Camelops hesternus, C. mexicanus, C. mini-
dokae, Eschatius conidens, Hemiauchenia macrocephala, Procamelops
minimus, Odocoileus halli, Platygonus compressus, and P. ticuli); the
probocideans, and the possible litoptern.
Other species became extinct in Mexico, but currently live
outside, such as the carnivoran Canis rufus, now living in temperate
North America. Others disappeared from their Rancholabrean
morphotectonic province, but survived elsewhere within Mexico,
such as the rodents (e.g., Microtus californicus, is now restricted to
northern BCP; Neotoma albigula, now thriving in NW; and N. phenax
currently living in southern NW); and the perissodactyl Tapirus
bairdii is now restricted to the tropics.
4.2.4. Southern provinces: SMS, CHI, GCP (southern part), and YPL
In this region lie some important palynofloral localities (Figs. 9
and 10), paleosol sites (Figs. 11 and 12) and major fossil mammal
localities (Figs. 5 and 8; Table 12). However, the sampling is strongly
biased toward the Late Pleistocene (Rancholabrean s.l.). As dis-
cussed elsewhere, the paleosol (Acrisol) record points to an
equable, largely tropical climate regime in the region for the last
40–50 ka. The palynological record, particularly that of Tehuacán,
Pue., northeastern SMS just south of the TMVB (Canul-Montañez,
92 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
2008), indicates moister and cooler conditions in the latest Pleis-
tocene, so that the now prevailing xeric vegetation is a Recent
phenomenon.
The mammal record (Table 12 and preceding environmental/
taxonomic review, Sections 3.1 and 3.2.4) mainly includes tropical
taxa, although cosmopolitan and temperate taxa are less frequent,
but not uncommon. These include Ursus americanus, Eumops per-
otis, and Spilogale putorius. This indicates that during the Rancho-
labrean, the climate regime, although warm and moist in general,
probably was punctuated at least locally, by less warm (¼ cooler)
and/or drier episodes, which allowed temperate taxa to south-
wardly expand their range, or to disjunctly occupy southern habi-
tats. This region’s geomorphic complexity, as well as limited
sampling, both in space and time (Figs. 5, 7 and 8), does not permit
the recognition of the specific pattern of changes, but only delin-
eates broad climate change trends (Fig. 16).
Nevertheless, as in the other provinces, important mammal
composition changes took place at the end of the Rancholabrean.
Medium and large size mammal taxa became extinct including the
carnivoran Canis dirus, the perissodactyl Equus conversidens, the
artiodactyls Navahoceros fricki, and Hemiauchenia sp., as well as
the proboscidean Mammut americanum. Other largely cosmopol-
itan or temperate taxa survided further north in temperate habitats
(e.g. the chiropteran Eptesicus fuscus, and Eumops underwoodi, the
carnivoran Ursus americanus; and the artiodactyl Bison bison),
perhaps returning to their primary range or a close approximation
(see preceding discussion).
Summing up, the combined results of individual species range
modifications affected and changed the biota’s/fauna’s physiog-
nomy and taxonomic makeup, so that the present one is quite
different from the (Late) Pleistocene one. However, the available
fossil record does not portray this major biogeographic shifting of
species. The analysis of disjunct and of demonstrably relict species
may be an alternative to this end. Biogeographic aspects of the
mammal record are treated in the last section of this paper.
4.3. Provinciality and other biogeographic aspects
4.3.1. Provinciality
As biogeography developed, major regions based on the distri-
bution of different groups were recognized and variously named
(e.g. birds, Sclater, 1858; mammals, Wallace, 1876; angiosperms,
Engler and Diles, 1936). In North America, the boundary between
the tropical and temperate biota, bearing ‘‘regions’’ was placed
across Mexico, by the TMVB. The boundary’s conception varies from
moderately sharp, as held by the majority (Wallace, 1876; Good,
1963; Darlington, 1966; Brown and Lomolino, 1998; Zunino and
Zullini, 2003), to arbitrary because, supposedly, the whole country’s
territory can be considered a broad transition zone (Cox and Moore,
2005; Woodburne et al., 2006). Further subdivision of Mexico into
biotic/biogeographic provinces was pioneered by Smith (1940),
who based his scheme on the distribution of Sceloporus spp. (an
iguanid). Since then, other schemes based on different groups were
proposed (Goldman and Moore, 1946; Hall and Kelson, 1959 (‘‘life
zones’’); Leopold, 1977; Casas-Andreu and Reyna-Trujillo, 1990;
Ramı́rez-Pulido and Castro-Campillo, 1990; Flores-Villela, 1993).
The Mexican Commission for the Knowledge and Use of Biodiver-
sity (CONABIO in Spanish) organized a 2-day workshop in 1997 to
draw a consensual biotic scheme for the whole country, which
would serve as the basis for establishing a nation-wide conserva-
tion-oriented ecoterritorial policy. The diverse schemes were dis-
cussed and evaluated, but the methodologies and defining criteria
of each were so different, that an integrated scheme acceptable to
all was not agreed upon and the situation persists today.
Given that species/faunas adjust in response to environmental
change, and that in Mexico it could be divided into broad areas
based on the morphotectonic provinces, each possessing particular
conditions (e.g. similar geologic makeup, geomorphic structure,
and to a certain degree soil and climate as well), these geologically
based provinces become important factors in shaping the distri-
bution of species and hence the faunal composition of the province.
As such they can be used in combination with distribution patterns
of different taxonomic groups, to establish a broader provincial
pattern, with more meaningful biotic and physiogeographic
boundaries. Such a biotic province scheme (Ferrusquı́a-Villafranca,
1990) agrees closely with those based on terrestrial vertebrates
(e.g. amphibians and reptiles, Casas-Andreu and Reyna-Trujillo,
1990; mammals, Ramı́rez-Pulido and Castro-Campillo, 1990), and
partially with that of angiosperms (Rzedowski and Reyna-Trujillo,
1990). The latter is expected given the dependence of plants on the
soils derived from the geology.
In order to test if this biotic scheme may hold for the Pleisto-
cene, the major fossil mammal localities were plotted on this
template (Fig. 17). Some broader patterns could be detected, such
as the Nearctic Region/Neotropical Domain boundary, and some
provinces are also recognizable, like the Neovolcanian (TMVB),
Coahuilan (CH-CO), Yucatanian (YPL) and Chiapasian (CHI) among
others. In the transitional provinces, localities bear a mixture of
tropical mammals (e.g. Veracruzian, northern GCP, and Sinaloan,
NW), temperate mammals (e.g., Oaxacan, SMS). The Chihuahuan
and Altiplanian Provinces, where currently xeric conditions prevail
appear wide in comparison with the Sierras Madres Occidentalian
and Orientalian Provinces, probably reflecting the upward forest
retreat, as a result of the warmer and drier climate regime estab-
lished in post-Rancholabrean time. However, to formally propose
a biotic province scheme for the Pleistocene would require a more
representative record time- and space-wise than the currently
available one, as well as an in-depth biogeographic analysis, not
possible here. Thus, in lieu of that, and as a working biogeographic/
faunistic province model or scheme for Mexico’s Late Rancholab-
rean, the proposed scheme (Fig. 17), seems to adequately reflect
the complex Pleistocene environmental/biotic history. Further-
more, this scheme could be regarded as a partial improvement
upon broader faunal provincial schemes proposed for the Late
Cenozoic of North America, such as those of Martin and Neuer
(1978), Graham (1979), Martin and Hoffman (1987), Fejfar and
Repenning (1992), Repenning et al. (1995), FAUNMAP Working
Group (1996), and Bell (2000). In all of them, Mexico is considered
a single broad province, the Mexican one (see also Mead et al.,
2006), which may include Florida (Fejfar and Repenning, 1992), or
exclude it (Bell, 2000). This variable definition of the province
perhaps reflects the scarcity of published relevant information,
rather than faunal uniformity. The review and discussion of Mex-
ico’s Pleistocene mammal record glaringly demonstrates the
complexity and heterogeneity of the faunal makeup and distribu-
tion, thus making untenable the simplistic interpretation of an
overall faunistic uniformity of this country that has been implied in
these schemes.
The role of Southern Mexico–Central America related to
mammalian cladogenesis during the Tertiary and subsequent
dispersal has been considered by many researchers (Scott, 1937;
Simpson, 1947, 1951; Patterson and Pascual, 1968; Ferrusquı́a-Vil-
lafranca, 1978; Webb and Perrigo, 1984; Webb and Rancy, 1996)
among others), and has recently been further elaborated by
Woodburne et al. (2006), who dubbed it the Central America
Province, without explicitly defining either its time or space limits,
although some Pleistocene mammal taxa are discussed. It appears
that at least for the Pleistocene, the Central American Province
included Mexico south of the TMVB, i.e., it largely corresponds to
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 93

Fig. 17. Mexico’s chief Pleistocene terrestrial mammal-bearing localities set on a biotic provinces template (adapted from Ferrusquı́a-Villafranca, 1990), the Nearctic Region/
Neotropical Realm boundary is highlighted. This template may serve too as a working Pleistocene biogeographic province scheme for this country. Main fossil mammal record
sources: Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales et al., 2002), and Arroyo-Cabrales et al. (2007a,b). For further information see Table 3 and Appendix.

the Middle American Region of the biogeographic scheme here
proposed (Fig. 17).
4.3.2. Other biogeographic aspects
4.3.2.1. Some major post-Pleistocene mammal distribution change-
s. Given that extinction and/or extirpation have a biogeographic
component, and may lead to the overall faunistic physiognomy
change (Table 2), the following are some additional remarks.
Extinction at the end of the Pleistocene included the Orders
Notoungulata and ?Litopterna, along with six families (four
xenarthrans and two proboscideans), 29 genera (seven xenar-
thrans, three carnivorans, one rodent, two lagomorphs, ten
artiodactyls, and four proboscideans), and included 77 species
(one didelphimorph, ten xenarthrans, two chiropterans, 11
carnivores, 13 rodents, four lagomorphs, 25 artiodactyls, and five
proboscideans). In terms of body mass, this includes 10.6%
microbaric species, 29.6% mesobaric species and 91.5% megabaric
species.
This event not only severely changed the distribution and faunal
physiognomy of mammals in Mexico, at the ordinal and family
levels, it also reiterates the well-established observation that meso-
to megabaric mammals fared the worst, not only in Mexico, but in
North America as a whole, as well as South America and Europe
(Osborn, 1911; Scott, 1937; Romer, 1945, 1966), and treated at
greater depth more recently in the context of Quaternary extinc-
tions (e.g., Martin and Klein, 1989).
Another well known fact, but important enough to be
mentioned here is that other suprageneric taxa managed to survive
hundreds or even thousands of kilometers away from Mexico. They
include the Proboscidea, now living in Africa and India; the Hyae-
nidae, currently extant largely in Africa and Asia; Hydrochoeridae,
still thriving in South America; the Equidae, living at present in
Africa and Asia; and the Camelidae, now extant in Asia and South
America (Honacki et al., 1982; Wilson and Reeder, 2005).
In addition to the above there are numerous records of taxa now
extirpated from some morphotectonic provinces where they are
represented by fossils but surviving elsewhere in Mexico. This
includes two murids (Rodentia), one dasypodid and one myrme-
cophagid (Xenarthra); one antrozoine vespertilionid, one embal-
lonurid, two molossids, and one phyllostomid (Chiroptera); two
cebids (Primates); one erethizontid, one heteromyid, eight murids,
and three sciurids (Rodentia); and one tapirid (Perissodactyla).
Finally, 27 species are extirpated in the morphotectonic province(s)
where they are represented by fossils, but persist at present else-
where in Mexico. Two species of rodent are now extinct in Mexico
but extant outside the country.
These biogeographic modifications also influenced the mammal
fauna physiognomy of the morphotectonic provinces involved, and
provide evidence of important environmental changes, at least at
provincial level. Most species are associated with temperate zones
and their distribution rarely exceeds five latitude degrees (Figs. 5, 8
and 13; Table 12). However, the environmental interpretation
derived from some fossil species may be at variance with that
inferred from other records (mammalian or of other kind), thus
making them hard to integrate into a conceptually coherent whole.
The following example illustrates this fact.
The myrmecophagid fossil-extant species records stand at
least 15 latitude and longitude apart, thus reflecting a biogeo-
graphic displacement of equivalent magnitude. Could it be
explained by climate change alone? If so, the change involved
would have been of a geographic extension equal to the country’s
north–south and east–west length. A climate change of such
magnitude would have affected the biota nation-wide, and must
have left a tangible fossil record displaying approximately the
94 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
same pattern. The mammal record as a whole does not conform
to this pattern (Table 12), nor do the palynologic and paleosol
ones. The sedimentary lake record (Lozano-Garcı́a et al., 1993,
Ortega-Ramı́rez et al., 1998; Caballero-Miranda et al., 1999) does
not support this pattern either. A parsimonious integration of
such diverse and conflicting information would rule out a nation-
wide change in climate to explain the myrmecophagid example,
thus requiring a combination of other factors, such as oppor-
tunism, dispersal capabilities of this myrmecophagid, and the
former distribution of colonial insects (its primary food source)
as an alternative tentative explanation.
4.3.2.2. Possible Pleistocene mammal dispersal routes and their
bearing in understanding the current mammal fauna physi-
ognomy. The available fossil record (QMMDB, Arroyo-Cabrales et al.,
2002) does not permit mapping of the geographic path followed by
these 27 species from their Pleistocene range to the present-day
one, nor to trace changes in their distribution through time, but only
portrays broad patterns of biogeographic change. On a greater scale,
the current distribution of these and other species falls within the
long known complex, present-day Mexico’s biotic and biogeo-
graphic pattern, whereby the temperate and tropical biotas extend
far southward and northward respectively relative to the Tropic of
Cancer (Hall and Kelson, 1959; Miranda and Hernández-X., 1963;
Leopold, 1977; Rzedowski, 1993), which is the practical boundary
between these biotas (Good, 1963; Darlington, 1966; Neill, 1970;
Brown and Lomolino, 1998; Zunino and Zullini, 2003; Christo-
pherson, 2006). These extensions call for past routes that allowed
dispersal or other mechanisms that could explain the observed
current pattern of distribution. Unfortunately, neontologists have
made very little use if any of relevant paleontologic and geologic
information to delineate such routes. It has only been broadly
postulated that the lowlands around the Gulf of Mexico and the
Pacific Ocean served for northward advance of tropical species, and
that mountain ranges permitted southward motion of temperate
species. The TMVB has been regarded as a barrier to the wholesale
north/south movements of tropical and temperate taxa. In an effort
to go beyond this stage, Ferrusquı́a-Villafranca and Gonzı́lez-Guz-
mı́n (2005); Ferrusquóa-Villafranca et al. (2005) summarized the
Cenozoic geologic and biotic evolution of Northern Mexico,
concluding that the Sierras Madres Oriental and Occidental func-
tioned as plausible southward dispersal routes of temperate species,
and the GCP and NW served also as northward dispersal routes for
tropical/subtropical species. With broadly the same aim, Ferrus-
quı́a-Villafranca et al. (2007) also reviewed the geologic evolution of
the TMVB, highlighting its biogeographic role as a major boundary
between Mexico’s temperate and tropical biotas.
Briefly, by Middle Paleogene time no major east–west high
mountain range hindered north–south biotic exchange between
central and southern Mexico, thus this region (w18 to 20 N)
functioned as a corridor (the CM/SM corridor), and the biota most
probably showed physiognomic change (from temperate in the
north to tropical in the south) along temperature/humidity gradi-
ents related to latitude, and also to local geomorphic features of this
corridor. The relevant plant record supports this contention (Gra-
ham, 1993; Rzedowski, 1993; Magallon-Puebla and Cevallos-Ferriz,
1994; Weber and Cevallos-Ferriz, 1994; Velasco-de León et al.,
1998; Ramı́rez and Cevallos-Ferriz, 2000; Ramı́rez et al., 2000;
Calvillo-Canadell and Cevallos-Ferriz, 2002; Calvillo-Canadell,
2008). Later, the CM/SM corridor became reduced in width from
west to east (Urrutia-Fucugauchi and del Castillo, 1977; Demant,
1982; Ferrari et al., 1994a; Pardo and Suárez, 1995), giving way to
narrow, low-lying passes, as the nascent TMVB ranges developed
by Middle-Late Miocene time (Ferrari et al., 1994b, 2000).
Continued volcanic and tectonic activity further built up the TMVB,
encroaching on the passes, and hindering north/south biotic
exchange. Eventually, by Late Pliocene–Early Pleistocene time, the
exchange became possible only through the TMVB and the adjacent
SMS drainage network. The precise routes used by particular
species, and the timing of such events, remain to be worked out.
Probably, as the TMVB developed, some communities and/or
species became isolated, but dating their occurrence on the extant
species record alone is not possible.
Combining the available geologic, paleontologic and environ-
mental information reviewed here, a broad hypothesis of Mex-
ico’s Pleistocene mammalian biogeographic dynamics is possible
(Fig. 18). It is still preliminary, given that it will take considerable
time to reduce the deficiencies of the record and to promote and
implement the necessary long-term multidisciplinary studies. In
the model presented here, temperate and tropical participants are
differentiated. Considering that the geographic position of the
routes s.l. involved is significantly different (low lands: plains and
plateaus on one hand, versus high lands: mountainous territories
in the other), both conditions were distinguished. The biogeo-
graphic polarity was inferred from present-day geographic
distribution of taxa; yet given that during the Pleistocene, broad
climate zones shifted latitudinally southward and northward, any
particular corridor could have been traversed in either direction
as climate conditions and species convenience demanded it.
Bearing all this in mind, the following corridors are proposed
(Fig. 18):
A. Low land, largely tropical mammal, biogeographically north-
ward routes:
(a) Gulf Coastal Plain Corridor. Tropical and subtropical species
traversed this corridor. It may have been available at least
since the Late Blancan, as attested by numerous xenarthran
and hydrochoerid (Neochoerus) records in Florida (Morgan
and Hulbert, 1995; Bell et al., 2004). The taxonomic makeup
of local faunas 19, 20, 34, 35 and 36 (Fig. 5; Table 12), partly
lends credence to this route. The current distribution of
mammals in southern YPL (Hall, 1981; Ceballos and Oliva,
2005), and in Guatemala (Hall, 1981) lead us to the postulate
that mammals from each subregion independently dispersed
east and northward, and thus two separate branches of the
GCP corridor are inferred. Outside Mexico, the presence of
xenarthrans in Blancan and Pleistocene faunas in many sites
of the Continental Interior (Bell et al., 2004, Fig. 7.4 and
references therein), seems to require at least some other
route northward, stemming from the GCP corridor.
(b) Pacific Corridor. Tropical and subtropical species dispersed
along this route. The TMVB western end divides this route
into a southern segment and a northern segment, here
designated the Sonoran Branch (c1), which probably remained
active at least until Irvingtonian–Early Rancholabrean time,
as indicated by the number of tropical/subtropical species in
El Golfo l.f. (Shaw, 1981; Shaw and McDonald, 1987) and
Térapa l.f. (Mead et al., 2006). It is possible that some tropical/
subtropical mammals could have passed the Gulf of California
(by Tiburón Island?) and reached the BCP’s eastern end
(Californian Branch, b2). The taxonomic composition of local
faunas 4, 40, 41, and 43 (Tables 3 and 12; Fig. 5; Appendix)
partly lends support to this contention.
(c) Balsas Depression Corridor and associated minor northward
routes: This depression is a WNW–ESE trending geomor-
phic feature southwardly adjacent to the TMVB, drained by
the Rı́o Balsas. Its northern slope is traversed by numerous
N–S trending tributaries. The presence of tropical aquatic
species (e.g. Neochoerus sp. and Crocodylus sp.) in the Cha-
pala-Zacoalco l.f., and of other tropical/subtropical
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104 95

Fig. 18. Chief Mexico’s probable Pleistocene temperate and tropical mammal biogeographic dispersal routes set on the morphotectonic provinces template pictured in Fig. 1. Low
land, largely tropical mammal routes: A, Gulf Coastal Plain Corridor; B, Pacific Corridor including its Sonoran (B1) and Californian (B2) branches; C, Balsas Depression Corridor and
associated minor northward routes. High land, largely temperate mammal routes: D, Rocky Mountains Corridor; D1, Southern Rockies–Sierra Madre Occidental Branch; D2,
Southeastern Rockies-Sierra Madre Oriental Branch; E, Cascadian/Nevadan-Western Baja California Corridor; F, Trans-Mexican Volcanic Belt Corridor, ‘‘Biogeographic Mixer,’’ and
Speciation Region; G, Southeastern Mexico Corridor; H, Southern Sierra Madre del Sur Corridor. Low land, largely temperate mammal routes: I, Great Basin-Sonoran Corridor; J,
Great Plains-Western Chihuahua/Central Plateau Corridor. For sources and further information see text.

mammals including the giant sloth Eremotherium (Table 12)
in several TMVB local faunas indicates that their most likely
source was the Balsas Depression, and perhaps the northern
Balsas tributaries and associated slopes functioned as
northward dispersal routes. Probably the Zacoalco Graben,
as well as the NW–SE trending fault and fracture zones
within the TMVB (Ferrusquı́a-Villafranca et al., 2007, Fig. 7),
also served as northward dispersal routes for tropical
species. The taxonomic makeup of local faunas 30 and 85
(Table 3; Fig. 5; Appendix) partly provides support for the
existence of this route.
B. High land, largely temperate mammal, biogeographically
southward routes:
(d) Rocky Mountains Corridor. The Rocky Mountains Cordillera
south of the Colorado Plateau broadly joins the Sierra
Madre Occidental, thus furnishing a corridor branch here
named Southern Rockies-Sierra Madre Occidental Branch
(d1). The composition of local fauna 22 (Fig. 5 and Table 12)
partly supports this contention. East of the plateau,
a narrow range parallel to the Rio Grande Rift extends
southward to the Coahuilan plateaus, and through them to
the eastern sector of the Sierra Madre Oriental, forming
another corridor branch, here named Southeastern Rockies-
Sierra Madre Oriental Branch (d2). Numerous local faunas of
the SMOr, e.g. San Josecito, Mina de San Antonio, and El
Cedral (Fig. 5; Table 12) include temperate, highland species
of mammals, like Marmota flaviventris, Microtus pennsylva-
nicus, and Synaptomys cooperi.
(e) Cascadian/Nevadan-Western Baja California Corridor. This
corridor involves the southwestern ranges of North Amer-
ica, lying close to the Pacific Coast. Highland-prone species
living in the southwest probably could have used it.
(f) The Trans-Mexican Volcanic Belt Corridor, ‘‘Biogeographic
Mixer,’’ and Speciation Region. Both highland and lowland
species moving south encountered this east–west oriented
mountain range barrier, which effectively deterred south-
ward dispersal for most temperate, low land-prone
mammal species. Further southern dispersal was addi-
tionally hindered by the tropical climate regime and biomes
existing south of the TMVB. On the other hand, highland-
prone species could continue southward, perhaps through
two routes (see below). Possibly during the Pleistocene, the
TMVB diverse physical-geographic scenario and fluctuating
climate regime had resulted in frequent disjunct distribu-
tions of species, subsequent genetic isolation, and eventu-
ally to speciation. The available information on the fossil
and geologic records is insufficient to establish the precise
dating of such events, or to map specific routes. However,
the fact that numerous extant TMVB species show discon-
tinuous and/or highly endemic, isolated biogeographic
distribution areas, such as Romerolagus diazi, Zygogeomys
trichophus, and Neotomodon alstoni (Hall, 1981; Ceballos
and Oliva, 2005) lends credence to this contention. This
shows the major role played by the TMVB in shaping the
composition and physiognomy of Mexico’s mammal fauna
during the Quaternary. The highly diverse taxonomic
96 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
composition of the local faunas, e.g. 22, 23, 24, 25, 26, 27,
and 28, found in the TMVB (Table 12; Fig. 5) lends strong
support to this and the above interpretations.
(g) Southeastern Mexico Corridor. Temperate, high land-prone
species that reached the TMVB’s eastern part could continue
southward traversing along the maize of mountain ranges
present in the SMS and CHI (see below). There is not enough
information to detect specific routes. In some regions, like
northern Oaxaca–southern Puebla, the complex relief and
microclimate promotes habitat diversification, allowing the
coexistence of tropical and temperate species within short
distances (Rzedowski and Reyna-Trujillo, 1990; Ferrusquı́a-
Villafranca, 1990; Flores-Villela, 1991). This complex
biogeographic picture is also partly reflected in the fossil
record (Tables 6–10). It should be noted that the taxonomic
makeup of some local faunas, e.g. 27, 30, 32, 84, 85 and 93
(Tables 3 and 12; Fig. 5; Appendix), indicates a complex
biogeographic picture congruent with this interpretation.
(h) Southern Sierra Madre del Sur Corridor. This corridor is
currently restricted to the highest part of the SMS Pacific
Ranges, where pine and pine–oak forests occur (Rzedowsky
and Reyna-Trujillo, 1990), and some temperate mammals
(largely rodents, Ceballos and Oliva 2005) are present. So, it
seems that some species that reached the TMVB’s western
part, and continued southward following this route. The
meager fossil record, e.g. local faunas 32 and 37 (Fig. 5;
Appendix), as well as the QMMDB (Arroyo-Cabrales et al.,
2002), lends credence to this contention.
C. Low land, largely temperate mammal routes:
(i) Great Basin-Sonoran Corridor: The low land region laying
between the Rockies/Colorado Plateau and the Pacific
Ranges, probably functioned as a southward biogeographic
dispersal route for mammals living in southwestern North
America. The corridor narrows to the south, and because of
its rather dry climate regime, it might have functioned as
a filter rather than as a corridor. The taxonomic composition
of local faunas 3, 5, 6, and 44 (Tables 3 and 12; Fig. 5;
Appendix) partly lends credence to this route.
(j) Great Plains-Western Chihuahua/Central Plateau Corridor. The
vast plains of North America, narrow southward, and in
Mexico are bordered by the SMOc to the west and the
Coahuilan plateaus and ranges to east. Farther south, the
SMOr eastern sector, which lies at a much lower altitude in
the west, practically allows the Chihuahuan plateaus and
plains to reach the Central Plateau, which to the south
becomes bound by the TMVB. The climate is less moist than
the mountain ranges that border it. However, because of its
great N–S extension (w40 at least), moisture and temper-
ature vary along latitude-dependent gradients, so that in
Mexico at present, the climate regime is dry with strong
seasonal temperature changes, thus supporting largely
prairie and xeric communities. These gradients add a mild
filtering effect to this vast corridor. During the Pleistocene,
the advances/retreats of the Laurentide Glacier correlatively
caused concomitant displacements of the ‘‘climate belts,’’ to
the north in the United States (Ehlers and Gibbard, 2004),
however in Mexico, the available information does not allow
mapping of such displacements, which also involve other
components (e.g. atmospheric circulation, which in turn
influences moisture distribution/discharge), as discussed
earlier. At any rate, Pleistocene temperate mammal species
living in the southern United States/northern Mexico had
a readier opportunity to disperse southward than those
living at higher latitudes. As this review shows, some such
species (e.g., cervids and ursids and many others, see above)
are frequent components of local faunas from the Ch-CO and
CeP morphotectonic provinces, as shown by local faunas 2, 7,
8, 21 51, 52, 53 and 66 among others (Tables 3 and 12; Fig. 5;
Appendix). Some such species are not unfrequently present
in the TMBV.
4.4. Concluding remarks
This review shows how vast and varied is the information on
Mexico’s Pleistocene. However, it also points out the major weak-
ness in obtaining more from the enormous effort invested by so
many researchers, namely the little coordination existing among
the different working groups, both within this country and outside.
Perhaps the vitally important international concern in under-
standing climate change could be used as a common springboard to
launch a concerted effort to fill the large knowledge gap existing on
the Pleistocene of Mexico in general, and of its mammal fauna in
particular.
5. Conclusions
1 Mexico’s Pleistocene terrestrial mammal record includes 13
orders, 43 families, 146 genera and 278 species. Thus, is
quantitatively comparable to the Recent record (10 orders, 34
families, 161 genera and 479 species), but shows greater ordinal
and family diversity. Extinction near or at the Pleistocene–
Holocene boundary was important, and while very uneven
across taxonomic groups was not overwhelming. Greater losses
largely involved meso- and megabaric species of the extant
Orders Xenarthra, Carnivora, Perissodactyla and Artiodactyla;
Proboscidea were extirpated, while the Orders Notoungulata
and ?Litopterna became extinct. Thus, Mexico’s Recent terres-
trial mammal fauna is highly enriched in microbaric species.
2 The review of Mexico’s Pleistocene mammal, palynologic and
paleosol records shows that they are strongly biased
geographically, so that only a few morphotectonic provinces,
the TMVB and the SMOr, include many sampled localities,
whereas others, e.g. the SMOc, have very few. Geochronologi-
cally, the Late Rancholabrean NALM age is the best studied,
whereas the Irvingtonian and latest Blancan NALMAs remain
very poorly known. Therefore, only broad climatic trends could
be delineated for the entire Pleistocene across the country, as
well as in each of its component morphotectonic provinces. For
the Late Rancholabrean, such trends broadly coincide with
those known for the Wisconsinan. They are not however
strictly coeval throughout the provinces or across them, but
seem to be diachronous. The dating, chiefly biochronlogical
based on mammals, does not allow the necessary precision to
quantify the differences or the amount of displacement in time.
3 The mammal record of the different morphotectonic provinces
seems to show by latest Rancholabrean time (ca. 25–12 ka), an
overall moister and cooler climate regime than at present;
which coincided with a greater mammalian diversity than the
current one. Many of the local faunas of that time are dishar-
monious, much more so those from territories within and
north of the TMVB. The incompleteness of the record does not
allow the precise dating and correlation of this regime and
climate trend with related vegetation and paleosols changes,
but by and large, their records seem to be congruent. These
phenomena are less apparent in the tropical morphotectonic
provinces, namely SMS, CHI, GCP southern part, and YPL.
4 The Wisconsinan climatic fluctuations is reflected in major
changes in the distribution of species, which impacted the
mammal fauna resulting in a very different physiognomy than
 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104
that of the Recent fauna, both in general and in each province.
Such distribution and related physiognomic changes partly
represent the mammalian faunal response to climatic change,
which climaxed by the end of the Rancholabrean with the
ensuing of the current climate regime. The major extinction
event, already dealt with, also could be regarded as part of the
Pleistocene mammal fauna response.
5 The complex faunal response to the Wisconsinan (and earlier)
climatic changes, is reflected in the Recent mammal fauna’s
intricate, tapestry-like biogeographic pattern, whereby
temperate and tropical species associations (e.g. communities
or species groups) occur outside their respective latitudes
compared to modern distributions. The Pleistocene mammal
record does not allow precise mapping of the biogeographic
distributional changes involved, nor the particular path fol-
lowed by any given species to account for their Pleistocene and
current distribution. It does, however, allow, in combination
with geologic data and present-day mammal distribution,
possible major Pleistocene dispersal routes to be discerned:
high and low land, southward corridors for temperate species
and low land, northward ones for tropical species.
6 Finally, the existence of a single Mexican Rancholabrean faunal
or biogeographic province is incompatible with Mexico’s
Rancholabrean mammal record composition and distribution
in the different morphotectonic provinces, and with the
marked contrast between the faunas from territories within
and north of the TMVB compared with those from provinces
south of it. Instead, a multi-province scheme largely corre-
sponding with the current regionalization is needed, which
recognizes the environmental impact that each morphotec-
tonic province territory imposes on the biota it contains, is
offered as a working hypothesis for better understanding
97
Mexico’s Pleistocene mammal biogeography, faunistics and the
origin of Mexico’s modern mammalian faunal patterns.
Acknowledgments
The authors thank Eric Scott and H. Gregory McDonald for their
kind invitation to participate in the volume dedicated to recognize
the outstanding contributions of Dr. Ernest L. Lundelius to the field
of Pleistocene Mammalian Paleontology. The senior author partic-
ularly is indebted to Dr. McDonald for allowing extra time to finish
this paper, due to severe and long lasting health problems he
suffered, for his encouragement to go on, and for his careful and
sympathetic editing, which greatly improved the paper. We also
duly thank Dr. Gary Morgan, as his meticulous editing allowed us to
better the paper a great deal. The authorities of the Instituto de
Geologı́a, Universidad Nacional Autónoma de México at Mexico
City, wholeheartedly supported the project and provided the time
and resources to make it possible. The Instituto Nacional de
Antropologı́a e Historia, in Mexico City, and the Museum of Texas
Tech University at Lubbock, TX, also supported the project and
furnished valuable resources to develop it. The authors express
their sincere thanks to these institutions, and to their authorities.
Appendix
Figure A1 shows selected main Pleistocene terrestrial mammal
localities of Mexico set on the morphotectonic provinces template
pictured in Fig. 1. Time frame adapted from Bell et al., (2004). The
geographic position of the localities is approximate. Chief sources:
Alvarez (1965), Barrios-Rivera (1985), QMMDB (Arroyo-Cabrales
et al., 2002), and Arroyo-Cabrales et al., (2007a,b). For further
information see Table 3.
98 I. Ferrusquı́a-Villafranca et al. / Quaternary International 217 (2010) 53–104

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