Michael O. Woodburne1,* Alberto L. Cione2,**, and Eduardo P. Tonni2

Transcription

73
Woodburne, M.O.; Cione, A.L.; and Tonni, E.P., 2006, Central American provincialism and the
Great American Biotic Interchange, in Carranza-Castañeda, Óscar, and Lindsay, E.H., eds., Advances in late Tertiary vertebrate paleontology in Mexico and the Great American Biotic Interchange: Universidad Nacional Autónoma de México, Instituto de Geología and Centro de
Geociencias, Publicación Especial 4, p. 73–101.
Central American provincialism and the Great American Biotic Interchange
Michael O. Woodburne1,*
Alberto L. Cione2,**, and
Eduardo P. Tonni2,***
ABSTRACT
The age and phyletic context of mammals that dispersed between North and South America
during the past 9 m.y. is summarized. The presence of a Central American province of
cladogenesis and faunal differentiation is explored. One apparent aspect of such a province
is to delay dispersals of some taxa northward from Mexico into the continental United
States, largely during the Blancan. Examples are recognized among the various xenarthrans, and cervid artiodactyls. Whereas the concept of a Central American province has
been mentioned in past investigations it is upgraded here. Paratoceras (protoceratid artiodactyl) and rhynchotheriine proboscideans provide perhaps the most compelling examples
of Central American cladogenesis (late Arikareean to early Barstovian and Hemphillian to
Rancholabrean, respectively), but this category includes Hemphillian sigmodontine rodents,
and perhaps a variety of carnivores and ungulates from Honduras in the medial Miocene,
as well as peccaries and equids from Mexico. For South America, Mexican canids and hydrochoerid rodents may have had an earlier development in Mexico. Remarkably, the first
South American immigrants to Mexico (after the Miocene heralds; the xenarthrans Plaina
and Glossotherium) apparently dispersed northward at the same time as the first Holarctic
taxa dispersed to South America (sigmodontine rodents and the tayassuid artiodactyls).
The main (Phase One) Great American Biotic Interchange (GABI) bipolar episode
transpired from about 2.7–1.8 Ma, with laggards lasting until about 1.0 Ma. A later phase
occurred from about 0.8 Ma to virtually modern times and resulted in mainly southern
enrichment.
Key words: Great American Biotic Interchange, vertebrate paleontology, dispersal,
provinciality.
RESUMEN
Se presenta en forma sucinta la edad y el contexto filético de los mamíferos que se dispersaron
entre América del Norte y del Sur durante los pasados nueve millones de años. Se explora
la posible presencia de una provincia Centroamericana de cladogénesis y diferenciación
faunística. Un aspecto relevante de tal provincia es el retardo en la dispersión de algunos
taxones hacia el norte, de México a los Estados Unidos de América, principalmente durante
el Blancano; como ejemplos de ello se tiene a los diversos xenartros y los artiodáctilos
cérvidos. El concepto de una provincia Centroamericana de tales características ya ha
sido tratado en investigaciones pasadas; en el presente estudio, dicho concepto es afinado.
Department of Geology, Museum of Northern Arizona, 3101 N. Fort Valley Rd., Flagstaff, AZ 86001
*E-mail address: [email protected]
2
Departamento Científico Paleontología Vertebrados, Museo de La Plata, 1900 La Plata, Argentina
**E-mail address: [email protected]
***E-mail address: [email protected]
1
Carranza-Castañeda, Óscar, and Lindsay, E.H. eds., Advances in late Tertiary vertebrate paleontology in Mexico
2006
74
Woodburne, Cione and Tonni
Quizá los dos ejemplos más convincentes de cladogénesis en América Central sean el de
Paratoceras (artiodáctilo protocerátido del Arikareeano tardío al Barstoviano temprano)
y el de los proboscídeos ryncotherinos (Henfiliano a Rancholabreano); adicionalmente,
esta categoría incluye roedores sigmodontinos del Henfiliano y, quizá, una diversidad de
carnívoros y ungulados de Honduras en el Mioceno medio, así como de pecaríes y équidos
de México. Para América del Sur, los cánidos y los roedores hidrocoéridos pueden haber
tenido un desarrollo más temprano en México. Es notorio que los primeros inmigrantes
sudamericanos a México (después de los precursores del Mioceno: los xenartros Plaina y
Glossotherium) se dispersaron hacia el norte, a la vez que los primeros taxones holárticos se
dispersaron a Sudamérica (roedores sigmodontinos y artiodáctilos tayasuidos).
El episodio bipolar principal (Fase Uno) del Gran Intercambio Biótico Americano
(GABI, por sus siglas en inglés) transcurrió durante el intervalo 2.7–1.8 Ma, con rezago
que subsistió hasta hace cerca de 1.0 Ma. Una fase posterior ocurrió hace aproximadamente
0.8 Ma, en tiempos virtualmente modernos, y tuvo como resultado un enriquecimiento
principalmente meridional.
Key words: Gran Intercambio Biótico Americano, paleontología de vertebrados, dispersión,
provincialismo.
INTRODUCTION
Studies of land mammal dispersal between North and
South America usually focus on the Great American
Interchange or Great American Biotic Interchange
(GABI) which began about 2.7 Ma and lasted into virtually modern times. The dispersal episodes are populated by suites of animals that display taxonomic facies
which differ not only geographically, but temporally.
The GABI events are summarized here before moving
on to a discussion of faunal provinciality, evolution,
and dispersal with which participants in the GABI must
have been associated.
The Great American Biotic Interchange reflects
the end of the Cenozoic isolation of South America with
respect to the land mammals and birds of that region in
contrast to those of Central and North America (Marshall
et al., 1979). This episode witnessed land mammal dispersals across the newly established Isthmus of Panama,
and resulted in faunas of both areas being enriched by
taxa from the other. In South America, North American
mammalian immigrants included tayassuids, rodents,
canids, gomphotheriids, camelids, cervids, felids, and
ursids, although not all at the same time. For North
America, newcomers were largely edentates, but also
included rodents and marsupials.
It is well known that these dispersals took place
at different times in the past 9 m.y. and that certain taxa
(procyonids, gomphotheres) reached South America
by 7–9 Ma (Table 1) well in advance of 2.7–2.5 Ma
(e.g., Campbell et al., 2000; Carranza-Castañeda and
Miller, 2004; Morgan, 2005), at least in part coincident with the earliest immigration of ground sloths
(Pliometanastes, Thinobadistes) to North America (ca.
8–9 Ma; Hirschfeld, 1981; Webb 1985; Morgan, 2005).
After procyonids, the first North American taxa are
sigmodontine rodents in Montehermosan beds perhaps
at 5–6 Ma. For many years, a classical view that prevailed among workers maintained that representatives
of two families of North American origin occurred in
Chapadmalalan beds of southern South America and
eight in post Chapadmalalan-pre-Ensenadan beds (the
“Uquían”; see Marshall et al., 1984; Webb, 1985).
However, a new emphasis on biostratigraphical studies demonstrated that the first occurrences of different
species of North American families in the south are
distributed in beds that encompass more than 4 million years (one in the Chapadmalalan, one in the lower
Marplatan, two in the middle Marplatan, one in the upper Marplatan and four in the Ensenadan [Tonni et al.,
1992; Cione and Tonni, 1995, 2001, 2005]). In addition there are other species of five families of North
American origin in very recent beds or with no fossil
record at all (Webb, 1985; Cione and Tonni, 1995).
More recently, Carranza-Castañeda and Miller
(2004) and Flynn and others (2005) showed that
Universidad Nacional Autónoma de México, Instituto de Geología and Centro de Geociencias
Publicación Especial 4
Neotropical immigrants to NorthAmerica, Glossotherium
and Plaina, were present in the vicinity of Guanajuato,
central Mexico, by 4.6–4.8 Ma, and that Glossotherium
reached the USA somewhat later (3.0 Ma, Table 1), but
still prior to the beginning of the late Blancan and the
GABI. According to Tonni and others (1992) and Cione
and Tonni (1995), Plaina occurs between 3.3 and 5.0
Ma in the Chapadmalalan SALMA (South American
Land Mammal Age) of Argentina. Whether this reflects
an evolutionary role for Central America and southward dispersal cannot be solved at present (and the genus seemingly does not occur in the USA in any case).
Megalonyx also is known from the early Blancan sites at
Guanajuato but rather than being designated as an immigrant at Guanajuato by Carranza-Castañeda and Miller
(2004) and Flynn and others (2005), Megalonyx is considered as an endemic North American descendant of
Pliometanastes (e.g., Morgan, 2005). The reason(s) for
the delayed dispersal of these and other taxa (Neochoerus,
Glyptotherium) from central Mexico to the USA remain
cryptic, as discussed further below.
From the southern South American perspective,
a number of workers in the past decade have made
considerable advances relative to the composition and
timing of the units that contain North American immigrants. Chief among these are Tonni and others (1992),
Cione and Tonni (1995, 2001, 2005), Cione and others
(2001), and references cited therein. Other students of
the GABI also have made significant advances in synthesis and interpretation (e.g., Marshall and Sempere,
1993; Webb and Rancy, 1996; Carranza-Castañeda
and Miller, 2004; Morgan, 2005). The following is designed to bring these data and inferences into a general
synthesis and to offer suggestions as to implications
for the continuity of dispersal between the Americas.
The dispersal history is summarized in Table 1 which
shows the taxa in North and South America that pertain to land mammal families that participated in the
faunal exchanges between them. In some instances the
South American record includes notation regarding the
North American record of certain genera. Not all South
American taxa are immigrants, but are included to provide an impression of endemic radiations. Appendix I
provides comments on the timing and contextual aspects of the taxa involved, including endemic genera.
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TECTONIC/GEOLOGIC BACKGROUND
Coates and others (2004) indicate that the Panamanian
region was progressively uplifted from about 12 to at
least 4.8 Ma, with concomitant disruption of formerly
confluent marine depositional environments and faunas, and reflected in the overland dispersals across the
Panamanian region at 8–9 Ma. Continued tectonic activity raised the region still more, coincident with the
mammalian dispersals at ca. 4.8 Ma and subsequently (GABI), after a brief episode of eustatic sea level
rise that breached the peninsula at 6–7 Ma. According
to Haug and Tiedemann (1998) and Haug and others
(2001) the Panamanian isthmus was a firm barrier to
Caribbean-Pacific circulation from 4.7–4.2 Ma, and
Hoernle and others (2002) show how the Caribbean
plate presented an overland dispersal route in the Late
Cretaceous-Paleocene as well as in the Pliocene.
For central Mexico, the Transmexican Volcanic
Belt (Figure 1) experienced a significant episode of
bimodal volcanism in the Hemingfordian-Barstovian
interval (Carranza-Castañeda and Miller, 2004) but the
effects of this activity were followed in the Hemphillian
and Blancan, at least, by erosion and basin filling with
intermittent deposition of air-fall tuffs (e.g., Flynn et
al., 2005). There seems to be no obvious or convenient geological barrier between central Mexico and
the United States to account for the temporal disjunction of dispersals between those places. The Basin and
Range Province, that extends from Arizona and New
Mexico into northern Mexico (Henry, 1989; Kowallis
et al., 1998) likewise had largely ceased activity by the
Hemphillian and in any case also provides no boundary
setting.
ECOLOGIC FACTORS
Webb and Rancy (1996) summarize the ecologic context of Central America and adjacent areas. Prior to and
coeval with the GABI, the region supported a general
savanna setting with relatively open conditions. This
also continued during the interval of glacial cooling
(from about 2.5 Ma) with concomitant lowering of sea
level (Figure 2). Recent research confirms that the onset of significant glaciation in the Northern Hemisphere
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TMVB
Scale
Figure 1. Map of Mexico-Central America showing fossil localities discussed in the text. A = Aguascalientes; G = Guanajuato; H =
Honduras; M = Mixson’s Bone Bed, Florida; P = Panama; O = Oaxaca and Chiapas; Y = Yepómera. TMVB = Transmexican Volcanic
Belt.
began at 2.7 Ma (Haug et al., 2005). This climatic event
must have had a remarkable influence on the vegetation
as well as opening up areas of coastal habitat not previously available, and surely favored the dispersals in the
Pliocene that formerly were not so facilitated.
The first and main episode of interchange
transpired from late Blancan (ca. 2.7 Ma) into the
Irvingtonian (ca. 1.4 Ma), and was populated mostly
by savanna-dwelling forms living under perhaps climatically somewhat cooler conditions than had obtained in the Hemphillian. This is also reflected in the
immigrants largely being drawn from ancient South
American stocks and is demonstrated in the diversity of
such taxa appearing in Central America: descendants of
Oligocene rodents and primates, and older stocks such
as marsupials and xenarthrans (armadillos, anteaters,
and sloths).
By about the beginning of the middle Pleistocene
(ca. 0.8–0.6 Ma), rainforest conditions extended from
South America to Central America, and contributed to
the diminished dispersals from that time onward (Figure
2). Webb and Rancy (1996) point out that elevational
partitioning of subtropical versus more savanna-like
conditions in the Andes played a part in facilitating the
main GABI dispersals into North America of temperate South American taxa on one hand, and on the other
promoted provincialism in the later Pleistocene and
Recent (see also Cione et al., 2003). In South America
the mid- to late Pleistocene record shows diversification
of groups having arrived earlier from North America,
with remarkably wide distribution. Clusters of endemism are recorded by the north temperate mastodon,
mammoth, Bison and Camelops restricted to Central
America. A component of south temperate forms also
shows regional endemism (pampas deer, pampean
glyptodonts and armadillos) in southeastern Brazil.
DISPERSAL SCENARIO
Pre-GABI, Late Miocene and Early Pliocene
A summary of recent literature results in the following reconstruction of dispersal participants and timing (Table 1; Appendix I). As indicated above, megalonychid and mylodontid edentates (Pliometanastes,
Thinobadistes) apparently swam across the Caribbean
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Figure 2. Chronology of North and South American immigration episodes. South American record is after Cione and Tonni (2001). North
American episodes are as in the text. For North American dispersals taxa in () are younger USA occurrences subsequent to an earlier record
in Mexico. GABI = Great American Biotic Interchange. Relative chronological sequence in the late Blancan record is comparable to ages
in Table 1. Arrows show H (high) and L (low) sea levels, after Gradstein and others (2004). * = immigrant from USA.
2006
Table 1. First occurrences of taxa involved in the Great American Biotic Interchange.
78
about 9 Ma, possibly coincident with the Peruvian
gomphothere Amahuacatherium. Gutiérrez and others
(2005), and Alberdi and others (2004) express reservations about the age and taxonomy of this genus. López
and others (2001) consider proboscideans to first appear
in the Sanandresian of Argentina, or at about 2 Ma.
A procyonid carnivore followed at about 7 Ma
(Cione and Tonni, 2001; Baskin, 1989), resulting in the
endemic genus Cyonasua. The North American xenarthran Megalonyx apparently evolved endemically from
Pliometanastes by about 7 Ma, so its presence in central Mexico at 4.7 Ma (Carranza-Castañeda and Miller,
2004; Flynn et al., 2005) has no bearing on the early
development of the Panamanian isthmus (also Morgan,
2005).
The central Mexican record of Glossotherium and
Plaina (Carranza-Castañeda and Miller, 2004; Flynn et
al., 2005) at about 4.8 Ma appears to reflect geological evidence (above) regarding the establishment of the
Isthmus of Panama. In that the earliest record of Plaina
in South America appears to be of Chapadmalalan age
(within about 3.3–5.0 Ma), the Mexican occurrence
may be the oldest for that genus.
Baskin (1986) summarizes evidence in favor of
the Hemphillian origin and diversification of sigmodontine rodents in North America prior to their diversification in Mexico in the late Hemphillian (CarranzaCastañeda and Walton, 1992) and dispersal to South
America in the early Blancan. As revised here, these
rodents arrived in the late Montehermosan, considered (Pardiñas and Tonni, 1998) as about 5.8 Ma
(Figure 2), and about contemporaneous with their record in Mexico. Endemic members of this immigrant
group (Auliscomys, Necromys) are sparse but present
in the type Montehermosan in southern Buenos Aires
province (Table 1). Diego Verzi (2006, personal communication to Cione) considers that an indeterminate
sigmodontine from possibly coeval deposits in northern La Pampa Province (Argentina) is more primitive
than Auliscomys or Necromys (see also Montalvo et al.,
2000). Sigmodontine rodents are well represented (and
phyletically diverse; Table 1) by the Chapadmalalan at
ca. 4.0 Ma. Their presence in South America is consistent with the Clarendonian and younger record of
sigmodontines in North America. According to Baskin
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(1986), Abelmoschomys, from the latest Clarendonian
Love Bone Bed of Florida, is an early sigmodontine.
Antecalomys (Korth, 1998) is a contemporaneous
form from Nebraska. Baskin (1978) described the late
Hemphillian Bensonomys from Arizona and placed it as
a subgenus of the living South American genus Calomys
(but see Pardiñas et al., 2002, for a different opinion). Bensonomys also occurs in the late Hemphillian
Yepómera Fauna, northern Mexico (Lindsay and
Jacobs, 1985), and the Rancho El Ocote Fauna of
Guanajuato, central Mexico (Carranza-Castañeda and
Walton, 1992). Other North American sigmodontines
include Repomys (late Hemphillian to late Blancan;
Tedford et al., 1987) and Jacobsomys (early Blancan
Verde Fauna, Arizona; Czaplewski, 1987).
The Chapadmalalan tayassuid Platygonus is an
additional immigrant at about 4 Ma in association with
the endemic genus Argyrohyus. Wright (1998) indicates
that this peccary group occurs in late Hemphillian faunas of North America.
At about 3.5 Ma, Capromeryx apparently entered
the USA from a slightly earlier presence in Mexico,
and whereas this has nothing to do directly with the
GABI, it does address the question of dispersal barriers
between the continental United States and districts to
the south (see below). In any case, another precursor
of the GABI, Neochoerus, is found in central Mexico
about then (3.3 Ma) also attesting to the likely presence
of the Panamanian isthmus at that time.
If it had a southern source, the dispersal of
Neochoerus to Mexico apparently transpired at about
3.3 Ma, coeval with the latest Chapadmalalan (Figure
2). The question of source stems from the observation
that the earliest occurrence of Neochoerus in South
America apparently is in the Ensenadan, at levels
younger than about 2.0 Ma (Table 1).
The early Marplatan saw the immigration to
South America of camels (Lama) at about 3.0 Ma.
Glossotherium and the other taxa discussed next have
their earliest or an old occurrence in Florida, but many
also are found in other sites in the continental USA
(Morgan, 2005). At about 3.0 Ma, Glossotherium is
found in the USA subsequent to its Mexican earlier
occurrence. The USA record is part of the enigma of
dispersal delay from Mexico (along with Capromeryx,
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Neochoerus, and perhaps Glyptotherium) and the
absence of certain taxa found in Central Mexico
(Plaina).
The GABI
The regional basis for the main immigration episode
in North America indicates that it likely began about
2.8 Ma with the first occurrence of Glyptotherium in
central Mexico, and an entry into the United States
very shortly thereafter, along with Neochoerus. Most
taxa involved in the beginning of the GABI in the
USA are members of late Blancan faunas in North
America (2.7–1.6 Ma) and of mid and late Marplatan
(Vorohuean and Sanandresian) and Ensenadan faunas
in South America (from about 2.6 Ma; Figure 2; Table
1; Appendix I).
Beginning about 2.6 Ma or somewhat later, middle Marplatan (Vorohuean) immigrants include mustelids (Galictis and Stipanicicia), canids (Pseudalopex),
and equids (Hippidion, Onohippidium). Late Marplatan
(Sanandresian) immigrants include the first gomphothere (López et al., 2001). Cervids (Epieuryceros,
Antifer) appear in the Ensenadan. Appendix I indicates
that North American stocks from which the dispersants
derived were present prior to the time of the GABI.
For the GABI in the USA, the porcupine Erethizon
apparently has no Mexican precursor, but accompanies
the above other two taxa to the USA at about 2.7 Ma.
Erethizon was recently identified in Sanandresian beds
in the Uquía Formation of northwestern Argentina
(Reguero et al., 2006). The new South American presence of this taxon, if endemic, implies a record in
South America equal to or older than 2.7 Ma. If a South
American immigrant, this event could have transpired at about 2.0 Ma from North American Erethizon.
In any case, the interval from about 2.5–2.1 Ma saw
the North American immigration of the pampatheres
Pampatherium and Holmesina, the armadillos Dasypus
and Pachyarmatherium, the megathere Eremotherium,
and the rodent Hydrochoerus.
In South America, Ensenadan immigrants include
the tapirid Tapirus; the gomphotheres Cuvieronius
and Stegomastodon; the ursid Arctotherium; the canids Cerdocyon (with a Mexican progenitor) and
Chrysocyon (with a Blancan record in North America;
Berta, 1987); the felids Smilodon, Puma, and Panthera;
and the camel Palaeolama. Catagonus is known from
the late Hemphillian of North America and the Blancan
of Mexico, as well as the Ensenadan to Recent of South
America (Wright, 1998). As indicated in Appendix I all
of these taxa, possibly except Palaeolama, have a prior
occurrence in North America. Depending on the age of
the Ensenadan SALMA the GABI in South America
recorded its major pulse from about 2.6–about (or older
than) 1.0 Ma, fundamentally similar to the late Blancan
to early Irvingtonian in North America.
For North America, this “Phase One” (Webb
and Rancy, 1996) of the GABI appears to be largely
over by the end of the Blancan, at about 1.8 Ma, with
later gasps apparently represented by the megalonychid Paramylodon (Morgan, 2005), the megathere
Nothrotheriops (ca. 1.5 Ma), Myrmecophaga (El Golfo
Local Fauna, about 1 Ma; Webb, 1991), and Didelphis
(about 1.2–0.8 Ma in Florida and Texas; Morgan,
2005). As indicated above, a second wave of North
American taxa entered South America between about
1.8 and 1.0 Ma. If those correlations of Ensenadan taxa
are accurate, it appears that both these and the later
North American immigrations are in the terminal part
of “Phase One”.
The late Pleistocene phase begins about 0.8 Ma,
with the Bonaerian (the Lujanian is restricted to the
latest Pleistocene; Cione and Tonni, 2005) faunas
experiencing a number of immigrants (Marshall et
al., 1979). These include the procyonid Nasua, the
mustelids Galera and Lutra, the rodent Calomys, the
peccary Tayassu, and the cervid Ozotoceros, among
others (Table 1). If most of these are Lujanian as
restricted by Cione and Tonni (2005), then they
would be about contemporaneous with those of
Rancholabrean age in North America, with taxa such
as Cryptotis, Sylvilagus, Lycalopex, Mustela, and
Mazama. For North America, the late Pleistocene
invaders mostly do not seem to have come directly
from South America, but reinvaded North America in
the Rancholabrean after having moved southward in
the late Irvingtonian: Glyptotherium, Eremotherium,
Desmodus, Neochoerus, and Hydrochoerus (Morgan,
2005, p. 300L).
THE PANAMA-MEXICO FAUNAL PROVINCE
Figure 3 summarizes the record of taxa in central
Mexico and in Panama that seem to pre-date their
counterparts in the USA, in addition to those found on
Table 1. If the timing disjunction is accurate and pertinent, it suggests that some factor or factors seem(s)
to retard the dispersal to the USA from this southern
region, on the order of 1 m.y. in most cases. If the
Panamanian example is not one of dispersal lag, but
one of prior lineage development, that pattern also is
not recognized in the USA at this scale for the groups
involved. Rhynchotherium provides another likely example of the endemic development of a lineage with
only subsequent manifestation in the USA, as well as
implications for dispersal southward. Webb and others (2003) review the history of the Protoceratidae and
81
indicate that the group was present in Central America
(Chiapas, Mexico) from about 26 Ma (Paratoceras tedfordi), with the genus representing a ‘southern branch’
of the “Protoceratinae” and Kyptoceras the ‘southern
branch’ of the Kyptoceratinae.
Kirby and MacFadden (2005) evaluated the
mammals of the Panamanian Gaitán Fauna (Cucaracha
Formation) regarding the question of whether the body
size of these taxa reflects regional isolation of islandlike dimensions, or is within the ranges expected for
taxa that were not isolated from counterparts in distant
locations. The goal of the investigation was to evaluate whether Central America was a series of isolated
islands, of which Panama was one, or whether the
region was an interconnected landscape at least during the medial Miocene (ca. 15 Ma). They concluded
that the Cucaracha mammals were not isolated bio-
Figure 3. Age of selected early occurrences of taxa in Mexico and Central America relative to their USA record. Boldface numbers refer
to those used in the text.
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82
geographically from North American congeners, but
that genetic interchange occurred readily between and
within populations at both ends of the zoogeographic
extremes, north and south, at least for members of the
Canidae, Oreodontidae, Protoceratidae, Equidae, and
Rhinocerotidae. In this context the mammals surveyed
suggest that Central America was not a series of isolated islands in the medial Miocene.
In their survey of mid- to late Pleistocene faunas
in South and Central America, Webb and Rancy (1996)
show clusters of endemism, recorded by the north
temperate mastodon, mammoth, Bison and Camelops
occurring no farther south than Honduras, in Central
America, with the same faunas recording the northernmost occurrence of the South American notoungulate,
Mixotoxodon. A component of south temperate forms
also shows regional endemism (pampas deer, pampean
glyptodonts and armadillos) in southeastern Brazil and
other areas of South America (see papers in Tonni and
Cione, 1999). In general, however, Webb and Rancy
(1996) find little other regional endemism in this
part of the time scale, as well as earlier. MacFadden
(2006a) provides a stimulating summary of faunal and
ecological evolution in Central America and the northern part of South America, but notes no major provincial differentiation within that region or relative to
adjacent areas to the north or south during the past 25
m.y., except for the above-noted late Pleistocene pattern that the Mexican-Central American province records the southernmost limit of Bison and Odocoileus
virginianus (Mexico) and the northernmost limit of
toxodonts from South America (Costa Rica; Lucas et
al., 1997).
The evidence from some of the late Cenozoic faunas in the Mexican-Central American region suggests
a somewhat more complex pattern, on the other hand.
These examples are numbered for easier reference in
Figure 3.
recovered from amber-bearing beds of the Balumtum
Sandstone (Webb et al., 2003). This unit is considered
to be early Miocene in age, and generally considered to
range from 22–26 Ma. In Figure 3 the site is correlated
on that basis with the ‘late Arikareean.’
As reviewed by Prothero (1998), the Protoceratidae
originated in North America in the Uintan (medial
Eocene, ca. 46 Ma), underwent a modest radiation
until the end of the Eocene (33 Ma), and persisted in
diminished numbers until the end of the Hemphillian
(ca. 4.5 Ma). One genus, Pseudoprotoceras, is present
in the Duchesnean and Chadronian (ca. 40–33 Ma.)
and founded the “Protoceratinae” which also contains
Protoceras (Whitneyan to late Arikareean, ca. 32–19
Ma) and Paratoceras now known to range from late
Arikareean (ca. 26–22 Ma) to early Clarendonian (ca.
10 Ma). Kyptoceratine protoceratids (Syndyoceras
[23–18 Ma], Kyptoceros [ca. 5 Ma]) comprise the
Kyptoceratini. According to Webb and others (2003),
both the “Protoceratinae” and Kyptoceratini are represented by forms in a northern (High Plains) and
southern (Gulf Coastal Plain) region, including Central
America. In both cases the southern region taxa persist
after the extinction of the northern forms, and are characterized as having retreated southward “in apparent
association with humid subtropical habitats” (Webb et
al., 2003: 363–364). In that Paratoceras was present
in the Chiapas region of Mexico as early as perhaps 26
Ma, is recorded in the Cucaracha Fauna of Panama at
about 17 Ma (Kirby and MacFadden, 2005), as well as
about 15–13 Ma in southern Texas and at about 10 Ma
in the Texas Panhandle (Prothero, 1998), the Central
American record for the genus supports an interpretation that it represents a group that has been endemic to
this region since the late Arikareean and that subsequent
to its apparent earlier immigration to this area from the
USA it was a member of an endemic Mexican-Central
American biota.
Mexico
2. Early Hemingfordian. Aguascalientes (A, Figure 1)
1. Late Arikareean. Chiapas (O, Figure 1)
Zoyatal Local Fauna. First nominated as early
Barstovian by Dalquest and Mooser (1974), this fauna is considered to be early Hemingfordian in age
(Tedford et al., 2004:201R). The fossils occur in the
Simojovel Local Fauna. The protoceratid artiodactyl
Paratoceras tedfordi is the sole taxon from this site,
Zoyatal Tuff (not dated), that contains Merychyus cf.
elegans, Aguascalientia wilsoni, Dyseohyus cf. stirtoni,
and Menoceras. Tedford and others (2004) base the age
largely on the oreodont Merychyus, which is a common
early Hemingfordian taxon. If the peccary actually is
Dyseohyus, it is an oldest occurrence of that genus,
even if D. stirtoni is regarded as “Prosthennops” xiphodonticus (Wright, 1998).
Dalquest and Mooser (1974) gave measurements
of the M3 of MU 8686 (not 8696 as stated by them) as
16.0 and 15.8 mm long and 16.2 and 15.9 mm wide.
The length, but not the width of these teeth is comparable to Dyseohyus stirtoni and substantially greater than
for D. fricki or any species of Cynorca (Woodburne,
1969). The Mexican teeth also seem to have a more attenuated posterior portion of M3 than in Dyseohyus or
Cynorca. At this point they are unlikely to be referable
to any of these species or genera, even if D. stirtoni is
considered as a member of “Prosthennops” xiphodonticus, as in Wright (1998). Nevertheless the specimens
do suggest that they represent an early population of
the group to which these genera pertain and suggest the
presence of such a radicle in Mexico that is unknown in
coeval deposits in the USA.
As summarized by Wright (1998), the
Tayassuini begins with “Cynorca” occidentale of late
Hemingfordian age in North America (also Woodburne,
1969), which forms a polytomy with Dyseohyus, first
known from the late Barstovian or ca. 15 Ma. If future research shows that the Zoyatal specimens pertain to this group rather than to lineages known from
early Hemingfordian faunas in the USA (Hesperhys,
Floridachoerus), then an early tayassuine evolutionary
theater is suggested for at least the northern part of the
Mexican-Central American province.
3. Late Hemingfordian, Oaxaca (O, Figure 1)
Suchilquitongo Local Fauna. This fauna is from the
Suchilquitongo Formation which occurs stratigraphically 80 m above tuffs dated at 19.2 ± 0.3 and 20.6 ± 0.6
Ma (Ferrusquía-Villafranca, 2003). The taxa include
Merychyus minimus, Merychippus sp., a protoceratid
referred to Paratocereas, and a kyptoceratine protoceratid more evolved than Syndyoceras cookei, according
83
to Ferrusquía-Villafranca (1990, 2003), who assigned
the fauna an early late Hemingfordian age (Tedford
et al., 2004:202L). Syndyoceras cookei is from the
Harrison and Marsland formations (Prothero, 1998) or
Ar3 and 4, and Paratoceras is known elsewhere from
faunas of late Arikareean to Clarendonian age (Webb
et al., 2003; Prothero, 1998). The Suchilquitongo protoceratids are not otherwise age-diagnostic. Merychyus
minimus (if this is the same as M. elegans minimus in
Lander, 1998), is listed by him from the Martin Canyon
beds (early Hemingfordian to early Barstovian), the
Lower Cady Mtn. L.F. (early Hemingfordian), possibly from the Garvin Farm L.F. (early Hemingfordian),
and Tick Canyon Formation (Ar4). Woodburne (1998)
removed M. calaminthus (= M. elegans minimus of
Lander, 1998) from the Lower Cady Mts. Local Fauna
and indicated its age as Ar 3 (late early Arikareean).
In the context of Lander (1998), M. elegans minimus
ranges from early Arikareean to early Barstovian, of
little use in close correlation.
Even if the material designated as Merychippus
pertains to another taxon of comparable hypsodonty, such as “Merychippus” or Cormohipparion
(Woodburne, 2003, in press), it is unlikely to be older
than 17 Ma, even if specimens from the Box Butte
Fm. (Galusha, 1975) are so assigned (Hulbert and
MacFadden, 1991). The Suchilquitongo Local Fauna
is most useful in supplying a likely late Hemingfordian
age for its record of Paratoceras.
4. Late early Barstovian, Chiapas (C, Figure 1)
Ixtapa Local Fauna. Ferrusquía-Villafranca (1990),
and Jiménez-Hildago and others (2002) indicate that
the Ixtapa Local Fauna is from the Ixtapa Formation in
the Ixtapa Graben, in Chiapas, Mexico. The site is from
the lower part of the formation about 200 m below tuffs
that yielded K-Ar ages of 15.02 ± 0.35 and 16.02 ±
0.53 Ma on biotite and plagioclase, respectively. These
ages suggest that the fauna is not younger than early
Barstovian. The fauna contains Cormohipparion,
apparently near C. quinni, cf. Teleoceras and
Gomphotherium. Ferrusquía-Villafranca (1990, 2003)
compares the Cormohipparion with C. occidentale,
but this is presented as C. quinni in Tedford and oth-
2006
84
ers (2004). As reviewed by Woodburne (in press), the
oldest member of the C. occidentale group (C. occidentale, s.l.) is at about 12.5 Ma, in the Burge Member
of the Valentine Formation. Cormohipparion quinni is
first represented in the Cornell Dam Member of that
formation, at about 14 Ma. If the Cormohipparion material proves to be referable to C. quinni (C. sphenodus
in Ferrusquía-Villafranca, 2003), it would pre-date the
earliest occurrence of that taxon in North America by
about 2 m.y. (Woodburne, 1996).
The record of Gomphotherium is one of the oldest in North America, and indicates faunal continuity
with the west coast of the United States (late early
Barstovian North Coalinga Local Fauna), another early occurrence of the genus (ca. 15.2 Ma; Tedford et al.,
2004).
5. Late early Barstovian, Oaxaca (O, Figure 1)
Nejapa Fauna. As summarized in Tedford and others
(2004), the El Cameron Formation contains a late early
Barstovian fauna. The fossiliferous beds are interbedded with the Yautepec Tuff dated at between 15.08 ±
0.8 Ma (biotite) and 16.7 ± 0.71 Ma (plagioclase) and
contain material from the El Gramal Fauna site referred
to “Merychippus,” Gomphotherium, cf. Miolabis, cf.
Protolabis, and a protoceratid, plus two other small
horses. One of these is a small hipparionine comparable to Hipparion shirleyi otherwise known from late
Barstovian faunas of Texas (MacFadden, 1984). The
other equid may be referable to Calippus. Webb and others (2003) indicate that the protoceratid is Paratoceras.
At the correlative El Camarón site taxa include the
mustelid, Plionictis oacacensis, Gomphotherium, a
small protohippine horse, a merycodont, small to medium-sized camels as well as members of the Canidae
and Felidae (Tedford et al., 2004) who estimate a late
early Barstovian age for these sites.
The Nejapa Gomphotherium seems to correlate
with that from Ixtapa, and the presence of this proboscidean in these two sites is pertinent to the early
Hemphillian occurrence of Rhynchotherium blicki from
Honduras (see below). The late early Barstovian occurrence of Paratoceras adds to this taxon’s presence in
the endemic biota of Mexico-Central America.
6. Early Blancan (Bl1), Guanajuato (G, Figure 1)
San Miguel de Allende Fauna. Capromeryx
(Artiodactyla: Antilocapridae). Jiménez-Hidalgo and
others (2004) report the occurrence of Capromeryx
tauntonensis from early Blancan sites in San Miguel
de Allende, Guanajuato, Mexico. Previously the species was known from the Taunton Local Fauna (early
Blancan) of Washington, and tentatively from the Sand
Draw Local Fauna (“mid” Blancan) of Nebraska, correlated to about 3.05 to 2.7 Ma (Bell et al., 2004). The
Mexican occurrence is about 0.5 m.y. older than that
from Taunton, and is compatible with a possible southern center of origin and diversification of the genus.
Janis and Manning (1998) indicate that
Capromeryx is one of the more derived antilocaprids
and has a sister-group relationship with a clade that
produced Hayoceras, Stockoceras, Hexameryx, and
Hexobelomeryx, with an earliest record in the early
Hemphillian (ca. 9 Ma). The intervening gap (early
Hemphillian to early Blancan) in the USA illustrates a
significant absence of a record there which is compatible with a Central American evolutionary center leading to Capromeryx in Mexico, as well as to Tetrameryx
in the United States.
Panama
7. Late Hemingfordian or early Barstovian (P,
Figure 1)
Gaillard Cut Local Fauna. Kirby and MacFadden
(2005) review the fauna from the Cucaracha Formation
of Panama. This fauna contains Texomys stewarti,
Tomarctus brevirostris, cf. Cynorca, Merycochoerus
matthewi, Paratoceras wardi, Anchitherium clarencei, Archaeohippus sp., Menoceras barbouri
and Floridaceras whitei. Based on the analysis of
MacFadden (2006b), the assemblage is a coherent fauna of contemporaneous taxa which in North America
occur in faunas of Hemingfordian (Merycochoerus
matthewi, Menoceras barbouri), Barstovian (Texomys
stewarti, Tomarctus brevirostris, Paratoceras wardi) or
of Hemingfordian-Barstovian (Anchitherium clarencei,
Archaeohippus sp.) age.
Whereas it could be straightforward to choose the
youngest of the potential ages involved, and presume
that taxa of Hemingfordian affinity are locally persistent or undiagnostic (Archaeohippus sp.), the possibility still exists that the earlier age is correct and that the
taxa of Barstovian affinities point to an earlier pattern
of cladogenesis than would be recognized from North
America. If one presumes that the Gaillard Cut Local
Fauna is of late Hemingfordian age, and that this carries a correlated age of about 17 Ma, then the allegedly
Barstovian taxa would pre-date their North American
counterparts by about 1 m.y.
Honduras and El Salvador
8. Early Hemphillian (H, Figure 1)
Gracias and Corinto faunas. Webb and Perrigo (1984)
describe two early Hemphillian faunas from Honduras
and El Salvador, respectively. The Gracias Fauna is
composed of Rhinoclemmys, Geochelone, Osteoborus
cyonoides, Rhynchotherium blicki, Hipparion placatile, Pliohippus hondurensis, Teleoceras cf. fossiger,
Prosthennops cf. serus, Protolabis cf. heterodontus,
Procamelus cf. grandis and Pseudoceras skinneri. The
most age-significant taxa, O. cyonoides, H. plicatile, T.
fossiger, indicate an early Hemphillian correlation for
the fauna, with the Mixson’s Bone Bed Fauna of Florida
being a strong correlate, suggesting a date of about 8–9
Ma (Morgan, 2005) for these Central American faunas.
Webb and Perrigo (1984) discuss the phyletic
position of Rhynchotherium blicki and point out that
it is not only more primitive than its chronologically
next counterpart (R. edensis) from the Mt. Eden and
Warren local faunas of California (correlated at about
5.2 Ma; Tedford et al., 2004), but that this is the oldest American occurrence of the genus. Lambert and
Shoshani (1998) posit a Central American origin for
the group which then dispersed northward to North
America, including the Blancan of Mexico (CarranzaCastañeda and Miller, 2004). This is shown on Figure
3 as Rhynchotherium tlascalae.
Lambert and Shoshani (1998) indicate that
Rhynchotherium is the basal member of a clade including
Stegomastodon (early record is Hh 4), and Cuvieronius
85
(Bl 1). This clade is part of a trichotomy (New World
forms) that phyletically precedes a clade composed of
‘shovel-tuskers’, of which Gomphotherium is the basal
member and has an early occurrence of early Barstovian
in both the USA and Mexico (Nejapa, Ixtapa; Figure
3). Tobien (1973) indicates that Rhynchotherium has
affinity with Cuvieronius and Webb and Perrigo (1984)
conclude that subtropical North America (= Central
America) was an evolutionary center that resulted in
the notiomastodonts Cuvieronius, Haplomastodon,
and Notiomastodon dispersing from there to South
America. The presence of Gomphotherium in late early
Barstovian faunas of Mexico indicates its presence in
the Central American realm, compatible with a hypothesis of the provincial derivation of Rhynchotherium.
DISCUSSION
The proposal that the Neogene fossil mammal record
of Central America contains examples of endemic
evolution is discussed by Webb (1974), FerrusquíaVillafranca (1984), Lindsay (1984), Webb and Perrigo
(1984), Slaughter and Ubelaker (1984; CarranzaCastañeda and Walton, 1992), and Jiménez-Hidalgo
and co-workers (2002), among others. All are consistent with the suggestions made herein based in part on
more recent information.
Many of the examples cited here are based on fragmentary or otherwise only generally diagnostic material,
but the published taxonomic allocations are taken as
stated for the purpose of this discussion. Whereas some
of the examples may be solved by upgrading correlations
(numbers 3, 4, 7; and Figure 3) others seem to definitely
indicate the likely presence of evolutionary provincialism in the Central American region. The late Arikareean
occurrence of Paratoceras tedfordi in Chiapas seems
to herald the presence of a protoceratid clade that underwent a distinct endemic evolutionary radiation in
Mexico-Central America that persisted to within the early
Barstovian (numbers 1, 3, 5, 7; and Figure 3). The Zoyatal
“Dyseohyus” (number 2; and Figure 3) either represents
a major range extension of pre-tayassuine taxa from the
USA, or is evidence favoring an early development of
the tayassuine clade not identified to the north. Perhaps
the most compelling example in addition to Paratoceras
2006
86
is number 8, where the presence of Rhynchotherium
blicki nearly demands that the origin of this group (and
its notiomastodont descendants) were unique to this region. The example of Capromeryx tauntonensis (number 6) speaks favorably for there having been a Central
American center of origin for this group of antilocaprids,
for which there is no record in the USA. At the moment,
the potentially earlier age of Plaina in Mexico versus the
Chapadmalalan of South America (Table 1) also permits
consideration of the role played here by the MexicanCentral American province.
In addition to Cerdocyon, immigrant to South
America in the Ensenadan from a prior apparent ancestor in Mexico, Berta (1987) reviews the remaining
South American canids and comments as to a possible Central American center of diversification for the
group. In that context it is informative that new taxa
appear in both the Marplatan and Ensenadan on Table
1, suggesting at least two separate dispersal episodes.
Regarding rodents, Carranza-Castañeda and
Walton (1992) point to a late Hemphillian diversification of sigmodontines in Mexico prior to their dispersal to South America, and the Mexican occurrence
of Neochoerus pre-dates its Barrancalobian record in
South America.
At this point, it is obvious that with the potential
exception of the late Pliocene and Pleistocene canids
(if they had a Central American radiation), and the
earlier record of Tomarctus in Honduras, the groups
involved all are herbivores and mostly ungulates. But
mesodont to early hypsodont horses (merychippines
and Cormohipparion) are not likely to have been members of the same ecologic or adaptive facies as were
peccaries, proboscideans, antelopes, or sigmodontine
and hydrochoerid rodents, even if all were basically savanna-dwellers (e.g., Webb, 1991). Preceding sections
recall numerous references attesting to changes in climate and tectonic setting in this region over the past 9
m.y., as well. Thus it is unlikely that whatever kinds of
provincialism operated, there was no single cause for
the patterns articulated above. Whatever their causes,
patterns of secular evolution as well as chronologic
disjunctions in dispersal demonstrate the importance of
Central America’s role in the Neogene faunal development of North and South America.
ACKNOWLEDGMENTS
This manuscript was initiated at the invitation of Drs.
Oscar Carranza-Castañeda, Centro de Geociencias,
Universidad Nacional Autónoma de México, Querétaro,
and E.H. Lindsay, Department of Geosciences,
University of Arizona, Tucson; the first author thanks
them for not only the invitation, but also help and encouragement along the way. Other colleagues who provided assistance include Drs. Gary Morgan, New Mexico
Museum of Natural History, Albuquerque, and Bruce
J. MacFadden, Florida Museum of Natural History,
Gainesville, with especial insight into Neotropical faunas both north and south of the Panamanian isthmus.
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APPENDIX I
Time and Entry
This summarizes the evidence for the timing and taxa
involved in the dispersal. *Underlined taxa are immigrants
to South America (also on Table 1). Taxa indicated as e are
endemic to South America, having been locally derived after
immigration. All chronologic references to Vrba (1993)
reflect only the earliest record of the taxon in question, not its
entire geochronological range. M&B 00 refers to McKenna
and Bell (2000), the searchable version of McKenna and
Bell (1998). SALMA units are underlined.
Early Hemphillian, ca. 9 Ma
Pliometanastes (Xenarthra: Megalonychidae) is known
only from North America, but the subfamily
Megalonychinae is known from coeval deposits in
South America (Megalonychops) according to M&B
00. Morgan (2005) indicates that Pliometanastes has
its earliest occurrence in the McGehee Local Fauna of
Florida, of early Hemphillian (Hh 1) age, or at about
8–9 Ma.
Thinobadistes (Xenarthra: Mylodontidae) is only known
from North America. Its earliest occurrence is in
the McGehee Farm Local Fauna at ca. 8.5–9 Ma
(Morgan, 2005), but the tribe Thinobadistini is known
from approximately coeval Huayquerian deposits of
South America (M&B 00). Morgan (2005) indicates
that Thinobadistes is only distantly related to
Glossotherium. The >8 Ma age for the McGehee Farm
and Mixson’s Bone Bed sites advocated by Morgan
(2005) is older than in Tedford and others (2004; ca.
7.5–8 Ma). Morgan (2005) suggests an age of 8.0–8.5
Ma for the Mixson’s site, following Hulbert (2001).
Megalonyx (Xenarthra; Megalonychidae). This sloth is
endemic to North America (Morgan, 2005), and
apparently evolved from Pliometanastes. It is known
only from North and Central America (M&B 00).
Megalonyx curvidens is recorded in the Aphelops
Draw Fauna of early Hemphillian age (Hh 2; Tedford
et al., 1987:171L; Tedford et al., 2004; Fig. 6.2; ca. 7.2
Ma). Morgan (2005) shows Megalonyx present in the
Palmetto Fauna of Florida (ca. 5.2 Ma), comparable to
its 5.2 Ma age in the Pinole Local Fauna of California
(Hh 3; Tedford et al. [1987, 2004]). Flynn and others
(2005) report Megalonyx as occurring at about 4.8
Ma in the Guanajuato area of Mexico. Contrary to
Flynn and others (2005), the Mexican record does not
represent a pre-USA element of GABI.
Kraglievichia (Xenarthra, Pampatheriidae). Chasicoan to
Huayquerian of South America. Contrary to other
opinions, the North American records are dismissed
(Edmund, 1987; Scillato-Yané et al., 2005).
Huayquerian ca. 9.0–6.8 Ma
Flynn and Swisher (1995) indicate an age from 9.0–7.0 Ma.
Butler and others (1984) and Marshall and others (1992)
present evidence that the Huayquerian-Montehermosan
boundary is about 6.0 Ma. Cione and others (2001) summarize
evidence in favor of an upper limit of about 6.8 Ma. See
Cione and Tonni (2005), and discussion under Cyonasua.
*Cyonasua (Carnivora: Procyonidae). Cyonasua is known
from the Huayquerian to the Chapadmalalan of South
America (Cione and Tonni, 1995; Cione et al. 2001).
Chapadmalania is a closely related taxon that is known
from the Chapadmalalan to Vorohuan and perhaps
Montehermosan; the Huayquerian record is erroneous
(Cione and Tonni, 1996; Daniel Berman, 2006,
personal communication). The subfamily Procyoninae
is recorded from the early Miocene of North America
(M&B 00; Marshall et al., 1983). Butler and others
(1984) correlated northwestern Argintinian specimens
of Huayquerian age referred to Cyonasua as between
7.0 and 7.5 Ma. This appears to be the oldest known
record of carnivores in South America and the oldest
known representatives of the interchange in South
America. Fossils attributed to the more advanced
endemic genus Chapadmalania by Butler and others
(1984) and Marshall and others (1979) from Corral
Quemado, Cataqmarca, northwestern Argentina, are
correlated to an age of about 5.4–5.8 Ma. However,
the provenance is uncertain because the material was
“collected between units 15 and 32 (probably from
unit 21) of the Corral Quemado Formation” (Marshall
et al., 1979, p. 276). Units 15 to 32 have radioisotopic
dates that range from 6.7 Ma to less than 3.54 Ma.
Chapadmalania is certainly known in the type upper
Chapadmalalan in beds dated approximately as old as
3.3 Ma and in correlative beds in the Huayqueras de
San Carlos, Mendoza Province, Argentina (Cione and
Tonni, 1996; Schultz et al., 1998).
Late Hemphillian ca. 6.8–4.9 Ma and Montehermosan, ca.
6.8–5.0 Ma
Traditionally, Chapadmalalan and Montehermosan were
considered as separate units (Kraglievich, 1934; see
discussion in Cione and Tonni, 1995, 1996). However Pascual
and others (1965) incorporated the Chapadmalalan into the
Montehermosan, mixing the faunal lists. Marshall and others
(1984) recognized the Chapadmalalan as a separate unit,
but the faunal list was mixed and included taxa from units
of Huayquerian, Montehermosan, and Chapadmalalan age
(see Cione and Tonni, 1995). Reliable Chapadmalalan and
Montehermosan lists appear beginning in papers by Tonni
and others (1992a, b), Cione and Tonni (1995, 1996, 1999,
2005), Deschamps (2004). Marshall and Sempere (1993)
combined the Chapadmalalan and “Uquían” in a single unit;
this view is not followed here.
The upper boundary of the Chapadmalalan is at
about 3.3 Ma (Schultz et al., 1998). However the age of
the lower boundary of the Chapadmalalan is unknown as is
the duration of the Montehermosan. The 4.0 Ma age for the
Chapadmalalan-Montehermosan boundary shown in Cione
and Tonni (2005) is conventional. Actually little is known of
the ages within the interval from about 7.0 to 4.0 Ma. It is highly
probable that Chapadmalalan faunas could be older than 4.0
93
Ma, with paleomagnetic evidence suggesting a 4 Ma age for
the base of the upper Chapadmalalan (Orgeira and Valencio,
1984). Bolivian sediments with mammals were considered
to be Chapadmalalan sensu lato by Cione and Tonni (1996),
and could be as old as 5.4 Ma according to radioisotopic
ages obtained from underlying tuffaceous beds (Cione and
Tonni, 1996). On this basis the lower Chapadmalan should
be older. In Puerta de Corral Quemado, Catamarca Province,
Argentina, a specimen of Chapadmalania, Chapadmalalan
genus, “probably” comes from a level dated at 5.4–5.8 Ma
(Butler et al., 1984, p. 634). The 5.0 Ma age chosen in Figure
2 reflects these uncertainties.
Furthermore, Montehermosan taxa derive only from
the type of that unit near Pehuencó, Argentina, where it is
about 2-m thick and underlies the type Chapadmalalan. All
other sections considered to be Montehermosan in age are
doubtful or appear to be Chapadmalalan (Cione and Tonni,
1996).
*Sigmodontine rodents. According to Baskin (1986)
Abelmoschomys, from the latest Clarendonian Love
Bone Bed of Florida, is an early sigmodontine rodent
from the USA. Antecalomys (Korth, 1998) is a
contemporaneous form from Nebraska. McKenna and
Bell (1998) include Abelmoschomys in the subfamily
Cricetodontinae, however. Baskin (1978) described
the late Hemphillian Bensonomys from Arizona and
placed it as a subgenus of the living South American
genus, Calomys. Bensonomys also occurs in the late
Hemphillian Yepómera (Lindsay and Jacobs, 1985)
and El Ocote (Carranza-Castañeda and Walton,
1992) faunas of Mexico. Other North American
sigmodontines include Repomys (late Hemphillian to
late Blancan; Tedford et al., 1987), and Jacobsomys,
early Blancan Verde Fauna, Arizona (Czaplewski,
1987). Jacobsomys is tentatively assigned to the
sigmodontines by McKenna and Bell (1998).
Baskin (1986) summarizes evidence in favor of the
origin and diversification of sigmodontine rodents in
North America prior to their dispersal to South America
in the Montehermosan at about 3.5 Ma (modified here
to about 5.7 Ma; Figure 2). Sigmodontine records are
extremely rare in the Montehermosan (Pardiñas and
Tonni, 1998). Remarkably, all South American cricetids
pertain to endemic genera. Reig (1981) proposed that
cricetids entered South America in the late Miocene.
New evidence from the Cerro Azul Formation in
La Pampa Province (central Argentina) could bring
2006
94
new insights regarding the problem (Montalvo et al.,
2000).
The South American genus Calomys is not a synonym
of Bensonomys according to most recent authors
(Pardiñas et al., 2002), who consider Bensonomys to
be a peromyscine, not a sigmodontine. Recognizing
that the North American genera Abelmoschomys and
Bensonomys are older than the oldest-known South
American sigmodontines, it is important to establish
their phyletic affinities (Pardiñas et al., 2002: 240).
*Auliscomys (Rodentia, Sigmodontinae). Cione and
Tonni (2005) indicate this genus is characteristic of
the Montehermosan Trigodon gaudryi biozone in
Argentina (see also Pardiñas and Tonni, 1998; M&B
00. See also Necromys.
Necromys (Rodentia, Sigmodontinae). Cione and Tonni
(2005) indicate this genus is characteristic of the
Montehermosan Trigodon gaudryi biozone of
Argentina. It was cited as Bolomys by Vrba (1993) and
M&B 00. Bolomys is currently considered as a junior
synonym of Necromys (Wilson and Reeder, 2005).
e
Early Blancan (Bl1, ca. 4.8–3.5 Ma), and Chapadmalalan,
ca. 5.0–3.3 Ma
See the discussion above regarding the Montehermosan.
As indicated in Figure 2, the Chapadmalalan is considered
to range in age from about 5.0 to 3.3 Ma, but its base is
not calibrated. An “escorias” bed near the top of the upper
Chapadmalalan near Mar del Plata, Buenos Aires Province,
Argentina, is dated about 3.3 Ma (Schultz et al., 1998).
Cione and Tonni (1995) indicate that the Chapadmalalan is
pre-Uquían in the sense that the Uquían is replaced by the
Marplatan, which begins about 3.2 Ma in their correlation
(Cione et al., 2001).
Glossotherium (Xenarthra: Mylodontidae). Glossotherium
is known in South America from the type upper
Chapadmalalan, Buenos Aires Province, Argentina,
to uppermost Lujanian in different locations in the
Pampean region, according to Cione and Tonni
(1995; but not Cione and Tonni, 2005), or from levels
somewhat older than 3.3 Ma to 0.01 Ma. The genus
Glossotherium Kraglievich, 1934 is a junior synonym
of Glossotherium Owen, 1839. In North America
Glossotherium typically is found in the Blancan.
According to Flynn and others (2005) Glossotherium
occurs in early Blancan (4.7–4.8 Ma) deposits in
Mexico, but not until the late Blancan, about 2.8 Ma
(Lower Cita Canyon and Blanco faunas; Bell et al.,
2004:268) in the United States. Morgan (2005) shows
Glossotherium as present in the late Blancan Macasphalt
Shell Pit, Kissimmee River and Haile 15A local faunas
in Florida considered by him to be at about 2.2–2.7
Ma. Tomida (1987) reported Glossotherium from the
11 Mile Wash Local Fauna in Arizona, correlated to
about 3.5 Ma (revised to about 3.0 Ma in Bell et al.,
2004; Country Club fauna).
Esteban (1996) considers that all North American
specimens referred to Glossotherium (including those
of the Hemphillian of Mexico) are referable to the genus,
Paramylodon. According to this study, Glossotherium
should be an endemic South American genus. Thus
Esteban (1996) considers that the specimens identified
as Glossotherium chapadmalensis from North America
do not belong to this genus and/or species. This Ph. D.
dissertation remains unpublished, however.
Plaina (Xenarthra: Pampatheriidae). Scillato-Yané and others
(2005) consider Plaina a valid genus. Morgan (2005)
indicates that Carranza-Castañeda and Miller (2004)
have distinguished as Plaina material from Mexico
formerly referred to Holmesina or Pampatherium.
Flynn and others (2005) record Plaina from the
Guanajuato district in late Hemphillian faunas at about
4.6–4.7 Ma. Cione and Tonni (1996, 2005) indicate
its oldest South American record is in the type early
Chapadmalalan near Pehuencó, southern Buenos Aires
Province, Argentina (Neocavia depresidens biozone;
about 4 Ma or older; Figure 2), and the younger beds
of the type late Chapadmalalan near Mar del Plata,
both in southern Buenos Aires Province, Argentina
(Paraglyptodon chapadmalansis biozone; somewhat
older than 3.3 Ma). It also occurs in Chapadmalalan
beds of the Tunuyán Formation, Mendoza Province,
Argentina, and at Inchasi, Bolivia (Cione and Tonni,
1996).
Pampatherium and Holmesina (Xenarthra: Dasypdidae).
M&B 00 consider Pampatherium as a junior synonym
of Holmesina which Morgan and Hulbert (1995) show
as having a first appearance in the Santa Fe River Local
Fauna correlated as about 2.4 Ma and late Blancan.
Scillato-Yané and others (2005) suggest that both
Pampatherium and Holmesina are valid genera, the
latter probably derived in North America from Plaina
or Kraglievichia and dispersed to South America in the
latest Pleistocene (Lujanian; Cione and Tonni, 1999).
Flynn and others (2005) consider Pampatherium as a
synonym of Kraglievichia. Kraglievichia is restricted
to South America according to Edmund (1987). Flynn
and others (2005) may believe that Kraglievichia
pertains to Holmesina floridanus. Edmund (1987) and
Scillato-Yané and others (2005) retain H. floridanus as
separate from Kraglievichia.
Pampatherium is a South American taxon with an
Ensenadan oldest record. It probably descended from
Vassallia and dispersed to North America during the
Pleistocene (Scillato-Yané et al., 2005). The only
certain North American Pampatherium appears
to be P. mexicanum from the Pleistocene beds of
Puebla, Hidalgo, and Jalisco, Mexico (Scillato-Yané
et al., 2005). However, Carranza-Castañeda (2006,
personal communication) indicates that scutes closely
resembling those of Pampatherium occur at Arroyo
Belén, Guanajuato, in early Blancan beds dated at
about 3.6–3.9 Ma. Pampatherium may have dispersed
to North America in the late Blancan.
*eChapadmalania (Carnivora: Procyonidae). Chapadmalania
is a descendant of Cyonasua. Chapadmalania appears
to be exclusive to the Chapadmalalan (Cione and
Tonni, 1995).
*eAkodon (Rodentia, Sigmodontinae). This genus ranges
from the late Chapadmalan to Recent, following Tonni
and others (1992) and Cione and Tonni (1995). Akodon
apparently is an endemic taxon. M&B 00 agree. Vrba
(1993) lists the genus as early Chapadmalalan.
*eDankomys (Rodentia, Sigmodontinae). This genus ranges
from Chapadmalalan to Vorohuean (Tonni et al., 1992;
Cione and Tonni, 1995). M&B 00 extend the range to
Sanandresian. Vrba (1993) lists the endemic genus as
early Chapadmalalan.
*eGraomys (Rodentia, Sigmodontinae). This endemic genus
ranges from the late Chapadmalalan to Sanandresian
(M&B 00; Cione and Tonni, 1995, 2005). Dankomys,
Graomys, Reithrodon, and Zygodontomys were included
incorrectly in the early Chapadmalalan by Vrba (1993).
In Table 1 a late Chapadmalalan age is shown.
95
*eReithrodon (Rodentia, Sigmodontinae). Reithrodon
ranges from Chapadmalalan to Recent (Tonni et al.,
1992; Cione and Tonni, 1995; M&B 00). This is an
endemic genus. Ptyssophorus (in Vrba, 1993) is a
subgenus of Reithrodon in M&B 00, but is considered
a junior synonym of Reithrodon in Pardiñas and others
(2002).
*eZygodontomys (Rodentia: Sigmodontinae). The taxon
ranges from Pleistocene (only in Aruba) to Recent
of South America, and Recent of Central America
according to M&B 00.
*eArgyrohyus (Artiodactyla: Tayassuidae). M&B 00 list
this endemic genus as Chapadmalalan, consistent
with Cione and Tonni (1995). It first occurs in the
late Chapadmalalan Paraglyptodon chapadmalensis
biozone of Cione and Tonni (2005), correlated as about
3.3 Ma or somewhat older.
*Platygonus (Artiodactyla: Tayassuidae). Platygonus occurs
in the upper Chapadmalalan to Ensenadan according
to Cione and Tonni (2005). It is present for the first
time, along with Argyrohyus in the late Chapadmalalan
(see above). Tedford and others (2004) indicate the
genus is present in Hh2 faunas of North America at
about 7.2 Ma.
Marplatan, ca. 3.2–2.0 Ma
The chronology follows Cione and others (2001) and
Cione and Tonni (1995, 2005) and effectively replaces
the Uquían SALMA, which Flynn and Swisher (1995)
correlate to about 3.0–1.5 Ma. The Marplatan is composed
of three subages, the Barrancalobian (about 3.2–3.0 Ma),
Vorohuean (about 3.0–2.4 Ma) and Sanandresian (about
2.4–2.0 Ma).
*Dusicyon (Carnivora: Canidae). Berta (1987; see also
Prevosti et al., 2005) indicates that this genus (as
Pseudalopex) ranges from Uquían to Recent in South
America, with Uquían here refined to Marplatan
(Vorohuean), and correlated to about 2.5 Ma in
Cione and Tonni (2001). Vorohuean is represented
by the Akodon (Akodon) lorenzinii biozone in Cione
and Tonni (2005). The reference to Canis in the
Marplatan by Tonni and others (1992) corresponds
to Dusicyon.
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96
*eStipanicicia (Carnivora: Mustelidae). This endemic genus
is known in Sanandresian and Ensenadan deposits
according to Cione and Tonni (1995), followed here.
*eGalictis (Carnivora: Muselidae). M&B 00 show it as from
the Recent of North, Central and South America,
with a probable record in North and South America
(Ensenadan) into the early Pleistocene. Cione and
Tonni (1995) show it in the Vorohuean to Recent. The
Vorohuean is the oldest record of the genus in South
America (a Chapadmalalan record was considered as
highly dubious by Cione and Tonni, 1995).
Scapteromys (Rodentia: Sigmodontinae). M&B 00 indicate
an Uquían to Recent range for this endemic genus.
Vrba (1993) lists it as “Uquían.” Tonni and others
(1992) and Cione and Tonni (1995) show a Vorohuean
to Recent range, followed here.
e
*Lama (Artiodactyla: Camelidae). Cione and Tonni (1995,
2005) indicate a Marplatan (Barrancalobian) to Recent
age for this immigrant genus of a Holarctic family. The
genus Vicugna is shown as Ensenadan to Recent in
M&B 00, and as Ensenadan in Vrba (1993). However
it is included within Lama in Marshall and others
(1983) and Cione and Tonni (2005).
*Hippidion (Perissodactyla: Equidae). Alberdi and Prado
(1992, 1993) consider Onohippidium as a junior
synonym of Hippidion. Other authors consider
it a valid genus (MacFadden and Skinner, 1997;
MacFadden, 1997). Cione and others (2003) list the
immigrant equid as Marplatan to uppermost Lujanian.
MacFadden and Skinner (1979) record the genus in
the early Hemphillian of Texas (ca. 9 Ma) as well
as in the Irvingtonian of California (Tedford et al.,
2004).
*Onohippidium (Perissodactyla: Equidae). Specimens
assigned to Onohippidium in Vorohuean beds near Mar
del Plata, Buenos Aires Province, Argentina, represent
the oldest record of Equidae in South America (see
Reig, 1957; Tonni and Cione, 2000; Cione and Tonni,
2005). Prado and others (1998) mistakenly proposed
that the oldest material of Equidae in South America
came from Sanandresian beds in the Uquía Formation,
in Jujuy, northwestern Argentina (see Tonni and Cione,
2000).
Erethizon (Rodentia: Erethizontidae). The genus is known
from the Wolf Ranch Local Fauna of Arizona (Bell
et al., 2004), at about 2.7 Ma. Other occurrences
cited in Bell and others (2004) are younger than that.
M&B 00 show its range from late Pliocene to Recent.
Morgan (2005) indicates that the oldest known record
of Erethizon dorsatum is from the site Haile 16A,
correlated at about 1.6 Ma, but other (extinct) species
in Florida are late Blancan, ca. 2.2–2.7 Ma, compatible
with the Wolf Ranch record.
Reguero and others (2006) reported the occurrence
of Erethizon from Sanandresian beds of the Uquía
Formation of northwestern Argentina. This is the only
record of the North American genus in South America,
and might reflect a dispersal about 2.0 Ma, but also
could imply an earlier record in South America with
dispersal northward.
Late Blancan ca. 3.0–1.8 Ma
Glyptotherium (Xenarthra: Glyptodontidae). M&B 00
indicate a range of late Blancan to Rancholabrean for
this Central and North American genus, consistent
with Bell and others (2004) and Morgan (2005), who
note that Glyptotherium appears about 2.5 Ma in
the Upper Cita Canyon Local Fauna and in the 111
Ranch fauna of Arizona. The first occurrence of about
3.6 Ma in Figure 7.2 (Bell et al., 2004) apparently
is a drafting error (E. Lundelius, 2-17-06, personal
communication). Flynn and others (2005) indicate
that Glyptotherium first appears at about 2.8 Ma in
the Guanajuato district of Mexico. Morgan (2005)
notes the presence of Glyptotherium in early Blancan
faunas of North America (nominally the USA) at about
2.5–2.7 Ma. At the level of resolution, the geochron of
Glyptotherium is the same in Mexico as in the USA.
Kraglievichia (Xenarthra: Pampatheriidae). Although it has
been cited for North America, it is not present in this
region (see comments on Pampatherium, above).
Holmesina (Xenarthra: Pampatheriidae). This genus is
known from deposits in Florida at ca. 2.4 Ma (Bell
et al., 2004). Morgan (2005) cites the presence of
Holmesina in the Macasphalt Shell Pit and Haile 15A
local faunas correlated by him at about 2.2–2.7 Ma
(Haile 15A is correlated at 2.3 Ma). See comments on
Pampatherium, above.
Dasypus (Xenarthra, Dasypodidae). Species of Dasypus
inhabit South and North America today. This genus
is known from deposits in Florida at ca. 2.4 Ma (Bell
et al., 2004). Morgan (2005) cites the presence of
Dasypus bellus in the Macasphalt Shell Pit and Haile
15A local faunas correlated by him at about 2.2–2.7
Ma (Haile 15A is correlated as 2.3 Ma). If this generic
assignment is correct, Dasypus would be older in North
America than in South America. The species D. bellus
actually should be referred to Propraopus (Alfredo
Carlini, 2006, personal communication). Propraopus
is known from the Ensenadan (Cione and Toni, 2005),
but it might also occur in Chapadmalalan or Marplatan
strata (Ameghino, 1908).
Pachyarmatherium (Xenarthra: Glyptodontidae). Morgan
(2005) records this apparently immigrant glyptodont
in the Kissimmee River, Haile 16A and Leisey Shell
Pit local faunas at about 2.4–1.4 Ma in Florida.
Paramylodon (Xenarthra: Mylodontidae). Morgan (2005)
records this North American taxon (P. harlani) from
faunas at about 1.8 Ma in Florida (Inglis 1A, 1C, Haile
16A, Leisey Shell Pit, Haile 21A).
Eremotherium (Xenarthra: Megatheriidae). Bell and others
(2004) indicate that Eremotherium occurs in Florida
faunas at about 1.77 Ma (De Soto Shell Pit; Inglis 1A).
Morgan (2005) concurs in the late Blancan age, but
apparently considers it to be somewhat older, from The
Kissimmee River Local Fauna (2.2–2.7 Ma).
Neochoerus (Rodentia: Hydrochoeridae). The genus is
known from the 111 Ranch Local Fauna in Arizona
(Bell et al., 2004), about 2.6 Ma. Morgan (2005) cites
the presence of Neochoerus in the Macasphalt Shell Pit
Local Fauna correlated by him at about 2.2–2.6 Ma.
Flynn and others (2005) indicate that Neochoerus is
recorded in Mexican deposits at about 3.3 Ma. In South
America Neochoerus is known from the Ensenadan
and Lujanian (Deschamps, 1998; Cione and Tonni,
1999, 2005). It is tempting to consider a Neotropical
source for the South American record, although it may
turn up in the Marplatan.
Hydrochoerus (Rodentia: Hydrochoeridae). The oldest
record of this rodent is from the late Blancan Haile
15A site, Florida, by Morgan and Hulbert (1995). This
97
is correlated as about 2.2 Ma by Bell and others (2004)
and about 2.3 Ma by Morgan (2005).
Tremarctos (Carnivora; Ursidae). This genus is known
from two species, T. floridanus (Gidley) from the late
Pliocene and Pleistocene of North America, and T.
ornatus in the Recent of South America, where it is
the only living tremarctine. It has not been found as a
fossil in South America.
Ensenadan, ca. 2.0–0.7 Ma
This follows Cione and others (2001). Flynn and Swisher
(1995) correlate the Ensenadan from about 1.2 to 0.7 Ma.
MacFadden (2000) indicates an age for the Ensenadan/
Lujanian boundary as 0.7–0.6 Ma. This does not include
the Bonaerian (e.g., Cione and Tonni, 1999, 2001, 2005)
which relegates the Lujanian to very late Pleistocene. The
Bonaerian is ca. 0.7–0.125 Ma, whereas Lujanian ranges
from 0.125–0.08 Ma (Cione et al., 2003; Verzi et al., 2004).
Many taxonomic references to the Lujanian in prior literature
use the Lujanian as the interval next after the Ensenadan.
*Chrysocyon (Carnivora: Canidae). Berta (1987) lists this
genus as Ensenadan to Recent. However there are only
a few fossils from the Tarija Formation of Bolivia of this
age. Now known as the Tolomosa Formation, the unit
is composed of two parts. The lower interval consists
of poorly fossiliferous clays, and the upper, highly
fossiliferous, interval is composed of conglomerates.
The basal age of the lower unit is about 1 Ma and could
be regarded as Ensenadan on that basis, but not on its
fossil content. The upper unit is younger than 0.78 Ma
(Soibelzon et al., 2005). Chrysocyon was not listed in
Tonni and others (1992) or Cione and Tonni (1995)
because it is not present in southern South America
until late Platan (Prevosti et al., 2004). Berta (1987)
groups this genus with ‘true’ wolves of South America:
Canis gezi, Canis nehringi and Canis dirus (see
comments below). Chrysocyon is also known from the
early Blancan of North America, so is an immigrant to
South America.
*Cerdocyon (Carnivora: Canidae). Marshall and others (1983)
show this apparently immigrant taxon is Ensenadan
and Lujanian to Recent. M&B 00 indicate it has a
late Pleistocene to Recent record in South America,
but also a late Miocene to early Pliocene record in
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98
North America (USA, Mexico; late Hemphillian and
early Blancan, respectively; Torres and Ferrusquía,
1981; Berta, 1987). R.H. Tedford (2-21-06, personal
communication) indicates that the USA record of
Cerdocyon is from the late Hemphillian fauna of
Rentfro Pit 1, Ogallala Grp., near Channing, Texas.
Torres and Ferrusquía (1981) consider the Mexican
species closest to the South American C. thous. The
genus apparently arose in North America. It is not
listed in Tonni and others (1992) or Cione and Tonni
(1995) because the genus is not recorded in southern
South America.
*eTheriodictis (Carnivora: Canidae). Vrba (1993) indicates
an Ensenadan age as do Cione and Tonni (1995) for
Theriodictis. The species Canis gezi actually is a
junior synonym of Theriodictis platensis (Prevosti,
2001). Berta (1987) records Theriodictis from the
Lujanian of Ecuador, but Prevosti (2004) considers
that this material probably is assignable to Protocyon
troglodytes.
*Protocyon (Carnivora: Canidae). Tonni and others (1992)
and Cione and Tonni (1995) show the genus as present
in the Ensenadan and Lujanian. “Uquían” references
(Vrba, 1993) are erroneous.
*?Conepatus (Carnivora: Mustelidae). This genus is known
from Ensenadan to Recent deposits in South America,
according to Cione and Tonni (1995), who rejected
an early Chapadmalalan age (Vrba, 1993). In any
case, the genus is known only from Irvingtonian and
younger times in North America (e.g., M&B 00, Bell
et al., 2004). At present, it appears that the northern
and southern first occurrences might be coeval at the
available level of resolution.
extinct during the early Holocene, approximately
synchronous with their extinction in North America
(Soibelzon et al., 2005).
*Smilodon (Carnivora: Felidae). Tonni and others (1992)
and Cione and Tonni (1995) show that the genus
ranges from Ensenadan to Lujanian. Smilodontidion
is a junior synonym of Smilodon based on juvenile
material (Cione and Tonni, 1995). It was referred to
Vorohuean (M&B 00), but this is an error.
* “Felis” (Carnivora: Felidae). Vrba (1993) records this
immigrant genus from the Ensenadan. Tonni and
others (1992) and Cione and Tonni (1995) show it
from Ensenadan and Lujanian to Recent. However,
the only certain reference to Felis in South America is
based on Felis cattus from latest Holocene sediments
of Buenos Aires Province, Argentina, a probable
result of Hispanic immigration (Prevosti, in press).
“Felis” vorohuensis and “Felis” sp., both of uncertain
systematic position, occur in the Ensenadan of Buenos
Aires Province, Argentina (Prevosti, in press). Tedford
and others (1987, 2004) record Felis rexroadensis from
the Hh3 Upper Bone Valley Local Fauna, Florida.
*Felis (Carnivora: Felidae). Vrba (1993) records the genus
from the Ensenadan. Tonni and others (1992) and
Cione and Tonni (1995) show it from the Ensenadan
and Lujanian to Recent. This is consistent with M&B
00. It also is listed by M&B 00 from the early Pliocene
(= early Blancan).
*Puma (Carnivora: Felidae). The Puma is known from
the Ensenadan in southern Buenos Aires Province,
Argentina (Marshall et al., 1984; Pomi and Prevosti,
2005).
*eLyncodon (Carnivora: Mustelidae). This endemic genus is
known from the Ensenadan of Buenos Aires Province,
Argentina, to Recent (Cione and Tonni, 2005).
*eLynchailurus (Carnivora: Felidae). This South American
genus is known from Ensenadan beds at Camet,
Buenos Aires Province, Argentina (Prevosti, in press).
*eArctotherium (Carnivora: Ursidae). The Tremarctinae in
North America range from Hemphillian (Plionarctos
sp., Hh 3, Tedford et al., 2004) to Rancholabrean (P.
floridanus and Arctodus simus). In South America
the first record corresponds to the Ensenadan
(Arctotherium latidens) and continues to the present as
Tremarctos ornatus, but the short-faced bears become
*eCalomys (Rodentia: Sigmodontinae). See comments on
Sigmodontinae. M&B 00 cite Marshall and others
(1984) for an Ensenadan-Lujanian duration of this
endemic genus. Vrba (1993) cites an “Uquían” age.
It does not occur in the Marplatan but is known
from the Ensenadan according to Cione and Tonni
(1995). Carranza-Castañeda and Walton (1992) record
Calomys (Bensonomys) from the late Hemphillian El
Ocote Local Fauna.
*eLundomys, Oxymycterus, Phyllotis, Holochilus (Rodentia:
Sigmodontinae). All these genera occur in the
Mesotherium cristatum biozone (Ensenadan) to Recent
(Pardiñas, 1993, 2004).
*eCatagonus (Artiodactyla: Tayassuidae). This endemic
peccary is listed as Ensenadan to Recent in Marshall
and others (1983), Vrba (1993), M& B00, and Cione
and Tonni (2001; 2005).
*Hemiauchenia (Artiodactyla: Camelidae). Tonni and others
(1992) and Cione and Tonni (1995) show an Ensenadan
to Lujanian age for this immigrant taxon. M&B 00
show a middle Miocene (early Clarendonian) to late
Pleistocene (Rancholabrean) record in North America,
late Pliocene to Recent in South America. Meachen
(2005) discusses Hemiauchenia species from Florida
(1.8–2.5 Ma) and their possible relationship to the
ancestry of lamines in South America.
*eVicugna (Artiodactyla: Camelidae). This endemic camel is
shown as Ensenadan to Recent in M&B 00, is listed as
Ensenadan in Vrba (1993), and included within Lama
in Marshall and others (1983) and Cione and Tonni
(2005).
*Palaeolama (Artiodactyla: Camelidae). This genus is
recorded in Inglis IC Fauna of Florida, correlated to
about 1.8 Ma (Bell et al., 2004; Morgan, 2005). The
genus ranges from the Ensenadan to Lujanian (Cione
and Tonni, 1995). It is not present in the Marplatan,
contra Marshall and others (1983). Nevertheless,
Morgan (2005) noted Paleolama as a native North
American genus. At this time it appears that the
northern and southern first occurrences might be
coeval at the level of resolution available. In any case
it is an immigrant to South America.
*eEpieuryceros (Artiodactyla, Cervidae). Churcher (1966)
considers Epieuryceros as a junior synonym of
Blastoceros (see also M&B 00). However, Bagnalasta
(1980) validates Epieuryceros as a distinct genus. It is
exclusive to the Ensenadan (Cione and Tonni, 2005).
It represents the first record of the Cervidae in South
America.
99
*eAntifer (Artiodactyla: Cervidae). Bagnalasta (1980)
considers Antifer as a valid endemic genus. It ranges
from the Ensenadan to Bonaerian (Cione and Tonni,
1999).
*Tapirus (Perissodactyla: Tapiridae). This genus is listed as
“Uquían” in Vrba (1993), and Ensenadan to Recent
in Tonni and others (1992), Cione and Tonni (1995,
2005) and M&B 00, who also point out that the genus
is represented in the late Miocene of Holarctica.
*Stegomastodon (Proboscidea: Gomphotheriidae). Cione
and Tonni (1995, 2005) report Stegomastodon from
the Ensenadan and Lujanian. Prado and others (2003)
synonymized Notiomastodon with Stegomastodon
(see Cione et al., 2005 and references therein). An
indeterminate gomphotheriid was reported from the
Sanandresian beds of the Uquía Formation in Jujuy
Province, Argentina (López et al., 2001). M&B 00
report Stegomastodon from the early to late Pleistocene
of South America and from the early Pliocene to late
Pleistocene of North America. Bell and others (2004)
indicate that Stegomastodon is limited to the Blancan
(ca. 3.5 Ma) and Irvingtonian (ca. 1.2 Ma) in North
America.
*Cuvieronius (Proboscidea: Gomphotheriidae). Marshall
and others (1983) report this genus from the Uquían to
Lujanian. It is not listed in Tonni and others (1992) or
Cione and Tonni (1995) from the Marplatan = Uquían.
Cuvieronius is not present in Argentina. Alberdi and
Prado (1995) report it from the late Pliocene or early
Pleistocene to late Pleistocene. Here it is listed as
Ensenadan to Lujanian (see MacFadden, 2000). This
genus is recorded in late Hemphillian/early Blancan
faunas of North America, ca. 4.9 Ma in age (Pañaca;
Bell et al., 2004), 3.0–3.6 Ma (Camp Rice Fm., New
Mexico), as well as late Blancan sites in Arizona,
Florida and Texas (Lambert and Shoshani, 1998). It
also occurs in late Pleistocene sites in Honduras, Costa
Rica and Mexico (Lucas et al., 1997; Jackson and
Fernández, 2005).
Irvingtonian, ca. 1.8–0.25 Ma
Didelphis (Marsupialia: Didelphidae). Vrba (1993) lists this
didelphid immigrant as early Irvingtonian. Bell and
others (2004) show an Irvingtonian and Rancholabrean
2006
100
to Recent range for Didelphis and indicate (p. 282) that
the genus first occurs in early Irvingtonian faunas, such
as Fyllan Cave, Texas, with a likely age of 0.78–1.77
Ma. Table 1 shows a correlated age of about 1.2 Ma.
Nothrotheriops (Xenarthra: Megatheriidae). Vrba (1993)
lists this member of an otherwise South American
megathere as early Irvingtonian, but Bell and others
(2004) indicate a latest Blancan age. Lundelius
and others (1987) cite an age of about 2.1 Ma
for Nothrotheriops in the Vallecito Creek L.F. of
California, corroborated by Bell and others (2004,
fig. 7.2). Morgan (2005) suggests that this record is of
Megalonyx, so the earliest record of Nothrotheriops is
ca. 1.4–1.6 Ma (Leisey Shell Pit, Florida).
Myrmecophaga (Xenarthra: Myrmecophagidae). The sole
occurrence of this genus is in the El Golfo Fauna
of Sonora (Bell et al., 2004). The association of
Megalonyx wheatleyi in this fauna suggests an age of
about 1.0 Ma based on typical occurrences of that sloth
elsewhere in North America.
Bonaerian, ca. 0.7–0.125 Ma
*eParaceros (Artiodactyla: Cervidae). Paraceros was
considered a synonym of Habromeryx (Cabrera, 1929)
and M&B 00 indicate that Habromeryx is a junior
synonym of Morenelaphus. However it is presently
considered a junior synonym of Paraceros (see
Ubilla and Perea, 1999; Cione and Tonni, 1999). It is
known from the Bonaerian of Buenos Aires Province,
Argentina (Cione and Tonni, 1999, 2005).
*eArctotherium (Carnivora: Ursidae). The tremarctines
Arctotherium vetustum, A. bonariense and A. tarijense
occur in the Bonaerian (Soibelzon et al., 2005),
according to whom the genus does not occur in North
America.
*eHerpailurus (Carnivora: Felidae). A probable specimen of
this genus comes from Bajo San José, southern Buenos
Aires Province, Argentina (Prevosti, in press).
*Pecari (Artiodactyla: Tayassuidae). This peccary ranges
from Lujanian to Recent in Marshall and others (1983),
with an ?Ensenadan record. It begins in the Ensenadan
per Vrba (1993). Cione and Tonni (2005) indicate that
Pecari tajacu (= Dicotyles; Woodburne, 1969) first
occurs in the Megatherium americanum biozone, or
Bonaerian SALMA, or about 0.6 Ma.
*Morenelephus (Artiodactyla: Cervidae). Morenelephus is
known from the Bonaerian Cione and Tonni (2005).
Webb and Rancy (1996) indicate that this name
includes Hippocamelus of previous literature, but
Adriana Menegaz (2005, personal communication)
notes that this is not the case.
Lujanian, ca. 0.125–0.08 Ma
This follows Cione and Tonni (1995). Cione and Tonni (2001,
2005) separate Bonaerian from Lujanian, with Lujanian
being late Pleistocene.
*Lutra (Carnivora: Mustelidae). The genus is immigrant in
the Ensenadan according to Vrba (1993) and extends
to the Recent (Marshall et al., 1983; M&B 00).
The first record of this genus in Argentina is in late
Lujanian beds of Buenos Aires Province, Argentina
(Tonni et al., 1985). Other records of the genus are in
the early Pliocene of Europe and Asia. Tedford and
others (2004) list the genus from the late Hemphillian
(Hh4) of North America. Lontra is a synonym of Lutra
according to McKenna and Bell (1998).
*ePteronura (Carnivora: Mustelidae). This endemic South
American genus of a Holarctic family occurs in
Lujanian beds of Entre Ríos Province, Argentina (see
Carlini et al., 2002).
*Nasua (Brachynasua) (Carnivora: Procyonidae). Marshall
and others (1983) record this immigrant genus as
Lujanian and ?Ensenadan to Recent. M&B 00 suggest
it is known from the early Pleistocene (= Ensenadan).
Vrba (1993) lists it as Ensenadan. M&B 00 list
Brachynasua as possibly from the early and middle
Pleistocene of South America. The genus is not present
in southern South America (Cione and Tonni, 1995).
Eira (Carnivora: Mustelidae). This genus is restricted to the
Pleistocene according to M&B 00. Marshall and others
(1983) list the genus as Lujanian to Recent.
*eDusicyon (Carnivora, Canidae). See comments on Dusicyon
under the Marplatan.
*eOncifelis and Leopardus (Carnivora: Felidae). These South
American genera are known from late Lujanian beds
of the Buenos Aires Province, Argentina (Prevosti, in
press).
*eOryzomys (Rodentia, Sigmodontinae). The range is
?Pleistocene to Recent of South America according to
M&B 00. It is apparently not found in the Marplatan;
not listed in Cione and Tonni (1995). Tonni and others
(1992) show Recent only. Vrba (1993) lists the genus
as early Chapadmalalan. This is an endemic genus.
*eEuneomys (Rodentia, Sigmodontinae). M&B 00 cite
Marshall and others (1983) for a Lujanian occurrence
of this endemic genus. Vrba (1993) indicates an
“Uquían” age. Tonni and others (1992) and Cione and
Tonni (1995) do not list this genus.
*eHippocamelus (Artiodactyla: Cervidae). M&B 00 suggest
the most likely age is Lujanian to Recent, as Tonni
and others (1992) also consider. Vrba (1993) lists this
endemic genus as “Uquían.” Webb and Rancy (1996)
offer that this taxon is misidentified, and should be
Morenelephus.
* Ozotoceros (Artiodactyla: Cervidae). Cioni and Tonni and
others (2005) indicate a Lujanian to Recent age. M&B
00 are inconclusive except for a Recent age. Vrba (1993)
lists the endemic taxon as “Uquían,” which is an error.
e
Sylvilagus (Lagomorpha: Leporidae. This immigrant rabbit
entered South America in the Lujanian.
*Canis (Carnivora: Canidae). The first certain record of
Canis in South America is C. dirus in indeterminate
Pleistocene beds from the northern part of the continent
(Berta, 1987). In southern South America, the first
known definitive Canis entered with Homo sapiens
in the latest Lujanian. Another species, C. nehringi,
known from indeterminate levels of latest Lujanian
to Platan age in northeastern Buenos Aires Province,
Argentina (Kraglievich, 1928), appears to be a junior
synonym of C. dirus (Francisco Prevosti, personal
101
communication). Lundelius and others (1987) note
the genus is present in late Hemphillian (Hh 3, ca. 5
Ma) and younger faunas of North America. See also
Munthe (1998). Carranza-Castañeda and Miller (2004)
list Canis from late Hemphillian faunas of Mexico, but
some of these (Guanajuato) are likely early Blancan
(ca. 4.8 Ma).
*Equus (Perissodactyla: Equidae). This immigrant equid
is listed as Ensenadan to Lujanian in Marshall and
others (1983) and M&B 00. Cione and Tonni (2001,
2005) indicate that equids first occur in the Vorhuean,
or about 2.7 Ma. Equus is recorded in the Lujanian
in southern South America (sensu Cione and Tonni,
1999, 2005). Specimens of Equus occur in Bolivia
in levels assigned to the Ensenadan (MacFadden et
al., 1983). However, the stratigraphic level with the
specimens, of normal magnetostratigraphic polarity,
does not include typical Ensenadan taxa. Alberdi and
Prado (1995: 299) mention the same species in the
middle Pleistocene beds in the Río Chiche. However,
these beds are currently assigned to the late Pleistocene
(Hoffstetter, 1986).
Homo (Primates: Hominidae). Man entered South America
in the latest Lujanian.
Recent
Several immigrants do not have a fossil record. Some of
them are actively expanding their geographic ranges.
Cryptotis (Soricomorpha: Sorididae).
Heteromys (Rodentia: Heteromyidae).
Orthogeomys (Rodentia: Geomyidae).
Microsciurus (Rodentia: Sciuridae).
Sciurillus (Rodentia: Sciuridae).
Sciurus (Rodentia: Sciuridae).
2006

Michael O. Woodburne1,* Alberto L. Cione2,**, and Eduardo P. Tonni2

Transcription

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