• TABLE OF CONTENTS
HIDE
 Front Cover
 Copyright
 Abstract
 Table of Contents
 Acknowledgement
 Methods and abbreviations
 Systematic paleontology
 Glossotheridium chapadmalense (Kraglievich),...
 Mylodontinae, indeterminant
 Proscelidodon patrius (Ameghino),...
 Plaina intermedia (Ameghino),...
 Plohophorus sp
 Paraglyptodon sp
 Order Rodentia Bowdich, 1821
 Chapalmatherium cf. saavedrai Hoffstetter...
 Order Litopterna Ameghino,...
 Order Notoungulata Roth, 1903
 Posnanskytherium desaguaderoi Liendo,...
 Posnanskytherium inchasense, new...
 Hypsitherium bolivianum, new genus...
 Discussion
 Summary and conclusions
 Literature cited
 Back Cover






Group Title: Bulletin of the Florida Museum of Natural History
Title: Pliocene mammals from Inchasi, Bolivia
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Full Citation
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Permanent Link: http://ufdc.ufl.edu/UF00095795/00001
 Material Information
Title: Pliocene mammals from Inchasi, Bolivia the endemic fauna just before the great American interchange
Series Title: Bulletin - Florida Museum of Natural History ; volume 39, number 3
Alternate Title: Pliocene mammals from Bolivia
Physical Description: p. 87-140 : ill. ; 23 cm.
Language: English
Creator: Anaya, Federico
MacFadden, Bruce J.
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1995
Copyright Date: 1995
 Subjects
Subject: Mammals, Fossil -- Bolivia   ( lcsh )
Paleontology -- Pliocene   ( lcsh )
Paleontology -- Bolivia   ( lcsh )
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
Spatial Coverage: Bolivia
 Notes
Bibliography: Includes bibliographical references (p. 138-140).
Language: Abstract also in Spanish.
General Note: Cover title.
Statement of Responsibility: Federico Anaya and Bruce J. MacFadden.
 Record Information
Bibliographic ID: UF00095795
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 34089935

Table of Contents
    Front Cover
        Page 85
    Copyright
        Page 86
    Abstract
        Page 87
    Table of Contents
        Page 88
    Acknowledgement
        Page 89
        Page 90
    Methods and abbreviations
        Page 91
    Systematic paleontology
        Page 92
        Page 93
    Glossotheridium chapadmalense (Kraglievich), 1925
        Page 94
        Page 95
        Page 96
        Page 97
    Mylodontinae, indeterminant
        Page 98
        Page 99
    Proscelidodon patrius (Ameghino), 1888
        Page 100
        Page 101
    Plaina intermedia (Ameghino), 1888
        Page 102
        Page 103
        Page 104
        Page 105
    Plohophorus sp
        Page 106
        Page 107
        Page 108
    Paraglyptodon sp
        Page 109
    Order Rodentia Bowdich, 1821
        Page 110
        Page 111
    Chapalmatherium cf. saavedrai Hoffstetter et al., 1984
        Page 112
    Order Litopterna Ameghino, 1889
        Page 113
        Page 114
        Page 115
        Page 116
        Page 117
        Page 118
        Page 119
    Order Notoungulata Roth, 1903
        Page 120
        Page 121
    Posnanskytherium desaguaderoi Liendo, 1943
        Page 122
        Page 123
        Page 124
        Page 125
        Page 126
    Posnanskytherium inchasense, new species
        Page 127
        Page 128
    Hypsitherium bolivianum, new genus and species
        Page 129
        Page 130
        Page 131
        Page 132
        Page 133
    Discussion
        Page 134
        Page 135
        Page 136
    Summary and conclusions
        Page 137
    Literature cited
        Page 138
        Page 139
        Page 140
        Page 141
    Back Cover
        Page 142
Full Text




iuiAmumuuu


of the



FLORIDA
MUSEUM OF
NATURAL HISTORY


PLIOCENE MAMMALS
FROM INCHASI, BOLIVIA:
THE ENDEMIC FAUNA JUST BEFORE
THE GREAT AMERICAN INTERCHANGE


Federico Anaya and Bruce J. MacFadden


Volume 39, No. 3, pp. 87-140


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PLIOCENE MAMMALS FROM INCHASI, BOLIVIA:
THE ENDEMIC FAUNA JUST BEFORE
THE GREAT AMERICAN INTERCHANGE



Federico Anayal and Bruce J. MacFadden2




ABSTRACT

Anew, high-elevation (3200 m) mammalian local fauna is described from Communidad Inchasi, ca. 50
km SE of Potosi, Bolivia. The fauna consists of 4 orders (Edentata, Rodentia, tLitopterna, and
tNotoungulata), 9 families, 10 genera, and at least 11 species. Of these, one new genus and two new species
of notoungulates are described. This fauna contains two genera endemic to the Bolivian altiplano,
tPosnanskytherium and tHypsitherium n. gen., whereas the remaining eight genera (tGlossotheridium,
tProscelidodon, tPlaina, tPlohophorus, tParaglyptodon, tChapalmatherium, tCaviodon, and
tPromacrauchenia) are referable to previously described genera from the Pliocene of Argentina.
The biochronology of the Inchasi mammals indicates either a: (1) Montehermosan/ Chapadmalalan
(undifferentiated) age because the included taxa are either restricted to the Montehermosan, Chapadmalalan,
or span both mammal ages; or (2) Chapadmalalan age based on recent biostratigraphic redefinitions from
classic localities in Argentina. Previous magneto- and biostratigraphic data indicate that the Inchasi beds and
their contained mammalian fauna are middle Pliocene with an age range of between 3.3 and 4.0 myr ago.
The Inchasi local fauna lacks any immigrant taxa and thus represents an endemic fauna that occurred about
1 myr before the beginning of the Great American Interchange.

RESUME

Una nueva fauna local de mamiferos es descripta para la Comunidad Inchasi, aproximadamente 50
kilometros al sudeste de Potosi, Bolivia, a una altura de 3200 m.s.n.m. Esta fauna comprende cuatro
6rdenes (Edentata, Rodentia, tLitoptera y tNotoungulata), nueve families, 10 g6neros y por lo menos 11
species. Se describe un g6nero nuevo y dos nuevas species. Esta fauna tiene dos g6neros endmnicos del
altiplano Boliviano, tPosnanskytherium y tHypsitherium gen. n. y los ocho restantes (tGlossotheridium,
tProscelidodon, tPlaina, tPlohophorus, tParaglyptodon, tChapalmatherium, tCaviodon y
tPromacrauchenia) pertenecen a g6neros del Plioceno de Argentina.
Hay dos posibilidades para interpreter la biocronologia de los mamiferos de Inchasi: o (1) los f6siles
guia indican una edad Montehermosense/Chapadmalalense (no diferenciada) debido a que los taxones estan
restringidos al Montehermosense, Chapadmalalense, o existieron durante las dos edades juntas; o (2) indican
solo una edad Chapadmalalense en base a las investigaciones recientes en Argentina. Estudios previous de la
magneto- y bioestratigrafia indican que los afloramientos de Inchasi pertenecen al Plioceno medio, con una
edad entire 3.3 y 4 millones de aflos. La mastofauna de Inchasi carece de inmigrantes norteamericanos y


SThe senior author is Jefe, Departamento de Paleontologia, Museo Nacional de Historia Natural, Casilla 8706, Cota Cota, La Paz, Bolivia,
and Courtesy Curator, Department of Natural Sciences, Florida Museum of Natural History, University of Florida, P. O. Box I117800,
Gainesvile FL 32611-7800, U. S. A..
The junior author is Curator of Vertebrate Paleontology, Department of Natural Sciences, Florida Museum of Natural History, University of
Florida, P. O. Box 117800, Gainesville FL 32611-7800, U. S. A., and Research Associate, Departamento de Paleontologia, Museo Nacional
de Historia Natural, La Paz, Bolivia.

ANAYA F. and B. J. MACFADDEN. 1995. Pliocene mammals from Inchasi, Bolivia: The endemic fauna
just before the Great American Interchange. Bull. Florida Mus. Nat Hist. 39(3):87-140.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


represents un intervalo aproximadamente de 1 mill6n de aflos antes del comienzo del Gran Intercambio
Americano.

TABLE OF CONTENTS

Introduction .......................................................................................................... 88
Acknowledgements ... .................... ............. ....................... 89
Geological Setting and Previous Investigations.................................................................................... 89
Methods and Abbreviations ....................... ... ............. 91
System atic Paleontology ............................................ ............................................................ ......... 92
Order Edentata Cuvier, 1798
tMegatheriinae, indeterminant ................. .... ................................ .... .. 92
SGlossotheridium chapadmalense (Kraglievich), 1925 ........... ............. .... ....... 94
M ylodontinae, indete nant..... ................ ... .................................... 98
Proscelidodonparius (Ameghino), 1888 ..... .............................. .......... ... ...... ...... 100
tPlaina intermedia (Ameghino), 1888 ......... ............. ......... .............. ... ... 102
Plohophorus sp..................................................................... ................................ 106
tParaglyptodon sp ..... .............................................. 109
Order Rodentia Bowdich, 1821
Caviodon sp.................. ............................................ ........ 110
tChapalmatherium cf. saavedrai Hoffstetter et al., 1984........................ ................................ 112
tOrder Litoptema Ameghino, 1889
Prom acrauchenia sp................................................................. ............................................... 113
tOrder Notoungulata Roth, 1903..... .................... ............................ 120
tPosnanskytherium desaguaderoi Liendo, 1943........................................................................ 122
tPosnanskytherium inchasense, new species............... ........ ....................... 127
tHypsitherium bolivianum, new genus and species.......................... ................. .. 129
D iscussion.................. .............. .............................. 134
Faunal composition, diversity, and alpha-level taxonomy........................... ..... 134
Paleoecology and paleoenvironmental reconstructions.................... ............. 135
Biogeography, Pliocene regional and continental correlations,
and the problematic Montehermosan/Chapadmalalan interval................................................. 136
Summary and Conclusions........... .......................................... .............................................. 137
Literature Cited.................. ..... ................................. ..... .... ............... 38

INTRODUCTION


South America drifted as an island continent during most of the Cenozoic,
carrying along with it an endemic biota. This isolation changed with the formation
of the Panamanian Land Bridge and the consequent dry-land connection that
allowed dispersal of many land mammals during the Great American Interchange
(e.g. Stehli and Webb 1985). The main phase of this interchange began during the
late Pliocene, about 2.5 myr ago, and continued through the Pleistocene until about
10,000 years ago when an extinction wave decimated the diversity of medium- to
large-bodied mammals throughout the Americas.
The fossil record of late Pliocene and Pleistocene land mammals in South
America is very rich, with many localities recorded from numerous regions on that
continent (Marshall et al. 1983; 1984). In contrast, relatively little is known of the






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


endemic mammalian faunas during the late Miocene and early Pliocene just before
the interchange. Consequently, the paucity of these earlier faunas has led to
confusion about the biochronological characterization and distinctness of the
Montehermosan and Chapadmalalan land mammal ages. Several Pliocene, pre-
interchange mammal localities have been reported from Bolivia, e.g. at Ayo Ayo-
Viscachani south of La Paz (Hoffstetter et al. 1971; Hoffstetter 1986), and although
certain taxa have been described in detail (e.g. Hoffstetter et al. 1984), in most
cases these important faunas have never been fully described. Recently a rich
Pliocene, pre-interchange mammalian fauna has been discovered at Inchasi, about
50 km south of Potosi. This fauna offers significant potential to: (1) better
understand the endemic mammals just before the Great American Interchange; (2)
provide a further understanding of the Montehermosan and Chapadmalalan land
mammal ages; and (3) further constrain the timing of the beginning of the Great
American Interchange.
The purpose of this study is to: (1) describe the mammalian fauna from Inchasi;
(2) compare this fauna with other faunas of similar age in Bolivia and Argentina;
(3) discuss the overall significance of this fauna with regard to faunal diversity,
paleoecology, and land-mammal chronology. This report represents an
amplification of a previous, preliminary description of this fauna (Anaya 1994).



ACKNOWLEDGEMENTS

The senior author gratefully acknowledges financial support from the Bryan Patterson Award of the
Society of Vertebrate Paleontology in 1992. These funds allowed for one field trip to Inchasi and resulted in
the collection of numerous important specimens that further increased the knowledge of this fauna. The
authors thank Dr. Jaime Argollo of La Paz for showing us the Inchasi basin and for allowing us to pursue
our studies there. Bemardino Mamani and Yuko Okamura assisted in the field and laboratory. Gary
Morgan's expertise in specimen identification is gratefully acknowledged. Yuko Okamura skillfully
photographed many of the specimens presented here or used to prepare the illustrations. Barbara Harmon
prepared the line drawings and Linda Chandler assisted in the final preparation of the figures. Malcolm
McKenna, Bruce Shockey, and Fred Thompson assisted with points about taxonomic nomenclature. The
thorough and careful reviews of this manuscript provided by C. S. Churcher, G. Deluliis, and J. J. Flynn are
greatly appreciated. This research was also partially supported by a U.S. NSF grant EAR-8716207 and
Fulbright Senior Research Fellowship to BJMacF. This is University of Florida Contribution to
Paleobiology number 453.

GEOLOGICAL SETTING AND PREVIOUS STUDIES

Communidad Inchasi is located 8.5 km northeast of Puna and 50 km southeast
of the city of Potosi in the Province of Linares, Department of Potosi, at an







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Cara Nacloal 1 50 000
PUNA -. \M 5A
S(pot-Huayquwl.n) I I Huy
Figure 1. A. General location of Communidad Inchasi, about 50 km southeast of Potosi, and specific location of measured
sections. B. Composite magnetostratigraphy of the Inchasi beds, absolute age constraints, and correlation to the Geomagnetic
Polarity Time Scale and South American land mammal ages (modified from MacFadden et al., 1993).






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


elevation of 3220 m. Pliocene fossil mammals have been collected from around
Inchasi between latitudes 190 41' 20" -- 190 44' 35" S and longitudes 650 26' 30" --
650 29' 22" W (Fig. la) and can be located on the Miculpaya 1:50,000 quadrangle
(Instituto Geogrifico Militar Hoja 6534 IV).
The Pliocene Inchasi beds were deposited into a small basin with a topographic
extent of about 30 km2. The flanks of the basin consist of either undifferentiated
Ordovician sediments or volcanic rocks of the prominent middle Miocene Kari
Kari caldera. The Inchasi beds consist primarily of high-energy fluviatile gravels,
fine grained-clays and silts, and several tuffaceous horizons of varied thicknesses.
The Inchasi beds were originally mapped as Quaternary Alluvium (Pareja et al.
1978). They were not known to contain fossils until the first specimens were
discovered by Jaime Argollo in 1984. In 1988 a visit to Inchasi by Argollo,
MacFadden, and Peter Friend yielded several fossil mammals indicating a probably
late Pliocene age for these sediments. In 1990 Anaya and MacFadden spent one
week at Inchasi measuring sections and collecting paleomagnetic samples and
fossil mammals. Since 1990 the Inchasi beds have been prospected for fossils by
Anaya and crews from the Museo Nacional de Historia Natural, La Paz. All fossils
collected from there can be precisely located stratigraphically within the four
principal sections that we measured (Fig. la). Although radioisotopic age
determinations (analyzed by Carl Swisher of the Berkeley Geochronology
Laboratory, California) on tuffaceous samples collected from within the Inchasi
beds indicated middle Miocene detrital contamination (apparently from the Kari
Kari volcanics), the magnetostratigraphic analysis yielded a calibration of the
Inchasi beds. The composite section at Inchasi is dominantly of reversed polarity
with two normal events (Fig. lb). Using this polarity pattern and mammalian
biochronology, the Inchasi beds and its contained fauna correlate to the late Gilbert
and early Gauss chrons with an age range of about 4.0 to 3.3 myr (MacFadden et
al. 1993). A minimum of 11 species of middle Pliocene mammals were reported in
that study. Since the work published in MacFadden et al. (1993), Anaya and
associates have greatly added to the number of specimens collected from Inchasi
and revealed that two new species are contained in that fauna, as are described
with the entire fauna below. The age of the Inchasi fauna with regard to the South
American land mammal ages is within the problematic Montehermosan/
Chapadmalalan interval (see discussion below).

METHODS AND ABBREVIATIONS

All specimens from Inchasi described in this report are conserved in the
Departamento de Paleontologia, Museo Nacional de Historia Natural, La Paz (Cota
Cota), Bolivia.
Although we were able to assign most of the material (except for some
postcranial remains) to genus with reference to previous descriptions from either
the classic material from Argentina or Bolivia, for several taxa we were unable to






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


assign material to a particular genus. This resulted from either or both of the
following situations: (1) our material was too poorly preserved for adequate
characterization of diagnostic characters; or (2) the particular taxon originally
described from Argentina consisted of several similar, relatively poorly
characterized species nomina. In addition to our new descriptions, another taxon
described below (Chapalmatherium cf. saavedrai) is almost certainly a new
species, but the available sample is too fragmentary for a formal taxonomic
description.
All measurements are presented in millimeters (mm) and were made using
calipers. Dental terminology follows the convention of upper case for upper teeth
(i.e. I, C, P, M) and lower case for lower teeth (i.e. i, c, p, m). The following
abbreviations are used in the text:
@; measurement approximate (usually because of fragmentary specimen).
CF/cf a caniniform tooth (upper/lower) of uncertain position and homology
relative to eutherian canines.
GB; Vertebrate Paleontology collection, Servicio Geol6gico de Bolivia, now
housed as part of the MNHN-Bol-V collection.
L; left side.
MF/mf; a molariform tooth (upper/lower) of uncertain position and
homology relative to eutherian premolars and molars.
MNHN-Bol-V; Vertebrate Paleontology collection, Museo Nacional de
Historia Natural, Bolivia,
R; right side.
Taxonomic nomenclature for higher categories follows Simpson (1945)
and Pascual (1967), although with some minor revisions proposed by
Mones (1994).

SYSTEMATIC PALEONTOLOGY

Class Mammalia Linnaeus, 1758
Order Edentata Cuvier, 1798
Infraorder Pilosa Flower, 1883
(Family Megatheridae, Owen, 1843
tMegatheriinae, indeterminate
Figure 2

Material.--MNHN-Bol-V 003354, R calcaneum.
Geographic and stratigraphic location.--350 m west of Communidad Inchasi,
17 m above the base of measured section 3B.
Description.--This large, robust calcaneum has a length of 240 mm. Dorsally,
it contains a robust tuber calcis with rough surface for tendinous and muscular
attachments (Fig. 2). The articular surface with the astragalus contains a






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 5

cm


Figure 2. Medial view of right calcaneum of Megatheriinae, indeterminate, MNHN-Bol-V 003354.




lateral sustentacular facet with maximum dimension of 67 mm and a medial,
convex ectal facet with a maximum dimension of 82 mm; these are distinctly
separated by a sulcus talis. The sustentacular facet of the calcaneum forms an
oblique angle with the cuboid articular surface. The main longitudinal sulcus in
the center of the ventral part of the calcaneal shaft is wide and contains vascular
foramina within its deeper part.
Discussion.--With the exception of Hoffstetter (1986; also see Marshall et al.
1983), who mentioned the presence of a possible ancestor (otherwise unspecified)
of Megatherium in the Pliocene of Bolivia, the specimen from Inchasi is the first
description of a pre-Pleistocene record of this subfamily in this country. Based on
our comparisons with the large collection of megatheriines from Tarija in the
MNHN-Bol, although of smaller size, the overall morphology and relative
proportions of the calcaneum from Inchasi make it distinctly referable to
Megatheriinae. However, without further, more diagnostic material, a more
precise taxonomic allocation cannot be made at present. The occurrence from
Inchasi extends the range of megatheres in Bolivia back into the middle Pliocene.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


tFamily Mylodontidae Ameghino, 1889
tSubfamily Mylodontinae Gill, 1872
tGenus Glossotheridium Kraglievich, 1934
tGlossotheridium chapadmalense (Kraglievich), 1925
Figures 3-5, Table 1

Material.--MNHN-Bol V 003358, partial mandible with Rcf, Rmfl-mf3,
Lmfl-mf3; MNHN-Bol V 003359, L partial juvenile ramus with cf, mfl-mf3 (Fig.
5); MNHN-Bol V 003371; mandible with Rcf, mf3, Lcf, Lmfl-mf3 (Figs. 3-4).
Geographic and stratigraphic location.--750 m east and 1200 m southwest of
Communidad Inchasi; stratigraphic position 56 m and 18 m, respectively, in
measured sections 2 and 4.
Description.--The mandibles are laterally (i.e. labiolingually) compressed and
have heavily fused symphyses (Fig. 3). The mandible is deepest ventral to mf3.
The symphysial region extends anterior to the cf. The anterolingual part of the
symphysis is relatively wide and the anteroventral region is spatulate. The dorsal
surface of the symphysis exhibits rugose bone containing small pits. The
anteroventral surface of the symphysis has a very well-developed convex crest in
the sagittal plane of the chin which becomes less prominent as it approaches the
dorsal surface. The mental foramina are large and well-developed, and one each is
located on the right and left sides of a well-developed ventral sagittal ridge of the
symphysis. In ventral view the anterior part of the mandible is very thin and forms
a thin crest extending posteriorly to below the mfl.
The teeth are hypselodont with a thick, central pulp cavity surrounded by
relatively thick dentine flanked by thinner orthodentine. The caniniforms are oval
to elliptical in cross-section and separated from the mfl by a small diastema with
an anteroposterior length of about 5 mm. In MNHN-Bol-V 003358 (not
illustrated) the cf is significantly more robust than in MNHN-Bol-V 003371. The
cfs have two distinct wear surfaces forming an acute angle (Fig. 4). The cfs lie
labially relative to the line of molariforms. In occlusal view all of the teeth contain
a large central pulp cavity surrounded by a thick cap of dentine and thinner cap of
orthodentine. Representative lower dental measurements are given in Table 1.
The mfl is triangular in cross-section; mf2 is trapezoidal in cross-section with the
longitudinal axis aligned obliquely with respect to the mfl. The mf3 has a
parallelogram or "figure-8" cross-section with a narrow constriction in the middle
of the tooth that connects the larger anterior and smaller posterior lobes. MNHN-
Bol V 003358 has a dentition with significantly greater wear than exhibited in
MNHN-Bol V 003371; nevertheless they both present the same dental morphology.
MNHN-Bol V 003359 (Fig. 5) represents a juvenile with essentially the same
dental morphology, although at an earlier wear stage than that exhibited in the two
adult specimens. The principal difference in the juvenile is the orientation of the
teeth in the jaw; in medial view (Fig. 5B) the cf and mfl are directed
posterodorsally, whereas the mf2 and mf3 are directed anterodorsally.







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 5

cm


Figure 3. Occlusal (A) and left lateral (b) views of mandible ofGlossotheridium chapadmalense, MNHN-
Bol-V 003371.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)






.............:.i..........................


ant



lab


D 5
I I I c
cm


Figure 4. Occlusal view of left lower dentition (cf, mfl-mf3) ofGlossotheridium chapadmalense, MNHN-
Bol-V 003371. Abbreviations: ant, anterior, lab, labial.





Table 1. Dental measurements (mm) for mandibles of Glossotheridium chapadmalense (Kraglievich),
1925 from Inchasi, Pliocene of Bolivia.


Specimen cfl mfl mf2 mf3
Number AP T AP T AP T AP T


MNHN-Bol-V
003359 8.0 5.3 7.4 5.9 5.6 8.9 13.2 9.0
MNHN-Bol-V
003371 14.4 10.5 13.0 9.1 18.5 11.4 33.0 16.3

AP; antroposterior length
cf, caninifom tooth
mt, moluifiom tooth
T; trnsverse width



Discussion.--Two Pliocene genera of closely related mylodont sloths, i.e.
Glossotherium Owen, 1840 (sensu Simpson 1945), and Glossotheridium
Kraglievich 1934, have been described in the literature. Some authors (e.g.
Simpson 1945) consider Glossotheridium to be a junior synonym of Glossotherium,
whereas others (e.g. Pascual 1967) consider the two genera to be distinct. We
follow Pascual (1967) and relative to Glossotherium, Glossotheridium is smaller,
has a narrower and less robust skull, and has a smaller caniniform. Both of these
genera are characterized by having a bilobate posterior-most molariform (mf3),


;'I







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 5

cm






Nk

A.












Figure 5. Occlusal (A) and medial (B) views of left mandible ofjuvenile specimen of Glossotheridium
chapadmalense, MNHN-Bol-V 003359.



a character clearly developed in the Inchasi specimens described above. The
available dental characters and general size preserved in the Inchasi mylodont
justify allocation to the genus Glossotheridium and common species G.
chapadmalense known from Argentina.
The genus Glossotheridium Kraglievich, 1934 was originally described from
the Pliocene of Argentina. Hoffstetter (1986) mentioned the presence of
Glossotheridium from Pliocene sediments of the northern altiplano of Bolivia. The






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


only previously collected Bolivian material of Glossotheridium available in the
GEOBOL collection (GB-078) was collected from Viscachani (some 10 km to the
south of Ayo Ayo) by Villarroel in 1973. This specimen preserves the anterior
region of the skull, but lacks the dentition. As such, it does not allow a detailed
description of the important dental morphology of this genus. Its primary
significance is to document the occurrence of this genus at another Pliocene site in
Bolivia.
The original description of the type species of Glossotheridium, i. e.
"Eumylodon chapadmalensis" Kraglievich (1925) is known from an associated
skull and mandible. The illustrations from that description allow an identification
of the most important characters of the dentition and mandible of the Bolivian
specimens. The angular orientation, slight wear, and small size of the teeth in
MNHN-Bol V 003359 clearly are evidence of juvenile morphology. However, the
less robust caninform of MNHN-Bol V 003371 relative to MNHN-Bol V 003358
could either be a result of ontogenetic differences in two adult specimens of
different age or could be an example of sexual dimorphism. Because of the small
sample from Inchasi, it is not possible to distinguish between these two possibilities
for the adult specimens; nevertheless, if more specimens are collected from this or
similar-aged localities in the future, perhaps this difference in dental morphology
can be better explained.

"Mylodontinae, indeterminate
Figures 6, 7

Material.--MNHN-Bol-V 00365, R central and distal portion of a femur (Fig.
6) and MNHN-Bol-V 003355, distal (ungual) phalanx (Fig. 7).
Geographic and stratigraphic location.--Respectively, 1100 m north and 350
m west of Communidad Inchasi; 50 m above the base of measured section 1 and 17
m above the base of measured section 3B.
Description.--MNHN-Bol-V 003365 represents most of a R femur, including
the central shaft and distal articular surface (Fig. 6). The shaft is anteroposteriorly
compressed and bowed laterally so that it is concave medially. Distally, there are
two condyles, a larger one medially and a smaller one laterally. There is a well-
developed intercondylar fossa in the middle of the two distal condyles.
The proximal region of the ungual phalanx, MNHN-V-Bol 003355, is divided
into two concave articular surfaces, one positioned laterally and the other medially.
Ventrally there are two medium-sized foramina. The medial and distal portions of
the phalanx are laterally compressed and claw-shaped (Fig. 7). Along the dorsal
surface there is a groove running from the distal tip to midway up the claw.
Discussion.--Hoffstetter et al. (1971; 1972) first reported the presence of small
mylodontines of an undescribed genus from Ayo Ayo, Viscachani, and Pomata.
Later, Hoffstetter (1986) referred the mylodontine to an unspecified taxon different






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


cm


Figure 6. Posterior view of right femur of Mylodontinae, indeterminate, MNHN-Bol-V 003365.


cm

Figure 7. Lateral view of ungual phalanx of Mylodontinae, indeterminate, MNHN-Bol-V 003355.


from Glossotheridium. We refer these specimens to the Mylodontinae because
they are relatively small and gracile and not Megatheriinae which would be larger
and more robust. The specimens from Inchasi, as well as material contained in the
GEOBOL collections from other Pliocene localities in Bolivia, are insufficient to
make a more specific taxonomic identification, or to describe a new genus.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


tSubfamily Scelidotheriinae Ameghino, 1909
tGenus Proscelidodon Bordas, 1935
tProscelidodon patrius (Ameghino), 1888
Figures 8, 9

Material.--MNHN-Bol-V 003353, skull with fragmentary R and L molariforms
MF1-MF5.
Geographic and stratigraphic location.--650 m northwest of Communidad
Inchasi, 13 m above base of measured section 3A.
Description.--In dorsal view, the skull is of medium size and is laterally
compressed from the anterior to medial regions; the posterior region forming the
cranium is expanded (Fig. 8). The premaxilla is small and anteroventrally
separated by two angular points. The nasals are fragmentary. In lateral view the
frontals are slightly convex, and form an undulating surface with the parietal along
the dorsal part of the skull. In dorsal view, the sagittal crest is weakly developed
and the occipital crests are convex. The occipital condyles are well-developed. In
ventral view the palatal suture is fused incompletely and there is a deep trough in
the center of the palate. The basisphenoid, basioccipital, auditory, and right
exoccipital regions are partially preserved. In the orbital region the skull is
constricted and concave with respect to the zygomatic arch. In lateral view the
zygomatic arches are positioned relatively high on the skull. The jugal is small,
the lacrimal does not contact the nasal, and the pterygoids form obtuse angles with
the posterior zygomatic processes. The auditory region is open and does not form a
bulla. Posterior to the orbit the squamosal surrounds the posterior zygomatic arch,
which is broken on both sides of this specimen.
Except for R MF4, the molariforms are fragmentary and broken at the level of
the alveolar margin (Fig. 8). The broken cross-sections of the teeth show
characteristic pulp, thick dentine, and a thinner, external rim of orthodentine. In
cross-section the MF1 is elliptical, and MF2-MF5 are triangular with shallow
lingual folds (Fig. 9).
Discussion.--The genus Proscelidodon was proposed by Bordas (1935) who
used Scelidotherium patrium Ameghino, 1888 as the type species. The original
material consists of a skull and left mandible, so the genus Proscelidodon is
relatively well characterized by a suite of diagnostic characters, including the
following: relative to Scelidodon, about one-third smaller; relative to
Scelidotherium, anterior portion of palate and frontal-parietal regions broader;
zygomatic arch more open, less flattened, and more dorsally positioned; and skull
less laterally compressed.
P. patrius is reported from Argentina in the typical Monte Hermoso and
Chapadmalal formations of Pliocene age (Pascual 1967). The presence of P.
patrius from Inchasi is the first reported occurrence of this taxon from Bolivia.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 5

cm


Figure 8. Dorsal (A), left lateral (B), and ventral (C) views of cranium ofProscelidodon patrius, MNHN-
Bol-V 003353.






102 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)






ant
S0 5
I I I I I I
lab cm
Figure 9. Occlusal view of right dentition (MF1-MF5, breakage below occlusal surface reconstructed) of
Proscelidodon patrius, MNHN-Bol-V 003353. Abbreviations: ant, anterior, lab, labial.



Infraorder Cingulata Illiger, 1811
tFamily Pampatheriidae Paula Couto 1954
fGenus Plaina Castellanos 1937
tPlaina intermedia (Ameghino), 1888
Figures 10-12

Material.--MNHN-Bol-V 003418, complete skull and mandible with anterior
region of vertebral column, L scapula, most of L forelimb, fragment of R forelimb
(Fig. 10); MNHN-Bol-V 003361 (Fig. 11), L mandibular fragment with partial
mf5, mf 6-8, partial mf9; MNHN-Bol-V 003362, portion of carapace with
articulated osteoderms; MNHN-Bol-V 003363 (Fig. 12a), rows of associated and
partially articulated mobile osteoderms; MNHN-Bol-V 003364 (Fig. 12b),
associated and partially articulated osteoderms.
Geographic and stratigraphic location.--MNHN-Bol-V 003418, 003362,
1100 m north of Communidad Inchasi, section 1, Unit A; MNHN-Bol-V 003361,
003363, 003364, 350 m west of Communidad Inchasi, section 3B, Unit A.
Description.--As represented in MNHN-Bol-V 003418 the skull is
anteroposteriorly elongated (Fig. 10) and laterally compressed. The dorsal region
of the skull was covered with oval to quadrate osteoderms during life, one of which
remains attached in this specimen. The premaxilla is small, and although the
sutures are difficult to distinguish in this region, this bone seems to house the CFI
(as represented by an alveolus) and CF2. The maxilla is relatively large and
contains the CF3 and molariform teeth (MF1-MF5; the MF6, which presumably is
present, is medial to, and hidden by, the ascending mandibular ramus). The nasal
is relatively elongated. The anterior portion of the nasal is situated dorsal to the
premaxilla and together they form the nasal opening. In the orbital region, the
lacrimal is small, the jugal is fragmentary, and the frontal contains a prominent
postorbital process. Dorsal to the anterior part of MF4 there is a well-developed
infraorbital foramen. On the parietal, the sagittal crest is weakly developed and in






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


/7 ( ;
*

/

igrn




fs~


Figure 10. Left lateral view of cranium, mandible, anterior part of vertebral column, partial scapula, and left
forelimb ofPlaina intermedia, MNHN-Bol-V 003418.


ant


cm


Figure 11. Occlusal view of left lower dentition (partial mf5, mf6-mf8, partial mf9; breakage below
occlusal surface reconstructed) ofPlaina intermedia, MNHN-Bol-V 003361. Abbreviations: ant, anterior,
lab, labial.


'"'
I
'XG~~'


It ~






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


the ventral region of this bone (near the occipital crest) there are about nine small
foramina on the left side (the right side is not prepared). The occipital crest is
relatively thick and better-developed than the sagittal crest. The mandible consists
of two well-fused rami. The left ascending ramus is complete and forms an angle
of about 900 with respect to the horizontal portion of the ramus. The
posteroventral gonion is rugose and forms a relatively shallow angle. Dorsal to the
gonion there is a notch below the condyle. Dorsal to the condyle there is a well-
developed sigmoid notch and a very high and delicate ascending coronoid process
which extends dorsal to the upper tooth row.
In the upper and lower dentitions it is impossible to distinguish the exact tooth
positions and homologies; therefore we refer to the teeth as caniniform if they are
simple cones and molariforms if they are bilobate. In the upper dentition (not
illustrated) there are positions for three anterior caniniforms, of which two are
preserved, followed by positions for six molariforms, of which 5 are preserved.
The preserved caniniforms are conical with CF2 being slightly smaller than CF3.
The molariforms are bilobate with a larger, better-developed labial groove than the
corresponding lingual structure, as is characteristic for this family.
Despite the fragmentary preservation of MNHN-Bol-V 003361, the general
morphology of the lower dentition is preserved (Fig. 11). The cheek teeth are
hypselodont with medium to high crowns and open roots. The internal dentine is
surrounded by moderately thick external orthodentine. Although mf2-mf6 are only
partially preserved and broken off at the top of the roots, the bilobulate occlusal
surfaces reniformm, sensu Edmund 1985a; 1985b) are preserved in mf3-mf5. The
teeth are strongly compressed labiolingually, with the long axis anteroposteriorly.
There are moderately developed anterior and posterior lobes separated in each
tooth by a large labial groove. As is characteristic for pampatheres (Edmund
1985b), and in addition to the characters described above, portions of the occlusal
surface show transverse striations (Fig. 11), indicating lateral motion of the jaw
during mastication.
With respect to the postcranial skeleton, the anterior portion of the vertebral
column, scapula, and most of the forelimb are preserved in articulation (Fig. 10).
In the proximal (anteromost) scapula there is a very large, recurved acromian
process. As also described by Edmund (1985a 1985b), at least two, and perhaps
three, cervical vertebrae are fused posterior to the atlas. The proximal articular
surface of the humerus is rounded, smooth, and large. The distal half of the
humeral shaft bears a well-developed deltoid crest. The radius and ulnae are
separate bones. Many of the carpal and metacarpal bones are preserved; the
phalanges are not preserved in this specimen.
The osteoderms of the scapular and pelvic bucklers of the carapace vary in
outline from hexagonal to square and are of different sizes (Fig. 12). The external
surface of these osteoderms are ornamented along the margins with numerous
small fossettes, and the central region is relatively smooth. In the dorso-central







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


cj~Y3


0 5
cm


Figure 12. Dorsal view of carapace (A) and close-up (B) view of individual osteodenns of Plaina
intermedia, MNHN-Bol-V 003363 and MNHN-Bol-V 003364, respectively.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


region of the carapace there are movable bands composed of large, elongated
osteoderms. The anterior one-third of each osteoderm has a raised region of
imbrication with the adjoining osteoderm of the postero-adjacent band. Posterior
to the raised region there is a slight transverse, concave depression with a line of
small fossettes. The remaining posterior portion (about half the area of each
osteoderm) consists of a small rugose periphery and a relatively large, internal part
which is relatively smooth except for very tiny pitting on the surface.
Discussion.--The genus Plaina was originally described by Castellanos (1937)
based on specimens collected from the Pliocene of Argentina. Edmund (1985b, p.
92) believes that: "The genus Plaina is based on inadequate material and will
probably be relegated to synonomy." Until this is done, however, we have no
choice but to rely upon this nomen for the most appropriate taxonomic assignment
for the material from Inchasi. In Bolivia, the presence of Plaina was previously
reported by Hoffstetter et al. (1971) and Hoffstetter (1986) from Pliocene sediments
of the Umala Formation from Ayo Ayo and Viscachani.
Plaina and the other genera of the extinct armadillo family Pampatheriidae are
characterized by large size, well developed and anteroposteriorly broad ascending
mandibular ramus, teeth open-rooted (hypselodont), dental formula 9/9 with the
anterior teeth conical (caniniform here) and posterior teeth bilobate or reniform
(molariform here), occlusal surface of molariforms have transverse striations,
presence of fused cervical vertebrae, and carapace with three imbricating bands of
relatively large osteoderms (Edmund 1985b).
Based on the dental, postcranial, and dermal armor morphology described
above, we are confident that the sample from Inchasi is clearly referable to Plaina,
as this genus is presently recognized. Although we were unable to make direct
comparisons with specimens from Argentina, we follow Paula Couto (1979) who
indicates that this is a monotypic genus consisting of the species P. intermedia
(Ameghino), 1888. Thus we are comparing the sample from Inchasi with the only
valid, existing species within Plaina.

fFamily Glyptodontidae Burmeister, 1879
tGenus Plohophorus Ameghino, 1887
tPlohophorus sp.
Figures 13-14

Material.--MNHN-Bol-V 003369, dorsal and lateral portion of a carapace.
Geographic and stratigraphic location.--1200 m southwest of Communidad
Inchasi, about 18 m above the base of measured section 4.
Description.--The articulated osteoderms are of moderate thickness and either
polygonal or hexagonal in shape (Figs. 13-14). The internal surfaces of these
osteoderms are rugose with medium-sized fossettes distributed irregularly. In the
central osteoderm the external surface is also rough, or rugose. Each of these







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA 107


0 10

cm


Figure 13. Dorsal view of carapace with articulated osteoderms ofPlohophorus sp., MNHN-BoI-V
003369.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


0 5
cm
Figure 14. Close-up, dorsal view of articulated central osteoderms ofPlohophorus sp., MNHN-Bol V
003369.



central osteoderms characteristically consists of a central, concave hexagon
flanked by 9-10 surrounding smaller (approximately half the diameter of the
central hexagon) sub-elements of either hexagonal, pentagonal, or irregular
outline. The lateral osteoderms are smaller, more simple in design, and generally
consist of a single, concave element with rugose surface of either pentagonal,
hexagonal, or irregular shape. The rugosities are in the form of small fossettes or
parallel lines. Slightly more than half of the portion of the preserved carapace
consists of the central, more complex osteoderms and the remainder consists of the
simpler lateral osteoderms. A notable characteristic of this carapace is that the
boundary between the two types of osteoderms is relatively indistinct and undulates
around the perimeter of central osteoderms.
Discussion.--As with almost all of the species of Plohophorus Ameghino
(1887), our knowledge of this genus is based primarily on the caudal armor. An
exception is P. figuratus which is known from carapace osteoderms from the
Pliocene of Monte Hermoso in Argentina. As such, our knowledge of the
morphology and specific differences within Plohoporus is quite incomplete. The






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


relatively complete, although partial, carapace from Inchasi allows a better
understanding of the morphological variation of individual osteoderms. We are
able to assign this specimen to Plohoporus based on the presence of highly
punctate plates of moderate thickness and of a general size similar to specimens
assigned to this genus from Argentina.
There are numerous other specimens of osteoderms and fragmentary carapaces
contained in the GB and MHNH-Bol collections attributable to Plohophorus from
the Pliocene localities of Ayo Ayo, Viscachani, and Pomata on the altiplano near
La Paz (also see Hoffstetter 1986). Although of generally similar morphology to
the Inchasi specimen, the lack of relevant diagnostic material (i.e. caudal armor)
and poor characterization of isolated osteoderms or partial carapaces from the
various species of Plohophorus from Argentina prevents a more specific
assignment. As such, this is the first description of portions of the carapace of this
genus from the Pliocene of Bolivia. In Argentina, Plohophorus has a range from
the middle to late Pliocene and it is characteristically found in Chapadmalalan-
aged sediments (Pascual 1967). Given the available paleomagnetic correlations
(MacFadden et al. 1993), the Inchasi occurrence is middle Pliocene in age.

tGenus Paraglyptodon Castellanos, 1932
tParaglyptodon sp.
Figure 15

Material.--MNHN-Bol-V 003366, 7 articulated carapace osteoderms and 2
caudal plates.
Geographic and stratigraphic location.--1100 m north of Communidad
Inchasi, approximately 36 m above the base of measured section 1.
Description.--The carapace osteoderms are relatively thin and hexagonal (Fig.
15). Each of these osteoderms consist of a surface covered with numerous tiny
fossettes and a central region delimited by a circular depression. The lateral
portions of each of these plates are separated by 6-9 shallow grooves radiating to
the periphery. The caudal osteoderms are pentagonal, relatively flat, and with a
rough external surface.
Discussion.--Hoffstetter et al. (1972) and Hoffstetter (1986) described the
presence of glyptodonts related to, but different from, Plohophorops from north of
Callapa and Anchocalla on the Bolivian altiplano. These forms, however, were not
assigned to a particular glyptodont genus. With the material from Inchasi we are
able to place on record the first-known occurrence of Paraglyptodon Castellanos
(1932) from Bolivia based on the unique characters of this genus including: (1)
size smaller than Glyptodon; (2) carapace osteoderms hexagonal; and (3) rugose
osteoderms with large, well defined central figure. We are unable to assign these
new specimens to either an existing species from Argentina, or to distinguish them
as new. Given the paleomagnetic constraints for Inchasi, the Bolivian occurrence
of Paraglyptodon sp. is middle Pliocene in age.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


cm
Figure 15. Articulated carapace osteoderms ofParaglyptodon sp., MNHN-Bol-V 003366.



Order Rodentia Bowdich, 1821
Suborder Caviomorpha Wood & Patterson, 1955
Family Caviidae Waterhouse, 1839
tGenus Caviodon Ameghino, 1885
tCaviodon sp.
Figure 16

Material.--MNHN-Bol-V 003367, fragmentary cheek teeth of uncertain
position.
Geographic and stratigraphic location.--1100 m north of Communidad
Inchasi, 59 m above the base of measured section 1.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 7
| I i U i I I
mm
Figure 16. Occlusal view of cheek-tooth fragments (probably P4/p4, Ml/ml, or M2/m2) of Caviodon sp.,
MNHN-Bol-V 003367.



Description.--These specimens have high-crowned, curved hypselodont teeth
with open roots. Two of the teeth from Inchasi consist of a single prism. Another
incomplete fragment consists of two small prisms. The better-preserved cheek
teeth consist of two larger prisms forming an "M" shape (Fig. 16). The apices of
each prism are anteroposteriorly compressed and are separated by a deep groove;
each prism also contains a shallow groove, and there are three columns extending
the height of the tooth. The crowns of all teeth are covered with enamel.
Discussion.--The genus Caviodon Ameghino (1885) was originally described
from Argentina and is known to range from the middle to late Pliocene (Pascual
1967). Prior to this description, Caviodon had neither been reported in the
literature nor was it present in any collections that we have examined from the
Pliocene of Bolivia. Given the fragmentary nature of the available specimens from
Inchasi, we cannot refer this sample to either an existing or new species, nor can
we confidently identify the exact position of individual fragments within the tooth-
row. Following the illustrations in Pascual (1967) the two better-preserved
specimens (Fig. 16) probably are either P4/p4, Mi/ml, or M2/m2 (distinction
between uppers and lowers are very difficult, if not impossible, in fragmentary
specimens of this taxon). Despite the incomplete nature of the Inchasi specimens,
they are confidently referred to Caviodon (rather that the closely related
Cardiomys) because of the presence in the former genus of the very deep grooves
(or, re-entrants) characterizing each lobe. The Caviodon sample from Inchasi is of
similar size to representatives of this genus from Argentina (Pascual 1967). Given
the paleomagnetic correlations for Inchasi (MacFadden et al. 1993), we assign the
Bolivian occurrence of Caviodon sp. to the middle Pliocene.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


Family Hydrochoeridae Gill, 1872
fGenus Chapalmatherium Ameghino, 1908
tChapalmatherium cf. saavedrai Hoffstetter et al., 1984
Figure 17

Material.--MNHN-Bol-V 003368, incisor fragments, anterior prism of ?L m2,
and fragmentary m3.
Geographic and stratigraphic location.--750 m east of Communidad Inchasi,
about 56 m above base of measured section 2.
Description.--The lower portion of the anteromost part of an incisor (not
illustrated here) shows diagnostic rodent-like enamel only on the anterior surface.
The fragmentary cheek teeth consist of a portion of the anterior prism and a
complete posterior prism which are separated by a deep groove (Fig. 17). The
M3/m3? consists of fragments of isolated plates characteristically separated by
cement. These teeth are relatively high-crowned. Although measurements are not
possible because of the fragmentary nature of the specimens, general comparisons
of the sample from Inchasi indicates that it is about 50% smaller than the
previously described hypodigm of Chapalmatherium saavedrai from the Bolivian
altiplano (Hoffstetter et al. 1984).
Discussion.--Ameghino (1908) originally proposed the genus
Chapalmatherium from Pliocene sediments in Argentina. Hoffstetter et al. (1971)
described the presence of this genus from Pliocene sediments of the Umala
Formation at Ayo Ayo and Viscachani on the Bolivian altiplano south of La Paz.
Hoffstetter et al. (1984) described the new species C. saavedrai based on a well-
preserved lower ramus and corresponding dentition (GB 081). The diagnostic















0 4
mm
Figure 17. Occlusal view of cheek teeth of Chapalmatherium cf. saavedrai, MNHN-Bol-V 00338. Given
the prism morphology, the specimen on the left is from a 3rd molar (upper/lower indeterminate); the
specimen on the right is a more anterior tooth (P4/p4, M /m or M2/m2).






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


generic and specific characters relate to the similar morphology of the enamel
prisms. Despite the paucity of material from Inchasi and the difficulty in
determining the exact position in the tooth row for individual fragments, we can
confidently refer it to Chapamaltherium based on the presence of diagnostic
characters, including: (1) cheek teeth (except M3/m3) consisting of triangular
prisms with deep re-entrants surrounded by cement; and (2) the middle and
posterior portions of M3/m3 consisting of a series of relatively flattened prisms
alternating with vertical layers of cement. We note, however, that the sample from
Inchasi is smaller than any of the species attributed to this genus from Argentina
(Pascual 1967), and it is about 50% smaller than the hypodigm of C. saavedrai
from Ayo Ayo and Viscachani (Hoffstetter et al. 1984). The Inchasi
Chapalmatherium appears to be a new species, but the fragmentary material is
insufficient upon which to formally describe it. Given the known range of this
genus from Ayo Ayo-Viscachani and those of Argentina, this occurrence indicates
a middle Pliocene age for Inchasi.

tOrder Litopterna Ameghino, 1889
fFamily Macraucheniidae Gill, 1872
tGenus Promacrauchenia Ameghino, 1904
tPromacrauchenia sp.
Figures 18-21, Tables 2, 3

Material.--MNHN-Bol-V 003351, L mandible with partial m2-m3 (Fig. 18a,
b); MNHN-Bol-V 003352 (Fig. 18C, D; Fig. 19), partial ventral symphysis and R
mandible with p2-m2; MNHN-Bol-V 003376, R mandibular fragment with m3
alveolus; MNHN-Bol-V 003372, R partial forearm (Fig. 20); MNHN-Bol-V
003374, partial R femur with shaft and distal articular surface (Fig. 21) and
associated partial tibia with distal articular surface and shaft; MNHN-Bol-V
003377, partial tibia and fibula with distal articular surface; MNHN-Bol-V
003378, partial R tibia with proximal articular surface and fragments of distal
articular surface.
Geographic and stratigraphic location.--The sample listed above was
collected from two localities: 350 m west and 1200 m southwest of Communidad
Inchasi, respectively, 17 m above base of measured section 3B and 18 m above base
of measured section 4.
Description.-MNHN-Bol-V 003351 consists of fragmentary cheek teeth which
are subhypsodont with a partial external cingulum located between the protolophid
and metalophid (Fig. 18). The alveolus for ml is double-rooted, only the posterior
of which is preserved. In the better-preserved MNHN-Bol-V 003352 two mental
foramina are present ventral to p3 and ml. The symphysial suture is well-fused,
indicating an adult. The incisors and canine are not preserved, but their alveoli
suggest teeth of moderate size relative to the cheek teeth. The lingual cingula are














A C o













0
cm cm














Figure 18. Occlusal (A, C) and lateral (B, D) views of mandibles of Promacruchenia sp., MNHN-Bol-V 003351 and MNHN-Bol-V 003352, W
respectively.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


cm .Fa


Figure 19. Occlusal view of right p2-m2 ofPromacrauchenia sp., MNHN-Bol-V 003352. Abbreviations:
ant, anterior, lab, labial.





Table 2. Mandibular measurements (mm) for Promacrauchenia sp. from Inchasi, Pliocene of Bolivia.


MNHN-Bol-V 003351 Length of series m2-m3 55.4
Mandibular depth below m3 42.8
Thickness of the mandible at position of m3 23.7

MNHN-Bol-V 003352 Length of the series p2-m2 96.7
Length of the series p2-p4 52.0
Length of the series ml-m2 46.6
Mandibular depth below m2 36.2
Mandibular depth below p3 30.6
Thickness of the mandible at position of m2 22.6@



incomplete in p2-p3 and better developed in ml-m2. The p2 is relatively simple
and lacks separate lophids (Fig. 19). In p3-p4 the protolophid is, respectively,
smaller and slightly smaller than the metalophid. In ml-m2 the protolophids and
metalophids are subequal in size. In occlusal view both specimens have well-
developed pulp cavities (Fig. 18; not indicated in Fig. 19) surrounded by dentine
and enamel. Representative mandibular and lower dental measurements are
presented in Tables 2 and 3.
As represented in MNHN-Bol-V 003372, the right forelimb is partially
preserved distal to the elbow (Fig. 20). The radius is anteroposteriorly compressed,
exhibiting a well-developed medial crest on the proximal part of the shaft. The
radius and ulnae are fused (indicated by dashed line in Fig. 20) for most of their
lengths except for the distal articulation with the proximal carpals. The distal
articular surface of the unfused portion of the ulnae articulates with the
proximolateral carpal, i.e. cuneiform. The cuneiform is elongated proximodistally


I



















Table 3. Measurements (mm) for lower dentitions ofPromacrauchenia sp. from Inchasi, Pliocene of Bolivia.


Specimen p2 p3 p4 ml m2 m3
Number AP T AP T AP T AP T AP T AP T


MNHN-Bol-V
003351 27.6 15.1 30.1 16.8
MNHN-Bol-V
003352 15.1 9.3 17.7 11.0 21.2 12.0 19.8 12.3 27.3 13.7
GEOBOL s/n 28.2 11.7 21.0 13.0 24.2 14.3 34.1 17.7 40.2 10.2 40.0 18.7

AP Antoposteior length
s/n = unnumbered specimen
T = Tansvere width





ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


radius


ulna








lunar,


Figure 20. Anterior view of right forelimb ofPromacrauchenia sp., MNHN-Bol-V 003372. Dashed line
indicates fusion of radius and ulna.


scaphoid

trapezoid
magnum












0 5 10
cm






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


X-


.4 :49
.." 4




'it
':


#i;zp' '


B

k b


0 5
cm


Figure 21. Anterior (A), medial (B), and distal (C) views of partial right femur of Promacrauchenia sp.,
MNHN-Bol-V 003374.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


proximodistally and narrow mediolaterally. The posterior portion of the cuneiform
has a well-developed facet for articulation with the pisiform (not illustrated). The
carpals that articulate with the radius are the lunar medially and the scaphoid
internally. The lunar is rectangular with an elongated proximodistal axis and is
compressed mediolaterally. The scaphoid is also generally rectangular and
elongated mediolaterally. The distal carpal series includes the uniform laterally,
magnum medially, and trapezoid internally. These bones are generally square,
with the uniform the largest, the magnum intermediate, and the trapezoid the
smallest carpal of this series. These bones articulate with convex facets with the
proximal carpal series. Distally the second carpal series articulates with
metacarpals II, III, and IV. Metacarpal III is the longest (167 mm) and MC II and
IV are of approximately equal length (151 mm and 140 mm, respectively). Distal
to these metacarpals, only the phalanges of MC III are preserved. Of these
elements, the proximal MC III is the longest (53 mm) and has widened proximal
and distal articular surfaces. The medial MC III is subrectangular with a length of
29 mm. The distal-most (ungual) phalanx has a widened bony distal terminus
forming a hoof.
As represented in MNHN-Bol-V 003374, the femur consists of a moderately
developed shaft with a third trochanter developed proximolaterally (Fig. 21). The
distal articular region consists of a fragmentary smaller lateral condyle and a larger
well-developed medial condyle. In anterior view the external trochlear crest is
smaller than the medial trochlear crest; these are separated by a deep intercondylar
fossa. The proximal part of the medial trochlear crest has a moderately developed
notch.
The tibia (not illustrated here) has a well-developed concave facet on the
proximal articular surface for articulation with the medial condyle of the femur.
There is a well-developed fossa for articulation with the patella. The posterolateral
region is not preserved in MNHN-Bol-V 003377, but in MNHN-Bol-V 003378
there is a moderately developed surface for articulation with the lateral condyle. In
anterior view the cnemial crest is well-developed. The central shaft is triangular in
cross-section. The distal articular surface of MNHN-Bol-V 003377 exhibits the
internal malleolus; the distolateral facet of the fibula is not preserved. The distal
portion of the fibular shaft is gracile, mostly distinct, but also partially fused to the
tibia.
Discussion.--The genus Promacrauchenia Ameghino (1904) was originally
described from the late Pliocene of Argentina (Pascual 1967) and is based on the
type species P. antiqua (=Macrauchenia antiqua Ameghino, 1887). This genus is
distinctly macrauchenine in dental and postcranial morphology, including: (1)
brachyodont cheek teeth with well-developed and distinct roots; (2) P2/p2 and
P3/p3 relatively small and not molariform; (3) P4/p4 submolarifiorm; (4) relatively
well-developed cingula; (5) shallow mandible; and (6) functionally tridactyl limbs
with slightly larger MP III relative to MP II and IV (the latter character
differentiates the family Macraucheniidae from their close relatives, the






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


Proterotheriidae). The basic difference between Promacrauchenia and the
relatively well-known Pleistocene Macrauchenia is that the former genus is about
one-third smaller in size (Pascual 1967).
In his thesis, Montafio (1968) cited the discovery of a femur and calcaneum of a
small macrauchenine, which he referred to "Microauchenia montaloi" (currently a
nomen nudum) from Quebrada Tijraska, which is an undescribed site located west
of Anzaldo, Bolivia. Marshall et al. (1983) and Hoffstetter (1986) referred this
material to cf. Promacrauchenia and assigned it a Pliocene age.
Hoffstetter et al. (1971) described two macrauchenid limb specimens from the
Pliocene of Ayo Ayo, which they refer to a small macrauchenid, possibly
Promacrauchenia. They also noted that this genus is present at other Pliocene
sites in Bolivia. Our comparisons with available collections indicate that
Macrauchenia is the common macrauchenine from the Pleistocene of Bolivia
whereas Promacrauchenia, which is of generally similar morphology but about
one-third the size of the former genus, is the correct taxonomic reference for
Pliocene litopterns from Bolivia, as they are currently known. As such, the
specimens from Inchasi represent the first definitive description of
Promacrauchenia from Bolivia and indicate a middle Pliocene occurrence based on
paleomagnetic calibrations at this site. This age is consistent with known
occurrences of this genus from the middle and late Pliocene in Argentina (Pascual
1967).

tOrder Notoungulata Roth, 1903
tSuborder Toxodonta Scott, 1904
fFamily Toxodontidae Gervais, 1847
tGenus Posnanskytherium Liendo, 1943
Figures 22-25, Tables 4, 5

Genoholotype and locality.--GB 004, P. desaguaderoi Liendo, 1943,
consisting of well-preserved skull and mandible. The exact locality is uncertain,
but as the type species name indicates, Villarroel (1977) states that it was collected
from Pliocene sediments along the Rio Desaguadero south of Lake Titicaca on the
Bolivia-Peruvian border. Other referred specimens described in Villarroel (1977)
come from the La Paz Valley and Ayo Ayo-Viscachani, both also known to be
Pliocene localities.
Generic diagnosis.--Dental formula and skull morphology referable to
toxodont subfamily Xotodontinae. Size smaller than Xotodon Ameghino. In
lateral view, the cranial vault of Posnanskytherium is high, relatively flat, and has
a weakly developed sagittal crest. Zygomatic arch less-well developed than in
Xotodon. Mandibular rami posterior to the symphysis subparallel. Dental formula
2/3, 0/1, 4/3, 3/3, which is characteristic of the subfamily Xotodontinae. Upper
and lower molariform teeth strongly compressed labiolingually. Incisors broad






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 5
cm


B

Figure 22. Dorsal (A) and left lateral (B) views of mandible ofPosnanskytherium desaguaderoi, MNHN-
Bol-V 003360.



transversely and slightly inclined posteromedially. P4 has a small posterolingual
fold covered with enamel. M1-M2 each consist of one very deep lingual fold. M3
posterior to the main lingual fold has a weakly developed fold with poorly
developed enamel. Lower molars with a single lingual fold; paraconid reduced,
and much smaller than the protoconid (modified from Villarroel 1977).
Discussion.-The genus Posnanskytherium consists of one previously described
species, P. desaguaderoi, and a new species described here, P. inchasense. This






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


genus is only known from the Bolivian altiplano. Its exact affinities, other than
being most closely related to xotodontine toxodonts, is otherwise unknown at
present.

tPosnanskytherium desaguaderoi Liendo, 1943
Figures 22, 23, Tables 4, 5

Material.--From Inchasi: MNHN-Bol-V 003360, symphysis and L mandibular
ramus with il-m3 and R il-p4 (Figs. 22, 23a); MNHN-Bol-V 003370, nearly
complete mandible with L & R il-i2, c, p2-p4, ml-m3; MNHN-Bol-V 003373,
upper dental fragments; MNHN-Bol-V 003379, L mandibular fragment with m3;
MNHN-Bol-V 003417, premaxilla with R and L il-i2. (For referred material from
other localities, see Villarroel 1977).
Geographic and stratigraphic location.--MNHN-Bol-V 003360, 750 m east
of Communidad Inchasi, 56 m above base of measured section 2; MNHN-Bol-V
003370, 003373, 003379, 003417, 1200 m southwest of Communidad Inchasi, 18
m above base of section 4.
Diagnosis.--As for genus (above), with the following specific differentia
relative to P. inchasense: P. desaguaderoi is larger, has a shallower mandibular
ramus, and there are well-developed posterolabial grooves in the upper molars and
anterolabial folds in the lower molars.
Description.--One fragmentary specimen, MNHN-Bol-V 003417 (not
illustrated here), preserves the premaxillary region with a portion of the nasal
crest. The Il is very wide transversely; the Ils from the right and left sides form a
linear cropping mechanism. 12 is positioned posterolabial to II and has a
triangular cross-section.
As exemplified in MNHN-Bol-V 003360, the rami are robustly fused at the
symphysis (Fig. 22). The symphysis is well-developed and extends posteriorly to a
position ventral to p4-ml. The ramus is deep, with the maximum depth developed
ventral to m3; it is laterally narrow extending anteriorly to the symphysis. The
cheek tooth rows are subparallel and converge slightly anteriorly. There is a
moderately well developed postcanine diastema. In MNHN-Bol-V 003360 the
diastema is relatively shorter than in MNHN-Bol-V 003370 and the holotype, GB
004. The main body of the ramus is compressed labiolingually. The posterior
region of the ascending ramus is preserved in MNHN-Bol-V 003370 but not in
MNHN-Bol-V 003360 or the holotype. In MNHN-Bol-V 003370 the gonion
(posteroventral surface of blade), coronoid, and sigmoid notch are all well-
developed. There is a well-developed complex of mental foramina ventral to p4
and ml. Representative mandibular measurements are presented in Table 4.
The il-i2 are anteroventrally compressed and form a distinctly linear cropping
mechanism with the long axis oriented tranversely. The i3 is relatively small,
triangular, and positioned posterolabial to il-i2. In MNHN-Bol-V 003360 the







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


Table 4. Mandibular measurements (mm) ofPosnanskytherium desaguaderoi and P. inchasense from the
Pliocene of Bolivia.

Measurement P. desaguaderoi P. desaguaderoi P. desaguaderoi P. inchasense
Inchasi Inchasi holotype holotype-Inchasi
MNHN-Bol-V MNHN-Bol-V GEOBOL MNHN-Bol-V
003360 003370 GB004 003350

Mandibular height
atml 63.8 65.5 73.3 92.0
Mandibular height
at m3 64.9 70.2 75.6 98.6
Length series
pl-m3 114.6 114.1 122.3 129.6
Length series
pl-p3 38.3 37.7 39.8 40.2
Length ml-m3 77.4 80.1 80.8 89.7
Length symphysis 77.9 111.2 106.2 117.8
Ant symphysial
width 22.0 27.9 33.1 33.8
Post. symphysial
height 30.0 35.8 45.0 44.0


A Posnanskytherium desaguaderoi



^Dcr'c&zGZ2P


a n t 0- 1


0 5
cm


B Posnanskytherium inchasense, n. sp.







Figure 23. Occlusal views of left lower dentitions (p2-m3) of (A) Posnanskytherium desaguaderoi,
MNHN-Bol-V 003360, and (B)Posnanskytherium inchasense, new species, MNHN-Bol-V 003350,
holotype. Abbreviations: ant, anterior, lab, labial.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


A 9 5
cm


L-... IA'


cm
Figure 24. Ventral skull and dorsal mandibular (A) and left lateral (B) views ofPosnanskytherium
inchasense, new species, MNHN-Bol-V 003350, holotype.








Table 5. Measurements (mm) for upper and lower dentitions ofPosnanskytherium desaguaderoi and P. inchasense
from the Pliocene of Bolivia.


Upper II I2 PI P2 P3 P4 Ml M2 M3
Dentition AP T AP T AP T AP T AP T AP T AP T AP T AP T


MNHN-Bol-V 003350,
P. inchasense
Holotype 13.2 18.4 8.1 5.4 13.5 8.5 19.0 10.5 21.6 11.1 34.6 11.8 41.8 11.7 41.0 11.9
GB 004,
P. desaguaderoi,
Holotype 26.2 8.5 13.6 18.4 6.1 5.2 13.8 9.6 17.1 9.9 22.4 9.3 30.7 12.4 33.0 11.6 35.5 12.2


Lower
Dentition


MNHN-Bol-V 003350,
P.inchasense,
Holotype
MNHM-Bol-V 003360,
P. desaguaderoi
GB 004,
P. desaguaderoi,
Holotype
MNHN-Bol-V 003370,
P. desaguaderoi

AP; Antroposterior length
T; Transverse width


il i2 i3 cl p2 p3 p4 ml m2 m3
AP T AP T AP T AP T AP T AP T AP T AP T AP T AP T


7.5 14.5 6.8 14.2 3.6 3.1 8.3 6.0 12.1 8.3 16.3 9.2 26.7 9.8 27.4 10.2 37.7 9.2

5.8 13.5 5.6 12.7 9.8 14.8 8.2 5.0 11.5 6.6 15.5 7.8 22.8 8.8 24.0 8.7 32.1 8.4


7.3 14.9 6.1 14.2 9.6 14.8 4.7 3.0 9.6 5.6 11.3 7.3 16.9 8.4 23.6 9.0 24.0 9.3 33.0 8.6

7.0 14.0 6.0 13.6 10.7 16.3 7.7 4.9 11.3 6.3 16.1 8.0 23.9 7.9 26.1 8.0 33.0 7.4






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


lab
O 5
ant cm

Figure 25. Occlusal view of left P2-M3 ofPosnanskytherium inchasense, new species, MNHN-Bol-V
003350, holotype. Abbreviations: ant, anterior, lab, labial.



canine is small, elliptical in cross-section and positioned posterior to i3; there is a
small precanine and larger postcanine diastema. The cheek teeth are very high-
crowned (hypselodont), very laterally compressed, and in cross-section consist of a
large pulp (not illustrated) and dentinal regions surrounded by relatively thin
enamel (Fig. 23). The p2 is triangular, p3 is roughly rectangular; both of these
teeth are relatively simple in cross-section; p4 is submolariform; p2-p4 have
concave lingual borders covered with enamel; these are progressively larger and
include double concavities in ml-m3; ml-m3 have well-developed, protolophids
and metalophids; m3 has a well-developed, elongated posterior heel. Each molar
has a single lingual fold between the protolophid and metalophid. In MNHN-Bol-
V 003370 the molar protoconid lobes are better developed than in MNHN-Bol-V
003360 and the holotype. Representative dental measurements are presented in
Table 5.
Discussion.--The genus and species Posnanskytherium desaguaderoi was
originally proposed by Liendo (1943) based on a nearly complete skull and
mandible from the Pliocene of Bolivia. The same material was later redescribed by
Villarroel (1977), who also included several other referred specimens from other
Pliocene localities on the Bolivian altiplano. The availability of the holotype (GB
004) and referred material (GB 098 and GB 099, also see Villarroel 1977) allow
the direct comparisons with the Inchasi specimens.
The exact provenience, horizon, and geographic locality of the holotype of P.
desaguaderoi is not certain at this time; various authors attribute different ages to
this species. Liendo (1943) attributed this occurrence to middle Tertiary (but not a
particular epoch), whereas Ortega (1970) considered it to be middle Pliocene and
Villarroel (1977) and Hoffstetter et al. (1971) consider it upper Pliocene. Based on
paleomagnetic data, MacFadden et al. (1993) correlate the Inchasi locality and its
fauna to the middle Pliocene, thus essentially confirming the age assignments for
this species of the previous authors.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


The combination of morphological characters of the skull and dentition makes
the Inchasi specimens indistinguishable from those previously referred to P.
desaguaderoi. However, the Inchasi specimens differ in being slightly smaller
than the holotype, (Table 5) but larger than the referred specimen from Alto
Seqiiencoma (GB 098), located in the La Paz Valley. We believe that this
difference is not significant, and, as such, can be confidently assigned to this
species. As mentioned above, these occurrences all appear to be middle Pliocene in
age.

tPosnanskytherium inchasense, new species
Figures 23-25, Tables 4, 5

Holotype.-MNHN-Bol-V 003350, partial cranium and mandible with R 12,
P2-P3; L P2-M3; R il-i2, c, p3-m3; L il-i2, p2-m3.
Etymology.--for Communidad Inchasi, location of the holotype.
Geographic and stratigraphic location.--1200 m southwest of Communidad
Inchasi, 18 m above the base of measured section 4.
Generic diagnosis.--See above.
Specific Diagnosis.--Dental formula 2/3, 0/1, 4/3, 3/3. Lower tooth row length
130 mm. Very high-crowned, hypselodont teeth. Relative to P. desaguaderoi,
shorter, deeper and more robust mandibular ramus, upper cheek teeth lacking
well-developed posterolingual grooves, and the absence of anterolingual folds in
the lower molars. P4 and M3 lack posterolingual folds seen in genoholotype.
Lower molars with weaker lingual folds, and m3 longer than in P. desaguaderoi.
Description.--MNHN-Bol-V 003350 represents a relatively well preserved
facial region and jaws of a mature individual in moderate wear (Fig. 24). The
nasal notch is developed over the premaxillary/maxillary suture dorsal to the
postcanine diastema. Only the anteromost portion of the nasals is preserved. Each
premaxilla is relatively short; dorsally these bones have a rugose crest which is
better developed than in P. desaguaderoi. The left maxilla is preserved up to the
nasal suture. Although fragmentary, the lacrimal extends anterior to the orbit to a
position lying above P2. The jugal extends to the same height as the squamosal
suture. The zygomatic arch is well-developed and forms a distinct ventral crest;
dorsally the zygomatic arch lacks the posterior orbital projection seen in the
genoholotype. In ventral view there are several palatal foramina.
Il is not preserved. 12 is triangular in cross-section. The alveoli for 13 indicate
a large tooth positioned posterolabially to 11-12. The upper canines are not
preserved. A large diastema (ca. 60 mm) separates the C from the alveolus of P1.
All of the cheek teeth are hypselodont and lack enamel on the antero- and
posteromost part of the tooth. P1 is relatively small and elliptical. P2 and P3 are
roughly oval (Fig. 25). P4 is the largest premolar and is submolariform; it lacks
the well-developed posterolingual fold seen in P. desaguaderoi. In general
morphology the MI-M2 are similar to the type species, consisting of a greatly






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


expanded anterolabial region and two lingual lophs divided by a well-developed
groove. The M3 of P. inchasense differs from that of P. desaguaderoi in that the
former lacks a shallow posterolingual fold. Representative dental measurements
are presented in Table 5.
The mandible has a heavily fused symphysis extending posteriorly to ml
similar to the genotypic species (Fig. 24). In general morphology, the mandible is
very deep and laterally compressed. Representative mandibular measurements are
presented in Table 4. The tooth row and sides of the mandible are subparallel in
the symphysial region and slightly diverge posteriorly. The maximum mandibular
depth is developed ventral to m3. Three mental foramina, which are partially
confluent, are present on the lateral side of the mandible ventral to p3-ml (Fig.
24B). The anterior portion of the ascending ramus rises at a steep angle (almost
900) relative to the tooth row; the posterior portion of the ascending ramus is not
preserved. The il and i2 are anteroposteriorly compressed and aligned to form a
linear cropping mechanism. Although the i3s are not preserved, their alveoli are
large and triangular. The canines are barely erupted and appear very small,
conical, and positioned relatively close to the il. There is a relatively large
postcanine diastema. The pi is absent. The p2 has a curved crown, is roughly
conical in cross-section, and has a very small lingual fold (Fig. 23b); p3 is
subrectangular; p4 is rectangular and submolariform. The p3-m3 are laterally
compressed, and consist of a large pulp cavity, surrounded by dentine and
relatively thin enamel, and exhibit lingual folds that are larger than the labial
folds. The ml and m2 consist of three principal lophids separated by folds. On the
lingual side there are two shallow folds each on ml and m2; these are less distinct
than in P. desaguaderoi. The ml-m3 each have a weak labial fold. The m3 is the
longest tooth and contains weak folds and an elongated posterior heel. The relative
length of m3 is greater in P. inchasense than in P. desaguaderoi.
Discussion.--Leindo (1943) originally proposed the genus and species
Posnanskytherium desaguaderoi based on a skull and mandible that was
redescribed by Villarroel (1977). While the latter author confirmed the Pliocene
age for this occurrence, the actual locality of the genoholotype is disputed (see
discussion above). Hoffstetter et al. (1971) reported the presence of mandibular
fragments of a xotodontine notoungulate comparable to P. desaguaderoi from
Pliocene sediments of the Umala Formation at Ayo Ayo. Later, Hoffstetter (1986)
recognized two species of Posnanskytherium, a larger one from Ayo Ayo,
Viscachani, and Pomata and a smaller, undescribed species from Achocalla, both
of Pliocene age from the Bolivian altiplano. It seems plausible that his second
species is what we are calling here P. inchasense.
The holotype of the new species P. inchasense, MNHN-Bol-V 003350, differs
in proportions from P. desaguaderoi in that it is larger and has a significantly
deeper and more robust jaw (Table 4). Some of the most notable differences in P.
inchasense include P4 and M3 that lack small posterolingual depressions and the






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


absence of anterolingual folds in the lower molars, both of which are present in P.
desaguaderoi (Fig. 22).
Based on the available sample of the genus Posnanskytherium, the differences
in size and proportions described here cannot be attributed to sexual dimorphism,
nor do they represent intraspecific variation. We therefore assert that the smaller
taxon from Inchasi represents a new species, described here as P. inchasense.

tFamily ?Mesotheriidae Alston, 1876
tSubfamily ?Mesotheriinae Alston, 1876
tHypsitherium bolivianum, new genus and new species
Figures 26-28, Table 6

Holotype.-MNHN-Bol-V 003356, partial L mandible with p4-m2, m3
partially erupted (Figs. 26, 27a).
Referred Specimen.-MNHN-Bol-V 003357, fragmentary R maxilla with Ml-
M3 (Figs. 27b, 28).
Geographic and stratigraphic location.--For MNHN-Bol-V 003356 and
003357, respectively, 650 m north and 1200 m southwest of Communidad Inchasi,
13 and 18 m above the base of sections 3A and 4.
Etymology.--Generic: Gr., hypsi-, high, on high, or high place; -therium,
mammal, beast, or wild animal; in reference to the discovery of this new genus
from the eastern Andean Cordillera at a current elevation of 3,200 m; specific:
bolivianum, for Bolivia.
Generic and specific (because of monotypy) diagnoses.--Small ?mesotheriid
(also see Discussion below for family characters) with M1-M3 anteroposterior
length about 36.5 mm; upper dentitions strongly curved hypselodont teeth with Ml
crown height exceeding 29 mm. Ascending ramus of jaw forms an angle of about
450 relative to the horizontal ramus; upper cheek teeth lacking well-developed
third (posterolingual) lobe distinctive of all other mesotheres; lower cheek teeth
with well-developed lingual folds and labial grooves; size comparable with
Microtypotherium; depth of mandible relatively deeper ventral to ml than in
Plesiotypotherium. Differs from other mesotheres in the presence of anterior lower
premolars (p3 and possibly p2).
Description.--As represented in the holotype MNHN-Bol-V 003356, the
mandible is relatively short and narrow with its greatest depth ventral to ml (Fig.
26). There is a groove containing mental foramina ventral to p4-ml. Part of the
symphysial region is preserved including the alveoli for p2-p3 (Fig. 26B). The
symphysial suture extends posteriorly to beneath the p3-p4 junction. Posteriorly,
only the anterior portion of the ascending ramus is preserved.
The lower cheek teeth are hypselodont. The dentine is covered with thin
enamel; the crowns are compressed labiolingually with the anteroposterior axis
being much longer than the transverse width (Figs. 26, 27a). Lingually there are






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


Table 6. Dental measurements (mm) forHypsitherium bolivianum from Inchasi, Pliocene of Bolivia.


Upper Ml M2 M3
dentition AP T AP T AP T

MNHN-Bol-V 12.4 6.9 14.5 6.2 11.8 6.2
003357



Lower p4 ml m2 m3
dentition AP T AP T AP T AP T

MNHN-Bol-V 11.2 4.2 11.6 4.15 12.7 3.8 unerupted
003356,
Holotype
AP; Anteroposteror length
T; Transverse width



relatively strong enamel folds situated about in the middle of each tooth; these are
better developed in ml and m2 than in p4. Labially, at about the position of the
lingual folds, there are distinctive grooves. The m3 is only partially erupted, and
therefore the characteristic dental pattern is not well exposed. Representative
dental measurements are presented in Table 6.
As represented in the referred specimen, MNHN-Bol-V 003357, the upper
molars are hypselodont, imbricate with one another, and have high, curved crowns
(Figs. 27B, 28). In occlusal view the molars have dentine surrounded by thin
enamel anterolingually and labially; posteriorly and posterolingually these teeth
either lack, or have very thin, enamel. The molars have a roughly triangular cross-
section consisting of large ectolophs with large, anterolabial projections; there also
are moderately developed protolophs and smaller metalophs. Lingually a deep
groove divides the protoloph and metaloph. Posterior to the metaloph there is a
well-developed lingual groove in m2; this structure is poorly developed in ml and
the relevant region in m3 is fragmentary.
Discussion.--Several recent publications describe the presence of the family
Mesotheriidae in South America. Francis (1960, 1965; in Pascual 1967) presented
a revision of what have been considered to be the valid genera from the Miocene-
Pleistocene of Argentina, including Eutypotherium Roth, 1901, Typotheriopsis
Cabrera & Kraglievich, 1931; Pseudotypotherium Ameghino, 1904 and
Mesotherium Serrds, 1867. To this list Villarroel (1974a; b) described the new






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 c

cm


Figure 26. Dorsal (A), lateral (B), and medial (C) views of left mandible ofHypsitherium bolivianum, new
genus and species, MNHN-Bol-V 003356, holotype.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


A
A -----


ant 0 2
t cm
lab


B ^ } lab





Figure 27. Occlusal views of(A) left lower (p4-m2; m3 unerupted) and (B) right upper dentitions (M1-M3)
ofHypsitherium bolivianum, new genus and species, respectively, 003356 holotypee) and MNHN-Bol-V
003357. Abbreviations: ant, anterior, lab, labial.



genera Plesiotypotherium and Microtypotherium from the Miocene of the
altiplano of Bolivia. Unlike the stratigraphic distribution of this family in
Argentina, Villarroel (1974a; 1974b) indicated that the Mesotheriidae did not
extend upwards into the Plio-Pleistocene (as currently calibrated) of Bolivia.
Many of the diagnostic characteristics that define this family and serve to
separate the valid genera of mesotheres are based on the incisor and cranial
morphologies, which are not represented in the sample from Inchasi.
Nevertheless, there are several diagnostic characters that justify the new genus
Hypsitherium. With respect to Plesiotypotherium, the mandible of Hypsitherium is
relatively deeper beneath ml and has an ascending ramus that forms an angle of
about 450 relative to the horizontal ramus, in contrast to about 900 in the former
genus. In the upper cheek teeth of Hypsitherium, the third, relatively well-
developed posterior lobe seen in Plesiotypotherium, Microtypotherium, and the
Argentine mesotheriines is absent. In the lower cheek teeth the lingual grooves
and labial folds are significantly more pronounced in Hypsitherium than in
Plesiotypotherium, Microtypotherium, and the Argentine mesotheriines.
We are confident that the two specimens from Inchasi represent a new genus
and species of notoungulate. However, we are less confident about its allocation to
an existing family. Taken together, the dental characters, in particular the
molarized p4, and the number, position, and relative development of the lingual
folds in the upper molars and lingual grooves and labial folds in the lower molars,
are most similar to diagnostic mesothere characters (Francis in Pascual 1967;
Villarroel 1974a, 1974b). There are, however, two seemingly important







ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


0 1
S c
cm


fi Ih.~


4





)LN


, d -
jc "-)''^ C'*4


Figure 28. Occlusal (A) and medial (B) views of right M1-M3 ofHypsitherium bolivianum, new genus and
species, MNHN-Bol-V 003357, referred specimen from Inchasi.


~~P~~Jt~C~h'P






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


exceptions to the mesothere dentition found in Hypsitherium. Firstly,
Hypsitherium lacks a well-developed third, posterolingual lobe in the upper molars.
Secondly, the number of lower premolars in Hypsitherium differs from mesotheres.
Previous descriptions of the dental formula for this family state that mesotheres
lack the pl-p3 (Francis 1965; Pascual 1967; Villarroel 1974a, 1974b). In contrast,
the lower jaw of Hypsitherium from Inchasi has alveoli for p2-p3 (the condition of
the pl is unknown). Despite these two morphological exceptions, in all other
dental characters Hypsitherium is most similar to mesotheres. Taking into account
the overall dental morphology, this genus differs significantly from that exhibited
in the two other typothere notoungulate families, i.e. Interatheriidae and
Hegetotheriidae (sensu Pascual 1967).
In summary, the specimens from Inchasi clearly represent a new genus and
species, Hypsitherium bolivianum, which is most closely referable to the existing
family Mesotheriidae. We cannot, however, rule out the possibility that
Hypsitherium belongs to a new, currently undescribed family, but we are not
prepared to propose one with the limited material available to us from Inchasi. If
indeed Hypsitherium is a mesotheriid, then its occurrence at Inchasi extends the
range of this family upward into the Pliocene of Bolivia, as the range of this family
is also known from Argentina. Although the latest genus of this family,
Mesotherium, is known from the Pleistocene of Argentina, no mesotherines are
currently known from this epoch in Bolivia.

DISCUSSION
Faunal Composition, Diversity, and Alpha-level Taxonomy

The Inchasi local fauna consists of 4 orders, 9 families, 10 genera, and at least
11 species of mammals. The diversity within these orders is similar to other
Pliocene sites from Bolivia (e.g. Ayo Ayo and Viscachani; Marshall et al. 1983;
Hoffstetter 1986) and Argentina (e.g. Tonni et al. 1992). However, no Pliocene
site from Bolivia has ever been as thoroughly sampled as, for example, the
extensive fauna from the Pleistocene of Tarija (Boule and Thdvenin 1920;
Hoffstetter 1963) or some of the classic late Cenozoic localities from the Province
of Buenos Aires and environs in Argentina (Marshall et al., 1984). There are some
notable absences in the Inchasi fauna, including non-mammalian vertebrates and
carnivorous mammals. In addition, micromammals are poorly represented. We
believe that these absences are an artifact of the available sample and the
underrepresentation results from the lack of screenwashing. Although four field
trips (amounting to about 50 person-days) have been spent recovering fossils by
surface prospecting at Inchasi, the discoveries to date must be viewed as
preliminary (this is in contrast to four centuries of sporadic, sometimes intensive
collecting at Tarija, for example). Given time for erosion to expose additional
fossils in this limited collecting area, we anticipate that additional surface
prospecting, and possibly screenwashing, for fossils will increase both the available






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


samples of currently poorly known taxa (e.g. Megatheriinae and Mylodontinae),
allowing more precise taxonomic allocations, and the recorded diversity both for
other mammals (most notably, from the orders Marsupialia, Carnivora, and
Rodentia) and probably the other vertebrate classes as well. The relatively low
diversity of the herbivores from Inchasi is similar to the known diversity of modern
mammalian faunas from the Bolivian altiplano (Anderson 1993; Teresa Tarifa,
pers. comm. 1994).
Four of our taxonomic allocations have been made to the level of genus, with
the species being indeterminant; these include Plohophorus, Paraglyptodon,
Caviodon, and Promacrauchenia. The two main reasons for this level of
allocation are either (1) there have been no comprehensive previous taxonomic
studies of these genera and species allocations from other Bolivian Pliocene sites;
and/or (2) the valid species for comparison from the classic, or better-known, sites
in Argentina are poorly characterized, or consist of portions of the skeleton not
present in the collections examined during this study. With regard to the latter, the
original descriptions of many glyptodont taxa are based primarily on caudal
morphology, whereas our specimens from Inchasi consist of carapace osteoderms.
Two other allocations could only be made to subfamily, i.e. Megatheriinae and
Mylodontinae, because the particular skeletal elements collected from Inchasi are
non-diagnostic at a more specific taxonomic level. Undoubtedly, future discoveries
of more characteristic elements, most notably teeth, and comparisons with
descriptions from Argentina will further refine these subfamilial allocations.
In summary, the current description of the mammalian fauna from Inchasi is
considered provisional. However, given the fact that all previous accounts of
Pliocene mammals from Bolivia either consist of preliminary faunal lists (e.g.
Hoffstetter et al. 1971), detailed descriptions of single taxa (e.g. Hoffstetter et al.
1984), or general taxonomic overviews (Marshall et al. 1983; Hoffstetter 1986),
our current description of the entire fauna from Inchasi, as it currently is known, is
therefore justified and of significance.

Paleoecology and Paleoenvironmental Reconstructions

A striking characteristic of middle Tertiary (Oligo-Miocene) mammalian
faunas from Bolivia and Argentina is the abundance of numerous taxa with either
very high-crowned (hypsodont), or ever-growing (hypselodont) teeth. This
predominance of high-crowned, presumed grazers, suggests a period during the
middle Tertiary of extensive grasslands as the predominant habitat and food
resource for the endemic notoungulates.
The relative tooth crown heights and mix of medium- to large-bodied
mammalian herbivores at Inchasi generally indicates local vegetation types and
broad climatic parameters. Based on analogies with other members of their
respective taxonomic groups, and/or the presence of short-crowned (brachyodont)
teeth, the occurrence of Mylodontinae, Megatheriidae, and the litopter






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 39(3)


Promacrauchenia at Inchasi suggest browsing and hence, the presence of either
scrub vegetation, woodlands, or forests. The diets and/or habitat preferences of the
other edentates (i.e. glyptodonts and armadillos) from Inchasi are difficult to
determine. At the other end of the spectrum, the presence of Hypsiitherium
bolivianum and two species of Posnanskytherium, all with hypselodont teeth and
the latter genus with a transversely linear cropping mechanism, suggest grazing to
hyper-grazing adaptations (e.g. Janis and Ehrhardt 1988). MacFadden et al.
(1994) has recently suggested from analysis of carbon isotopes of tooth enamel that
Posnanskytherium from Inchasi fed upon C3 vegetation, in this case probably a
predominance of montane grasses. Thus, so far as we know, the mammalian
herbivores from Inchasi probably represent a diversity of feeding adaptations
ranging from browsing to grazing. This balance and mixture of herbivores is in
contrast to Oligocene and Miocene mammalian faunas from Bolivia and elsewhere
in South America in which there usually is a strong predominance of grazers. The
smaller proportion of grazers in the Pliocene of Bolivia seems related to the
presence of different montane habitats resulting from Andean uplift.

Biogeography, Pliocene Regional and Continental Correlations,
and the Problematic Montehermosan/Chapadmalalan Interval

The fauna from Inchasi consists of 10 genera of Pliocene mammals. Eight of
these (Plaina, Glossotheridium, Proscelidodon, Plohophorus, Paraglyptodon,
Caviodon, Chapalmaltherium, and Promacrauchenia) are relatively widespread in
that they also are known from similar-aged deposits in Argentina. Only the genus
Posnanskytherium and its two species, P. desaguaderoi and P. inchasense, and the
new genus and species Hypsitherium bolivianum are unknown elsewhere and thus
appear to be endemic to the Bolivian altiplano. At first glance this high degree of
faunal similarity and low endemism is perhaps surprising. However, during the
middle Pliocene about 3-4 myr ago when these mammals lived, Inchasi probably
was at an elevation of about 2000 m and not 3200 m as it is today (MacFadden et
al. 1994). While high elevations of 3000 m or greater can effectively act as
biogeographic barriers to dispersal because of climatic effects and different
vegetation, lower elevations are less of a hinderance to dispersal. Take for
example, the middle Pleistocene mammalian fauna from Tarija in southern Bolivia
(e.g. Hoffstetter 1963), which today is at about 2000 m elevation. With its position
in the sub-Andean belt, the Tarija basin has probably experienced negligible uplift
(MacFadden et al. 1994) within the past 1 myr (when the fossiliferous Tarija
Formation was deposited). Its faunal composition both at the generic and in many
cases, specific, levels is almost identical to similar-aged faunas from Argentina.
This indicates an active dispersal corridor with regions at other latitudes in South
America during the Pleistocene. A similar situation probably was in existence at
Inchasi during the Pliocene.






ANAYA & MACFADDEN: PLIOCENE MAMMALS FROM BOLIVIA


The high degree of faunal similarity with classic and other well-known
Pliocene sites from Argentina allows an opportunity to assess the age of Inchasi
with regard to the Pliocene South American land-mammal ages (SALMAs). Two
names are generally applied to Pliocene SALMAs, i.e. the early Pliocene
Montehermosan and the middle Pliocene Chapadmalalan, both of which are based
on faunas from Argentina. Although the beginning of the Montehermosan and
end of the Chapadmalalan are fairly well accepted at, respectively, 5 and 2.5 Ma,
the boundary between these two land mammal ages is very poorly characterized.
There are two current hypotheses: (1) Some workers (e.g. Marshall et al. 1983)
have challenged the biochronological distinctness of the Chapadmalalan and argue
that both names should be included in an expanded concept of the Montehermosan
than spans the early to early-late Pliocene from 5 to 2.5 Ma. Assuming that these
two land mammal-ages are distinct, what can be said about the biochronological
assemblage of widespread taxa at Inchasi? Following Marshall et al. (1983), the
genera Proscelidodon and Plohophorus from Inchasi are restricted to the
Montehermosan s. s. as it is known from Argentina. The genera Glossotheridium
and Paraglyptodon are restricted to the Chapadmalalan. The longer-ranging
genera Plaina, Promacrauchenia, Chapalmatherium, and possibly Caviodon are
known from both the Montehermosan s. s. and Chapadmalalan. Based on these
biochronological occurrences, the Inchasi local fauna suggests that the
Monterhermosan s. I. is preferred over distinct Montehermosan s. s. and
Chapadmalalan SALMAs. (2) In contrast, Cione and Tonni (1994) argue that,
based on biochronological and magnetostratigraphic criteria, the Montehermosan
and Chapadmalalan are distinct land-mammal ages. In their study they cite the
importance of Inchasi to the resolution of these two biochrons. Their basic
argument is that most of the widespread Chapadmalalan s. s. index genera, i.e.
Glossotheridium, Plaina, Plohophorus, Paraglyptodon, and Chapalmatherium, are
found at Inchasi. Furthermore, Inchasi is of prime importance to resolution of the
definition of the land-mammal ages and Pliocene time in South America because
this locality clarifies some of the problems with the discontinuous stratigraphic
sections associated with the typical faunas from Argentina. Given their careful
redefinition of these two SALMAs, we favor the conclusion of distinct
Montehermosan and Chapadmalalan SALMAs proposed by Cione and Tonni
(1994).

SUMMARY AND CONCLUSIONS

Prior to the discovery of fossil mammals at Inchasi, knowledge of Pliocene
mammals of Bolivia was restricted to several sites in the northern altiplano, most
notably from Ayo Ayo, Viscachani, and Umala, but also including the poorer
known Alto Segiiencoma, Gualberto Villarroel, Achocalla, and Pomata (e.g.
Marshall et al. 1983; Hoffstetter 1986). Furthermore, although preliminary faunal







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL 39(3)


lists have been reported for these localities, only a few of the fossil mammals had
been described in detail.
The mammalian fauna from Inchasi consists of 4 orders, 9 families, 10 genera,
and at least 11 species of extinct mammals. Despite its somewhat fragmentary
representation, and difficult taxonomic assignments for some of the specimens,
description of this fauna represents a significant contribution to our knowledge of
Pliocene mammals from Bolivia. In the interpretation of Marshall et al. (1983),
the Inchasi mammals have faunal elements characteristic of both the
Montehermosan and Chapadmalalan SALMAs and thus are referred to as
Montehermosan s. 1. In contrast, the preferred interpretation here follows the
analysis of Cione and Tonni (1994), who recognize a distinct Chapadmalalan s. s.
SALMA; it is therefore concluded that the Inchasi mammalian fauna is
Chapadmalalan. In addition, previous magnetostratigraphic correlation of the
Inchasi beds to the Geomagnetic Polarity Time Scale further refines the age of this
fauna to middle Pliocene, between 4.0 and 3.3 myr ago (MacFadden et al. 1993).
We realize that the Inchasi fauna is only a first step in describing the
interesting mammalian communities that existed in the Bolivian Andes just prior
to the Great American Interchange. Further work at Inchasi and other high-
elevation Pliocene sites will add to our knowledge of some of the more poorly
represented taxa, allow further taxonomic refinements with respect to previous
specimen descriptions from elsewhere in Bolivia as well as from classic Pliocene
localities in Argentina, and provide further data relevant to refining the
traditionally problematic Montehermosan and Chapadmalalan SALMAs.

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Highly recommended as a guide is the CBE Style Manual, 3rd Edition 1972 (American Institute of
Biological Sciences, Washington, D.C.).
Manuscripts must be submitted in duplicate and satisfy the following minimal requirements. They
must be dwhle.Maced throumout including tables, figure captions, and literature citations. Figure
legends and tables should be typed on separate sheets. Also, please submit a copy of the complete text,
tables, figure captions, and literature cited on a floppy disk (software used does not matter, but we use
Word for Windows).
All illustrations are referred to as figures. They must comply with the following standards:
Photographs should be sharp, with good contrast, and printed on glossy paper, or the originals submitted.
If the background of photographs (especially those of specimens) is not desired, amberlith should be cut
out and used to cover the background. Drawings should be made with dense black waterproof ink on
quality paper or illustration board. All figures should have a cover sheet All lettering will be medium
weight, sans-serif type (e.g. Future Medium, News Gothic) in cutout, dry transfer, or lettering guide
letters. Make allowance so that after reduction no lower case letter will be less than 1 mm high (2 mm is
preferred) nor any capital letter greater than 5 nun high. The maximum size for figures is 9" x 14" (twice
BULLETIN page size); figures should not be less than typepage width (4A"). With soft lead pencil, on
the back of each illustration, designate the top and identify each by author's name, manuscript title, and
figure number.
All manuscripts not submitted in BULLETIN format will be returned to the author for retyping
and/or formatting.

Manuscripts, all editorial matters, and requests for more detailed preparation instructions should be
addressed to:

Managing Editor of the BULLETIN
Florida Museum of Natural History
University of Florida
P. O. Box 117800
Gainesville FL 32611-7800, U.S.A.







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Last updated October 10, 2010 - - mvs