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Pal&ontologische Zeitschrift 2008, Vol. 82•2, p. 139-152, 30-06-2008
New Glyptodont from the Codore Formation (Pliocene), Falc6n State, Venezuela, its relationship with the Asterostemma problem, and the paleobiogeography of the Glyptodontinae ALFREDO A. CARLINI, La Plata; ALFREDO E. ZURITA, La Plata; GUSTAVO J. SCILLATO-YANI~, La Plata; RODOLFO S,&,NCHEZ, Urumaco & ORANGEL A. AGUILERA, Coro with 3 figures
CARLINI, A.A.; ZURITA,A.E.; SCILLATO-YANI~,G.J.; S.~NCHEZ,R. & AGUILERA,O.A. 2008. New Glyptodont from the Codore Formation (Pliocene), Falc6n State, Venezuela, its relationship with the Asterostemma problem, and the paleo- biogeography of the Glyptodontinae. - Palaontologische Zeitschrift 82 (2): 139-152, 3 figs., Stuttgart, 30. 6. 2008. Abstract: One of the basal Glyptodontidae groups is represented by the Propalaehoplophorinae (late Oligocene - mid- dle Miocene), whose genera (Propalaehoplophorus, Eucinepeltus, Metopotoxus, Cochlops, and Asterostemma) were initially recognized in Argentinian Patagonia. Among these, Asterostemma was characterized by its wide latitudinal distribution, ranging from southernmost (Patagonia) to northernmost (Colombia, Venezuela) South America. How- ever, the generic assignation of the Miocene species from Colombia and Venezuela (A.? acostae, A. gigantea, and A. venezolensis) was contested by some authors, who explicitly accepted the possibility that these species could corre- spond to a new genus, different from those recognized in southern areas. A new comparative study of taxa from Argen- tinian Patagonia, Colombia and Venezuela (together with the recognition of a new genus and species for the Pliocene of the latter country) indicates that the species in northern South America are not Propalaehoplophorinae, but represent the first stages in the cladogenesis of the Glyptodontinae glyptodontids, the history of which was heretofore restricted to the late Miocene - early Holocene of southernmost South America. Accordingly, we propose the recognition of the new genus Boreostemma for the species from northern South America and the restriction of Asterosternrna to the Mio- cene of Patagonia. Thus, the available data indicate that the Glyptodontinae would in fact have arisen in the northern- most regions of this continent. Their arrival to more southerly areas coincides with the acme of the "Age of Southern Plains". The Propalaehoplophorinae are geographically restricted to Patagonia. Keywords: Boreostemma pliocena n. gen., n. sp. • Glyptodontinae • Propalaehoplophorinae ° paleobiogeography ° phylogeny ° South America Kurzfassung: Eine der basalen Gruppen der Glyptodontidae stellen die Propalaehoplophorinae (spates Oligozan - mittleres Miozan) dar, deren Gattungen (Propalaehoplophorus, Eucinepeltus, Metopotoxus, Cochlops und Aste- rostemma) man zuerst aus dem argentinischen Patagonien kannte. Darnnter ist Asterostemma dutch eine weite latitu- dinale Verbreitung gekennzeichnet, welche sich vom siidlichsten (Patagonien) zum ntrdlichsten (Kolumbien, Vene- zuela) Stidamerika erstreckt. Allerdings wurde die Gattungszugehtrigkeit der miozanen Arten Kolumbiens und Ve- nezuelas (A. ? acostae, A. gigantea und A. venezolensis) von einigen Autoren angezweifelt, die explizit die M6glich- keit in Betracht zogen, dass diese Taxa einer anderen Gattung angehtren, die sich vonder Gattung der stidlichen Brei- ten unterscheidet. Ein neuer Vergleich der Taxa aus Patagonien, Kolumbien und Venezuela (zusammen mit einer neuen Gattung und Art aus dem Plioz~in Venezuelas) zeigt, dass die Arten aus dem ntrdlichen Stidamerika nicht zu den Propalaehoplophorinae gehtren, sondern die ersten Stufen in der Kladogenese der glyptodontinen Glyptodonti- dae darstellen. Deren Geschichte war bislang auf das spate Miozan - frtihe Holozan des siidlichsten Siidamerikas be- schrankt. Dementsprechend stellen wir hier die neue Gattung Boreostemma ftir die Arten des ntrdlichen Sttdameri- kas auf; Asterosternma wird auf das Miozan Patagoniens beschrankt. Die Datenlage deutet somit darauf hin, dass Glyptodontinae sich tatsachlich im ntirdlichsten Teil S~idamerikas entwickelten. Ihr Auftreten in den stidlichen Ge- bieten fallt mit dem Hthepunkt des ,,Zeitalters der stidlichen Ebenen" zusammen. Die Verbreitung der Propalaehop- lophorinae wird geographisch auf Patagonien beschrankt. Schliisselwtrter: Boreostemma pliocena n. gen., n. sp. • Glyptodontinae ° Propalaehoplophorinae • Palaobiogeographie ° Phylogenie ° Siidamerika
Addresses of the authors: Alfredo A. Carlini, Pal~iontologisches Institut und Museum, Universit~it Ziirich, Karl Schmid-Stra- Be 4, CH-8006 Ztirich, Switzerland, and Departamento Cientffico Paleontologfa de Vertebrados, Museo de La Plata, Paseo del Bos- que s/n B1900FWA La Plata, Argentina. CONICET. C~ttedra de Anatomfa Comparada, Facultad de Ciencias Naturales y Mu- seo, Universidad Nacional de La Plata, Argentina; e-mail
0031-0220/08/0082-139 $ 6.30 © 2008 E. Schweizerbart'scheVedagsbuchhandlung, D-70176 Stuttgart 140 ALFREDO A. CARLINI et al.
Introduction pressa) (see SCOTT 1903-1904). In this context, a recent phylogenetic analysis of the southern Propalaehoplo- Among the particular palaeofauna that inhabited South phorinae (Propalaehoplophorus, Eucinepeltus, Mete- America during much of the Cenozoic, the cingulate su- potoxus, and Cochlops) has suggested that these taxa perfamily Glyptodontoidea are one of the most conspi- can be regarded as a monophyletic group (FERNICOLA cuous groups; several authors have recognized more 2006). than 65 genera during their diversification (late Eocene The "Friasian" lapse (middle - late Miocene) - early Holocene) (see MCKENNA & BELL 1997). In this (which comprises three successive "faunas", Friasian context, the Glyptodontidae Propalaehoplophorinae s. s., Colloncuran and Mayoan; SCILLATO-YANI~8z CAR- (sensu CASTELLANOS 1932) are among the best charac- LINI 1998) signals the beginning of the Panaraucarian terized basal clades (together with the Glyptatelinae; Faunal Cycle, Protoaraucarian Subcycle (Friasian - late Eocene - late Miocene; SCILLATO-YANt~ 1977; Chasicoan, of PASCUAL t~¢ ORTIZ JAUREGUIZAR 1990; CARLINI et al. 1997; VIZCMNO et al. 2003), and have PASCUAL et al. 1996). The Xenarthra, especially the been proposed by several authors as potential structural large cursorial Tardigrada (such as Megatheriidae, My- ancestors of later Glyptodontidae (SCOTT 1903-1904; lodontidae, and Scelidotheriinae; see Me DONALD SC[LLATO-YANg 1977). 1987) become larger and more frequent during this pe- The propalaehoplophorine glyptodonts are mor- riod (PASCUAL et al. 1996; ORTIZ JAUREGUIZAR t~z phologically characterized by the following traits: a) CLADERA 2006). This phenomenon could be associated low elongated skull, partly resembling those of dasypo- with the progressive aridization of the Southern Cone of did cingulates; b) dorsal carapace with mobile bands on South America, as a consequence of the Andean-Pata- the lateral margins (no less than 6 rows in Propalaeho- gonian Orogeny (see PASCUAL & BONDESIO 1982; PAS- plophorus and Cochlops), which is probably the primi- CUAL 1984; PASCUAL et al. 1984; ORTIZ JAUREGUIZAR tive condition for the Glyptodontidae; c) ornamentation 8,: CLADERA 2006; TONNI & CARLINI in press). This cli- of dorsal carapace osteoderms consisting of a polygonal matic-environmental context characterizes the last southern records of Propalaehoplophorinae (Propalae- central figure completely encircled by a row of smaller hoplophorus? and Eucinepeltus) (BONDESIOet al. 1980), peripheral figures, forming a rosette; d) caudal armor as well as the first records of Hoplophorinae, with the formed by a series of rings (generally four to six), each tribes Hoplophorini? (Eonaucum colloncuranun SCIL- ring formed by two rows of osteoderms, followed by a LATO-YANI~ & CARLINI, 1998) and Palaehoplophorini short caudal tube; e) five digits in both fore- and hind- (Palaehoplophorus AMEGHINO and Palaehoplopho- limbs; f) first three upper and lower molariforms sim- roides SCILLATO-YANt; & CARLIN 0 (PAULA COUTO ple, subcircular or subelliptic in transversal section. 1979; SCILLATO-YANI~" 8z CARLINI 1998). Many of these characters could be considered primitive; Argentinian Patagonia seems to have been, at however, some of them (c-d) do not occur in the Glypt- least until the "Friasian" (middle to late Miocene), the atelinae, or there is no evidence of their occurrence. geographic location where the cladogenesis and radia- From a chronological viewpoint, the subfamily tion of this group is best recorded (PASCUAL & Propalaehoplophorinae is first recorded during the De- ODREMAN RIVAS 1973; PASCUAL et al. 1984), consid- seadan (late Oligocene) (SCILLATO-YANI~ 1977), a time ering the good record of cingulates in the area (SCIL- of remarkable increase in glyptodontid diversity. The LATO-YAN~ 1986). Furthermore, PASCUAL et al. (1984) Deseadan also marks the end of the records of Glypt- and PASCUAL & ODREMAN RIVAS (1973) went so far as atelinae at high latitudes; however, they persist in more to propose this territory as a center of radiation and northern areas, until the middle Miocene in Colombia cladogenesis. (CARLINI et al. 1997; VILLARROEL 8¢ CLAVIJO 2005) Recently, other possible evolutionary scenarios and until the late Miocene in Uruguay (VIZCAiNO et al. have been proposed for the Cingulata at lower latitudes, 2003). They might even have occurred in the early These hypotheses are especially supported by fossil re- Pleistocene of North America (MCKENNA & BELL mains from the Honda Group ("Friasian") in the fossil 1997). locality of La Venta, Colombia (CARLINI et al. 1997). During the Colhuehuapian (early Miocene), and es- Several taxonomic and paleobiogeographic differences pecially during the Santacrucian (early Miocene), the between these materials and isochronous Patagonian Propalaehoplophorinae show great diversification, evi- paleofaunas support these hypotheses. These include denced in the recognition of five genera and eight spe- the remarkable survival of a representative of the glypt- cies in Argentinian Patagonia during this latter South ateline Glyptodontidae (Neoglyptatelus originalis) with American Land Mammal Age (SALMA): Propalaeho- very primitive characters (CARLINI et al. 1997), as well plophorus AMEGHINO, 1887 (P. australis and P. minor), as records of astegotheriine, dasypodine, and tolypeu- Cochlops AMEGHINO, 1889 (C. muricatus and C. debi- tine Dasypodidae, which are absent in Patagonia; in lis), Eucinepeltus AMEGHINO, 1891 (E. complicatus and contrast, representatives of Peltephilidae, Euphractini, E. petestatus), Metopotoxus AMEGHINO, 1898 (M. an- and Stegotheriini occur in the latter region (CARLINI et ceps), and Asterostemma AMEGH1NO, 1889 (A. de- al. 1997). New Glyptodont from the Codore Formation and the paleobiogeography of the Glyptodontinae 141
Within this paleobiogeographic context, the Mio- berifer from the "Araucanian"; late Miocene - Pliocene) Pliocene glyptodontid fauna in northern South America are not very frequent or diverse in late Neogene sedi- (Colombia and Venezuela), which remained unknown ments; only two species, Paraglyptodon chapalmalen- until the mid-1940s (see SIMPSON 1947), seems to agree sis from the middle Pliocene (Chapadmalalan) of the with this hypothesis. Traditionally, all the Miocene glypt- Pampean region and Paraglyptodon uquiensis CASTEL- odonts from this area were included within the subfamily LANOS, 1953, from the middle - late Pliocene (late Cha- Propalaehoplophorinae, and more precisely, within the padmalalan - Sanadresian) of the current territory of Ju- genus Asterostemma. Thus, three species have been de- juy province (see REGUERO et al. 2007), both from Ar- scribed, A. venezolensis SIMPSON, 1947, for the (mid- gentina, have been described for that period. However, dle?) Miocene of Venezuela, and A.? acostae VILLAR- the Glyptodontinae become very frequent in southern ROEL, 1983 and A. gigantea CARLINI et al., 1997, for the sediments since the Ensenadan (late Pliocene - early middle Miocene (Laventan SALMA) of Colombia. The Pleistocene; ca. 1.7 - 0.98 Ma) of the Pampean region, inclusion of these taxa in the southern genus Aster- achieving both noticeable taxonomic richness and exten- ostemma was accepted by most authors, though with the sive latitudinal distribution that ranged from the south of explicit acknowledgment that these taxa could in fact Buenos Aires province to, probably, Venezuela (see represent a new genus, different from the ones tradition- MARSHALL et al. 1984; TONNI & SCILLATO-YANI~ 1997; ally recognized for the Patagonian Miocene (i.e., RINCON 2006). Thus, approximately eight Glyptodon Propalaehoplophorus, Eucinepeltus, Metopotoxus, species have been recognized for the Pleistocene of Cochlops, and Asterostemma) (see SIMPSON 1947: 9; southernmost South America (SOIBELZON et al. 2006; VILLARROEL 1983: 30; CARLINI et al. 1997: 223). CARLINI & SCILLATO-YANt~ 1999). In this contribution we present a new Glyptodonti- Similarly, this subfamily is very well represented dae from the Codore Formation (Pliocene) of Falc6n in North America since the early Blancan (ca. 2.6 Ma), State, Venezuela (Fig. 1); concurrently, on the basis of that is, immediately after the establishment of a contin- both morphological and paleobiogeographic evidence, uous Panamanian terrestrial connection (ca. 2.7 Ma), we propose the recognition of a new genus for all the with the genus Glyptotherium (see CARLINI et al. 2008). Mio-Pliocene species recognized in northern South However, recently CARRANZA-CASTAIqEDA & MILLER America (Colombia and Venezuela) that have tentatively (2004) cited the presence of Glyptotherium OSBORN, been assigned to the southern genus Asterostemma. 1903 in Pliocene sediments from central Mexico, dated The new Glyptodontidae described herein allows by them at 3.9 Ma, although the generic assignation is the proposal of a different evolutionary scenario for the not secure, given that the authors themselves (2004: species from northern South America; particularly, it 258) highlight the marked differences with respect to allows us to elaborate a different hypothesis for the dif- known Glyptotherium species. It would not be surpris- ferentiation of the glyptodontine Glyptodontidae. This ing, if these Mexican findings were to correspond in subfamily is known from the "Araucanian" (late Mio- fact to a new taxon close to the one described here. cene - Pliocene) of NW Argentina, represented by Glypt- Abbreviations: AMNH, American Museum of Natural His- odontidium tuberifer CABRERA, 1944, a species with tory, New York, USA; AMU-CURS, Colecci6n de Paleonto- dorsal carapace formed by thick osteoderms with the logfa de Vertebrados de la Alcaldia de Ururnaco, Estado Fal- typical rosette comprising a central polygonal figure c6n, Venezuela; M, m, upper and lower molariform respec- and several marginal figures in a single row. Although tively; GABI, Great American Biotic Interchange; ICNUNC, this character would be primitive among the propalae- Instituto de Ciencias Naturales, Universidad Nacional de Co- hoplophorine and glyptodontine glyptodonts, the mor- lombia; IGM, Instituto de Geociencias y Mineria, Colombia ; n/n, without official catalog number; SALMA, South Ameri- phology of the sulci that delimit the pattern on the ex- can Land Mammal Age. posed surface of the osteoderms is different in cross section. While in the southern Propalaehoplophorinae the cross section of these sulci is parabolic ("V"-shaped), in the Glyptodontinae it is quadrangular ("U"-shaped), Geologic and stratigraphic context: with almost vertical sides and a wide, almost flat bot- the Codore Formation tom. In addition, the carapace osteoderms of glyptodon- tines are relatively thicker than those of propalaehoplo- The Codore Formation was recognized and described phorines, and the contact edges between successive by HODSON (1926) in Falc6n State, Venezuela, and as- osteoderms are definitely denticulated. In general, the signed chronologically to the Pliocene s.l., while always section of these sulci and the thickness of osteoderms, considered older than the overlying San Gregorio Fm. together with the greater lobation of the anterior molar- from the late Pliocene (GONZALEZ DE JUANA et al. iforms, are generally used to define the glyptodontine 1980). This geologic unit is divided into two (see LID- glyptodonts (HOFFSTETTER 1958; PASCUAL et al. 1966; DLE 1946) or three (see REY 1990) members, according PAULA COUTO 1979). to different authors. Lithologically, it comprises a suc- The history of the Glyptodontinae is rather particu- cession of ochraceous to dark brown sandstones with lar, given that their earliest records (Glyptodontidium tu- cross-stratification, and light-colored limolites with in- 142 ALFREDO A. CARLINI et al.
CARIBBEAN SEA
VENEZUELAN GULF Paleontological site Codore Fm. at ' Tio Gregorio p
100 km i
Fig. 1. Map showing the locality where the type specimen of Boreostemma pliocena n. gen., n. sp. was recovered.
tercalated carbonatic levels of varying thickness. The Systematic paleontology depositional environment has been interpreted by REY (1990) as characteristic of a distal fan-delta, with an al- Xenarthra COPE, 1889 luvial plain associated with small meandering channels, Cingulata ILLIGER, 1811 and some brief intervals of marine invasion halfway Glyptodontoidea GRAY, 1869 through the sequence. The presence of palynomorphs Glyptodontidae GRAY, 1869 allows to infer subhumid to dry climate throughout the Glyptodontinae GRAY, 1869 entire deposition of the Codore Fm. (LINARES 2004). The sequence is about 800 m thick and comprises sedi- Boreostemma n. gen. ments concordant with the underlying Urumaco Fm. (late Miocene). Recently, LINARES (2004), on the basis Etymology: Boreos from the Latin "borealis" "from the North", of the mammalian fauna, has assigned an age corre- in reference to its geographical distribution, restricted to nor- sponding to the Montehermosan SALMA (late Miocene thern South America, + "stemma" ending of Asterostemma, austral genus to which it was previously wrongly assigned. - early Pliocene) to the E1 Jebe Member of Codore Fm., Type species: Boreostemma pliocena n. gen., n. sp., from the and suggested a probable age of 6 Ma for the contact be- Codore Formation, E1 Jebe Member, Pliocene, northern Vene- tween the Urumaco and Codore formations. zuela. Referred species: The type species and Asterostemma vene- zolensis SIMPSON, 1947 (middle? Miocene, eastern Vene- zuela), Asterostemma? acostae VILLARROEL, 1983 [sic] New Glyptodont from the Codore Formation and the paleobiogeography of the Glyptodontinae 143
(middle Miocene, Huila Department, Colombia), Aste- tween the sulcus surrounding the central figure and the rostemma gigantea CARLINI et al., 1997 (La Victoria Forma- sulci between peripheral figures. Caudal rings imbri- tion, Honda Group, middle Miocene; Huila Department, Co- cated, formed by a single row of osteoderms. Caudal lombia). ring osteoderms with barely marked central figure and Diagnosis: Small to medium sized Glyptodontinae, without signs of peripheral figures, differing from ranging in size from that of Propalaehoplophorus aus- Propalaehoplophorus australis and Cochlops murica- tralis to Eucinepeltus crassus. Osteoderms arranged in tus (propalaehoplophorines whose osteoderms are well- transverse bands more regular than in Propalaehoplo- known), in which the central figure is clearly visible and phorinae, with rugose and very punctate surface. Cen- a row of peripherals is present at the proximal edge. tral figure polygonal, in a more central position. Sulci Posterior upper molariforms (at least M5-M7) very surrounding adjacent figures wide, which has (in cross close to each other, with almost no separation, differing section) nearly vertical walls and wide bottom, which from those of Propalaehoplophorinae, in which the sep- we call "U"-shaped, contrasting with the "V"-shaped aration between molariforms increases disto-proxi- sulci of Propalaehoplophorinae and Hoplophorinae ho- mally. Ornamentation of cephalic shield osteoderms plophorini with convergent walls and strait bottom. similar to that of dorsal carapace, as in Propalaehoplo- Area of contact and articulation of adjacent osteoderms phorus australis and the hoplophorine Hoplophorinae strongly denticulate, with deep osseous interdigitating Neosclerocalyptus PAULA COUTO, 1957, and very dif- projections. Generally, a single series of peripheral fig- ferent from that of Eucinepeltus complicatus and Meto- ures in each osteoderm, morphologically simpler than potoxus laevatus, which show no signs of rosette orna- those of Glyptodontidae Propalaehoplophorinae. Pe- mentation. ripheral figures of two or three adjacent osteoderms forming composite figures. Caudal rings imbricated, Description and assignation of material formed by a single row of osteoderms. Posterior region Palate: This is a posterior fragment with part of the al- of caudal tube with slightly imbricated osteoderms, veoli for molariforms M6 and M7 on the left side, and more similar to the caudal tube of Dasypodidae than to the last alveolar lobe of M6, that of M7 and the first of those of known Propalaehoplophorinae. Dorsal cara- M8 on the right side (Fig. 3). It is 53.5 mm long and pace formed by larger, thicker and fewer osteoderms 35 mm wide. The entire surface of the palate is pierced than in Propalaehoplophorinae. by foramina that are particularly evident on the midline. There are also six large foramina at the level of the first Boreostemma pliocena n. sp. lobe of M7, at the limit between M7 and M8, and at the Figs. 2A-D, 3 limit between the first and second lobes of M8. On the Etymology: "pliocena" from Pliocene, the age of the Codore right side, these foramina open at the level of the last Formation, Venezuela. lobe of M6, the second lobe of M7 and the first lobe of Holotype: Thirty complete, associated osteoderms, some M8. Apart from this, the particular arrangement of the fragmentary osteoderms, and a fragment of the palate with alveoli for the last molariforms shows virtually no sep- partial alveoli of M 5-7. aration between M7 and M8. This trait is clearly differ- • ~¢pe locality: Road to Tfo Gregorio locality, Urumaco, Es- ent from the condition in Propalaehoplophorinae, in tado Falc6n, Venezuela, 11° 15' 35" N and 70 ° 17' 30" W (Fig. which the separation between successive molariforms 1). increases disto-proximally. On the other hand, the mid- Stratigraphic range and geochronologic age: E1Jebe Mem- dle lobe seems to be more inclined with respect to the ber of the Codore Formation, Pliocene. longitudinal axis of the palate than in Propalaehoplo- Diagnosis: Osteoderms of dorsal carapace very large, phorus, Asterostemma, Eucinepeltus, and Cochlops, in with extremely punctuate dorsal surface, very similar in which this lobe is approximately at straight angles to the this respect to Boreostemma gigantea, B. acostae, B. anteroposterior axis of the dental series. venezolensis, Glyptodontidium tuberifer and Paraglypt- Osteoderms: Numerous osteoderms from the dorsal and contrasting with the condition ob- odon uquiensis, carapace, as well as some from the cephalic shield, have served in the southern Propalaehoplophorinae been preserved. They are categorized into five groups (Propalaehoplophorus, Eucinepeltus, Metopotoxus, according to their location on the dorsal carapace. and Asterostemma), which have a smoother exposed surface. Punctuation especially concentrated in a slight a) First group: these osteoderms correspond to the dor- central concavity at each central figure, more visible in sal or postero-dorsal region of the carapace (Fig. 2A). the largest (dorsal and posterodorsal) osteoderms. Sulci The larger ones are 51 mm by 47 mm in diameter, and dividing the central figure from the peripheral ones, and 14 mm thick, while the smaller are 39 mm by 33 mm and the latter from each other, wide and shallow, as in Glypt- 10 ram, respectively. Each osteoderm has five or six odontidiurn tuberifer, Paraglyptodon uquiensis and P. sides (in most) which are relatively isodiametric, with chapalmalensis; hair follicle (?) pits well developed and extremely punctate dorsal surface, as in Glyptodontid- situated (as in most of the Glyptodontidae with this ro- ium tuberifer, Paraglyptodon uquiensis, and Boreo- sette ornamentation pattern) at the intersections be- sternma acostae (n. comb.). The central figure is poly- 144 ALFREDO A. CARLINI et al.
A
B 't C
Fig. 2. Boreostemma pliocena n. gen., n. sp. A: Osteoderms corresponding to the dorsal or postero-dorsal region of the carapace (seven in external view, one in lateral view). B: Osteoderms from the sides of the carapace (nine in external view, one in lateral view). C: One osteoderm from the cephalic shield (external and lateral views). D: Os- teoderms from the caudal rings (two in external view, one in lateral view). - Scale bar = 2 cm. New Glyptodont from the Codore Formation and the paleobiogeography of the Glyptodontinae 145
16
/17
Fig. 3. Boreostemma pliocena n. gen., n. sp. Palate: stereopairs of a posterior fragment with part of the alveoli for molariforms M6 and M7 on the left side, and the last alveolar lobe of M6, that of M7 and the first of M8 on the right side. - Scale bar = 2 cm. gonal, with the number of sides coinciding with the raglyptodon chapalmalensis (see ROVERETO 1914; CAS- number of peripheral figures. The central figures of the TELLANOS 1953). A conspicuous foramen appears in larger osteoderms are between 23.5 by 21 mm, and some of the intersections between the sulcus that delim- 25 mm by 19 mm. The figures of the smaller osteoderms itates the central figure and the radial sulci; there are are 19 mm by 18 mm. The dorsal surface of the central three to eight of these foramina. The sulci separating the figures is slightly elevated with respect to the peripheral central figure from the peripherals, and the radial sulci ones, particularly in the larger osteoderms, whereas it is between peripherals are wide, shallow, with nearly flat flat in the smaller ones. As mentioned above, the entire bottom, and morphologically similar to those of Glypto- surface of each central figure is densely punctate (al- dontidium tuberifer. The sulci are similar in Paraglypto- though much less than in Glyptodon and Glyptotherium, don uquiensis and P. chapalmalensis, although in these in which the surface of the osteoderms is definitely ru- latter species the sulci are deeper and somewhat similar gose), but this condition becomes especially evident at its to those of the genera Glyptodon and Glyptotherium. central area, which bears larger-diameter foramina lo- The ventral surface of the osteoderms may be flat cated in a small concavity, somewhat similarly to the or slightly concave at the center. One to nine foramina morphology of Boreostemma acostae. As in the previ- are visible on this surface; generally the largest (one or ous case, this concavity is more conspicuous in the larger two of them) occupy a central position while the rest are osteoderms. Each central figure is surrounded by a row arranged peripherically in a circular pattern. of eight to ten peripherals, 4- or 5-sided, well developed and very angular. The number of peripherals seems to be b) Second group: osteoderms from the sides of the car- lower when compared to the southern Propalaehoplo- apace (Fig. 2B). These are rectangular with a disto-prox- phorinae forms; the latter may have up to twelve or thir- imal long axis, and have four to six sides. The largest are teen peripherals (see SCOTT 1903--1904). The entire sur- 37.5 mm by 23 mm and 14 mm thick, while the smallest face of these figures is clearly punctate, although the are 28 mm by 21 mm and 9.5 mm thick. The entire ex- punctuations situated toward the periphery are some- posed surface is visibly punctate, as in the dorsal and what radial in appearance (as they become oblique in- posterodorsal osteoderms. The central figure is flat, with stead of perpendicular to the surface), as observed in subelliptical to subcircular outline, and unlike the dorsal some osteoderms of Paraglyptodon uquiensis and Pa- osteoderms, does not show a concave area with larger fo- 146 ALFREDO A. CARLINI et al. ramina. The eight or nine peripheral figures are less de- the peripheral figures. As in the largest dorsal carapace veloped than in the dorsal osteoderms, and the lateral osteoderms, the largest punctuations are concentrated in ones are smaller. As in the Propalaehoplophorinae and the center of the figure, although with a slight tendency the most primitive hoplophorine Hoplophorinae (i.e., to occupy one of the margins. The twelve peripheral fig- Eosclerocalyptus tapinocephlaus and E. proximus), the ures are asymmetrically developed, and largest and anterior peripheral figures are larger than the posterior thickest at the margin/border. Eleven hair follicle (?) pits ones. The central and radial sulci are only slightly are clearly visible. marked, and the number of foramina ranges between The ventral surface of the osteoderm is notably three and six. concave at its center; much more so than any osteoderm from the first group, with three large foramina at the e) Third group: formed by very small osteoderms of the area of maximum thickness. dorsal carapace, possibly corresponding to the extreme antero4ateral and postero4ateral regions of the carapace. These osteoderms are quadrangular to hexagonal, and measure from 24 mm by 24.5 mm and 11 mm thick, to The problem of the genus Asterostemma 25 mm by 23 mm and 8.5 mm thick. The entire dorsal AMEGHINO in the context of the Glypto- surface has the same characteristics described above for dontidae / Propalaehoplophorinae other osteoderms, with three to five well-developed hair follicle (?) pits at the intersections between the sulcus sur- All the genera of propalaehoplophorine Glyptodonti- rounding the central figure and the sulci between periph- dae (Propalaehoplophorus, Asterostemma, Cochlops, eral figures, but in this case there is no differentiation be- Metopotoxus, and Eucinepeltus) were originally rec- tween central and peripheral figures. ognized on the basis of materials exhumed from the early and middle Miocene (Colhuehuapan, Santacruc- d) Fourth group: osteoderms from the caudal rings ian, and Friasian ages) of Argentinian Patagonia (see (Fig. 2D). These are clearly rectangular with a disto- AMEGHINO 1889, 1891; LYDEKKER 1894; SCOTT 1903-- proximal long axis. The largest osteoderms are 44 mm 1904; RUSCONI 1946, among others). Osteoderm orna- by 23.5 mm and 12 mm thick, while the smallest one is mentation in this subfamily is characterized by a cen- 34 mm by 20 mm and 9 mm thick. Each osteoderm tral figure encircled by a row of more-or-less angular shows two well-defined areas: a distal area with a flat peripheral figures, producing a characteristic rosette exposed surface, and a proximal area with an extremely pattern (HOFFSTETTER 1958; PAULA COUTO 1979) that rugose irregular surface, the latter corresponding to the remains almost unchanged in later groups (e.g., area of contact and articulation with the adjacent osteo- hoplophorine Hoplophorinae; see ZURITA et al. 2005). derm. The surface of the distal area is flat and punctate, This situation, together with the absence of isochronic although a disto-proximally elongated small convexity records in more northern areas, led some authors to was observed in the center of one of the osteoderms suggest that some of these Propalaehoplophorinae studied. In addition, these osteoderms bear a very weak were the ancestral stock for the remaining Glyptodon- sulcus that delimits a subcircular central figure, al- tidae lineages (SCOTT 1903-1904: 155). The diagnos- though there are no peripheral figures; this central fig- tic characters provided by AMEGHINO (1889: 822) for ure is 16 mm long by 15 mm wide in the smallest osteo- his genus Asterostemrna seem to be highly variable derms and 19 by 17 mm in the largest ones. This partic- within a single dorsal carapace, and occur also in the ular morphology differentiates this taxon from Propalae- remaining Propalaehoplophorinae species (e.g., num- hoplophorus australis and Cochlops rnuricatus, as both ber of sides of osteoderms in the dorsal carapace, size have a clearly visible central figure and a row of peri- of the central figure and development of peripheral pherals at the proximal margin. A series of foramina, figures). Later, SCOTT (1903--1904), as part of a sys- larger than those of the dorsal carapace osteoderms, is tematic revision of Propalaehoplophorinae, upheld the evident at the lateral and posterior margins. validity of the genus, but recognized only one of the Likewise, the proximal contact area is depressed three species originally described by AMEGHINO toward its proximal end, and has a very rugose surface (1889), namely as valid, whereas he syn- with large foramina. In lateral view, the osteoderms are A. depressa, onymized A. laevata and A. granata with Metopot- maximally thick at their center and become thinner to- oxus laevatus and Cochlops muricatus, respectively ward the edges, particularly at the proximal margin. (SCOTT 1903-1904). In addition, this author referred e) Fifth group: only one osteoderm from the cephalic new materials, particularly an isolated caudal armor, shield is well preserved. It is somewhat similar to those to A. depressa (but without providing an extensive ra- of Propalaehoplophorus, and clearly different from tionale for his decision). He also emphasized that: "The those of Eucinepeltus and Metopotoxus (Fig. 2C). The status of this genus is somewhat uncertain and more osteoderm is relatively isodiametric, 36 mm by 35 mm complete material is greatly to be desired...". (SCOTT and 13 mm thick. The central figure is subcircular 1903-1904: 154). Finally, RUSCONI (1946: 9-11), on (24 mm by 27 mm) and slightly elevated with respect to the basis of six associated dorsal osteoderms, recog- New Glyptodont from the Codore Formation and the paleobiogeography of the Glyptodontinae 147 nized a poorly-diagnosed new species, A. barraelense Paleobiogeography and morphology of RUSCONI, 1946, from the Miocene of San Juan prov- the propalaehoplophorine and glyptodon- ince. tine Glyptodontidae: a new scenario Recently, the genus Asterostemma was reported by GOIS-LIMA & COZZUOL (2005) and COZZUOL (2006) in Within this paleobiogeographic and morphologic frame- the upper Miocene of Acre (Brazil). Unfortunately, this work, a detailed analysis of the main traits that charac- Glyptodontidae material consists of one isolated osteo- terize the southern Propalaehoplophorinae forms indi- derm (not illustrated), and consequently its generic iden- cates that most of these traits do not occur in the more tification is uncertain. northern Mio-Pliocene forms (from Colombia and Vene- SIMPSON (1947) was the first to recognize and de- zuela). Thus, the southern Propalaehoplophorinae are scribe a new propalaehoplophorine in more northern re- characterized by: a) first two upper and lower molari- gions, a species from the middle Miocene of Venezuela forms simple, with subcircular or subelliptical outline, (Santa In6s Fm.) that he tentatively assigned to the genus and third molariform with slight indication of lobation Asterostemma AMEGHINO (A. venezolensis SIMPSON). in some species; b) in the dorsal carapace, the sulci sep- However, such traits as the evidently more complex arating the central figure from the peripherals (main morphology of the first lower molariforms, which are sulcus) and these from each other (radial sulci) are "V'- greatly elongated anteroposteriorly, the absence of mo- shaped in transversal section, a morphology that is con- bile bands in the antero-lateral region of the dorsal car- served in the hoplophorine Hoplophorinae; c) dorsal apace, and the particular ornamentation of the osteo- surface of the osteoderms smooth, without remarkable derms that delimit the caudal notch, do not occur in any features and, in many cases, with anterior accessory pe- of the southern taxa. This situation was recognized by ripheral figures; d) vestiges of imbrication between suc- SIMPSON himself (1947: 9), who stated that "There re- cessive osteoderm rows in the most anterolateral region mains, however, the distinct possibility that the Vene- of the dorsal carapace (as in Dasypodidae; see SCOTT zuelan specimen belongs in a distinct genus not defined 1903--1904). In addition, the monophyly of the southern from the Argentine". forms has been suggested by FERN[COLA et al. (2006), Later on, DE PORTA (1962) referred some isolated although the issue is problematic and there is no clear osteoderms from the middle Miocene of Colombia to consensus about it. SIMPSON's species (A. venezolensis) and to Propalaeho- As we have mentioned before, a new and more plophorus, but without explicit reasons for these assigna- thorough examination has shown that this combination tions. In 1983, VILLARROEL recognized and described a of characters does not occur in the taxa from northern new species from the Miocene of the same country, and, South America, which have been traditionally included like SIMPSON (1947), he assigned it provisionally to the in subfamily Propalaehoplophorinae on the basis of the genus Asterostemma (A.? acostae VILLARROEL [sic]). superficial similarity in the rosette ornamentation pat- More recently, CARLINI et al. (1997) described a tern of the dorsal carapace. new species from the middle Miocene (Laventan) of The type of Boreostemma venezolensis n. comb. Colombia, A. gigantea. Among the diagnostic charac- (n/n) shows a notable antero-posterior elongation and ters, the large size and evident complexity of the second greater complexity of the first two lower molariforms, lower molariform, which tends to be trilobed, are re- absence of mobile bands in the lateral region of the car- markable. Contrastingly, molariform lobation of the apace, and a particular ornamentation of the osteoderms southern forms is clearly defined from the third or adjacent to the caudal notch (SIMPSON 1947). fourth tooth. Following the ideas of SIMPSON (1947) and In Boreostemma acostae n. comb. (ICNUNC P- VILLARROEL (1983), CARLINI et al. (1997) stated that 225-1) the dorsal carapace osteoderms have a strongly "Asterostemma gigantea is clearly different from spe- punctate dorsal surface, particularly in the middle of the cies of Propalaehoplophorus AMEGHINO, Cochlops central figure; in addition, the sulci separating the cen- AMEGHINO, Eucinepeltus AMEGHINO, and Metopot- tral figure from the peripherals are wide and shallow oxus AMEGHINO. Regrettably, the carapace of the type (see VILLARROEL 1983). species A. depressa from the early Miocene of Patago- Other remarkable traits of Boreostemma gigantea nia is very poorly known. Like VILLARROEL (1983), we n. comb. (IGM 250928) are the large body sized and a cannot completely discredit the idea that the Propalae- beginning of trilobation already in the second lower hoplophorinae from northern South America, including molariform, which is morphologically more complex A. venezolensis SIMPSON, could belong to a new genus, than that of B. venezolensis, and much more so than in distinct from Asterostemma. In this case, Asterostemma the southern taxa. In addition, the carapace osteoderms would have a geographic distribution limited to Patago- are thick and lack secondary peripheral figures (CAR- nia. Into this new genus, we would include the species LINI et al. 1997). described herein." Finally, Boreostemma pliocena n. sp. (AMU- CURS 158) differs from the southern forms by the greater size and thickness of its osteoderms with strongly punctate dorsal surface, by its caudal ring osteoderms 148 ALFREDO A. CARLINI et al.
with the central figure only slightly marked and no trace gose in the terminal taxa, without secondary peripheral of peripheral figures, and by having the posterior upper figures; c) increasing osteoderm thickness; d) tendency molariforms (at least M6-M8) very close to each other of the articulation between osteoderms to become highly with almost no separation. denticulated; e) lobation of first upper and lower molari- In addition to this particular set of characters oc- forms. The first molariform is already trilobated in curring in these taxa, a detailed examination of the mor- Glyptodon (see SOIBELZON et al. 2006). phology of the sulci that separate the central figure from the peripherals and the latter from each other shows that it differs from the condition observed in the southern Discussion taxa (in which these structures are "V"-shaped in trans- versal section). Indeed, in these northern taxa, the trans- The Miocene Glyptodontidae of northern South Amer- versal section of these sulci has walls parallel to each ica (Colombia and Venezuela) have traditionally been other and an almost flat bottom. Additionally, the dorsal included within the Propalaehoplophorinae, and, more surface of the osteoderms is always strongly punctate, precisely, assigned to the genus Asterostemma, a taxon the osteoderms are thick and the area of articulation be- originally recognized from Argentinian Patagonia. tween them is notably denticulated. However, the rosette ornamentation pattern of the dorsal This notable morphological differentiation be- carapace osteoderms (which represented the main basis tween the Patagonian taxa and those from northern for these assignations) of southern Propalaehoplophori- South America agrees with the paleobiogeographic nae is not the same as that of the northern taxa that were scenario outlined by CARLINI et al. (1997) for the mid- originally assigned to this subfamily. In effect, as we dle Miocene. These authors have proposed other possi- stated, while the sulci shows a "V"-shaped transversal ble evolutionary scenarios for the Cingulata at lower lat- section in the Propalaehoplophorinae and hoplophorine itudes, given the distant phylogenetic relationship be- Hoplophorinae, this transversal section is clearly "U"- tween the taxa in both areas, among other elements. The shaped in the taxa from Colombia and Venezuela. This Dasypodidae that are present in northern regions in- is coupled to greater complexity of the anterior molari- clude the Astegotheriinae, Dasypodini, and Tolypeuti- forms, and possession of dorsal osteoderms with ex- nae, which are absent in Patagonia; on the contrary, the tremely punctate dorsal surface, greater thickness, and Peltephilini, Euphractini, and Stegotheriini occur in the markedly denticulated articulation surfaces. latter area (CARLINI et al. 1997). Accordingly, VIZCA~NO This morphologic differentiation between "south- et al. (1990) have suggested a probable intertropical ori- ern" and "northern" forms agrees with the paleobio- gin for the dasypodine Dasypodidae. geographic and evolutionary scenario proposed by Within the Glyptodontidae, the subfamily Glypto- SCILLATO-YANE (1986) and CARLINI et al. (1997) for dontinae maintains this primitive pattern of rosette or- the cingulate Xenarthra, which was also indirectly an- namentation ("Araucanian" - Lujanian) (see AMEGHINO ticipated by SIMPSON (1947) and VILLARROEL (1983), 1889; HOFFSTETTER 1958; PASCUAL et al. 1966; PAULA given the particular differences observed between most COUTO 1979). In fact, some authors (e.g., CABRERA Cingulata from these two areas. As we have previously 1944) have highlighted the notable similarity in dorsal discussed, in this context, the characters observed in carapace ornamentation between the earliest Glypto- Asterostemma venezolensis, A.? acostae, and A. gi- dontinae species (Glyptodontidium tuberifer; see CA- gantea justify their exclusion from the genus Astero- BRERA 1944) and the Propalaehoplophorinae. However, stemma and the Propalaehoplophorinae, and their as- in the Glyptodontinae, as in the Boreostemma species, signation to our new genus Boreostemma. Furthermore, the sulci are not parabolic in transversal section, but this set of evidences strongly suggests that these taxa have parallel sides and a flat bottom ("U"-shaped); this represent the first stages in the cladogenesis of the glypt- detail has already been noted by authors such as CAS- odontine Glyptodontidae. TELLANOS (1953) for Paraglyptodon. This is coupled to In agreement with this, the biogeographic and phy- the fact that the dorsal surface of the osteoderms is logenetic history of the Glyptodontinae was poorly strongly punctate in the earliest Glyptodontinae (CA- known until now, and it was practically restricted to the BRERA 1944; CASTELLANOS 1953), whereas, in the ter- southernmost regions of South America. Within this minal taxa (Glyptodon OWEN 1839 and Glyptotherium area, their Tertiary records are scarce, relatively late, OSBORN 1902), this surface is rugose, with deep and and show characters typical of the terminal Pleistocene wide sulci (BURMEISTER 1870-1874; AMEGHINO 1889; glyptodontines, which suggests that these are already SOmELZON et al. 2006; CARLINI et al. 2008). derived taxa. To sum up, the following trends seem to occur in Thus, the first record of a Glyptodontinae corre- the glyptodontine Glyptodontidae: a) progressive in- sponds to Glyptodontidium tuberifer from the "Araucan- crease of the width and depth of the sulci dividing the ian" (late Miocene - early Pliocene) of NW Argentina central figure from the peripherals; these always with (CABRERA 1944). The stratigraphic provenance of this parallel sides and a flat bottom; b) increasing rugosity taxon is somewhat uncertain, given that the "Araucan- of the dorsal surface of osteoderms, which is openly ru- ian" is represented by the Andalhuala and Corral Que- New Glyptodont from the Codore Formation and the paleobiogeography of the Glyptodontinae 149 mado formations. However, some datings have sug- Suggestively, the first Glyptodontinae records gested that the Andalhuala Fm. could correspond to 7 - (Glyptodontidium tuberifer) in southern South America 3.54 Ma (BUTTLER et al. 1984; BossI et al. 1987; BOSSI coincide with what PASCUAL & BONDESIO (1982) called & GAVRILOFF 1998; GAVRILOFF2000; HERRERA & OR- "Age of Southern Plains", which includes the Chasicoan, TIZ 2005). Likewise, the Corral Quemado Fm. is close Huayquerian, Montehermosan, and Chapadmalalan to 3.54 Ma in age (BUTTLER et al. 1984); however, dat- ages (ca. 11 - 3 Ma) (see PASCUAL et al. 1996) and whose ings from the base of this formation range between 4.61 beginnings were undoubtedly related to the Quechua and 4.83 Ma (STRECKERet al. 1989; BOSSI & GAVRILOFF Phase of Andean Orogeny (see YRIGOYEN 1979). More 1998), while the most modern ones correspond to about precisely, the evidence suggests that the first southern 2.9 Ma (BossI & GAVRILOFF 1998) or are even younger Glyptodontinae records correspond to the latest Mio- (2.4 Ma?; [sic] BOSSI et al. 1987). cene or Pliocene, when these great plains reached their The other Tertiary Glyptodontinae genus is Pa- maximum development (ORTIZ JAUREGUIZAR 1998), raglyptodon CASTELLANOS (Pliocene, Chapadmalalan extending from north Patagonia to Venezuela (MAR- - Marplatan) (see ROVERETO 1914; CASTELLANOS 1953) SHALL et al. 1983; ORTIZ JAUREGUIZAR • CLADERA from South America. One species of this genus (P. 2006). In fact, some authors have suggested that these chapalmalensis AMEGHINO, 1908) is biostratigraphi- climatic-environmental conditions greatly favored the cally important, since it is characteristic of the upper entrance of the first Holarctic immigrants, which al- Chapadmalan Age/Stage (ca. 3.5 - 3.2 Ma.) (see CLONE ready show clear adaptations to open grassland environ- & TONNI 2005). Similarly, the distribution of P. uquien- ments (e.g., Cricetidae and Tayassuidae) (VUCETICH sis seems to be restricted to the latest Chapadmalalan 1986; MENEGAZ & ORTIZ JAUREGUIZAR 1995; PAR- and the Marplatan (REGUERO et al. 2007). DISAS 1995). Thus, it is possible that these great open A remarkable species diversification occurred extensions also favored the dispersion of the glyptodon- among the Glyptodontinae during the Pleistocene (ca. tines into more southern areas through "biogeographi- 1.8 - 0.008 Ma), along with a wide geographic distri- cal corridors". bution that was probably stimulated by the expansion of The recognition of this new genus and species in open environments during the glacial periods (CLAP- northern South America, and the associated systematic PERTON 1993; CIONE et al. 2003; RABASSA et al. 2005). and paleobiogeographical consequences, highlight the im- In fact, these are probably the only Glyptodontidae to portance of research in continental sediments at these lat- successfully partake in the GABI, given that they occur itudes, especially considering the proximity to areas that in North America since the early Blancan (ca. 3.9 Ma) functioned as biogeographic corridors between both (CARRANZA-CASTAIqEDA & MILLER 2004), and evolved American subcontinents (WOODBURNEet al. 2006). These several taxa in that subcontinent, from the late Blancan corridors modified the biotic relationships in both Ameri- (ca. 2.6 Ma) to the Rancholabrean (late Pleistocene) cas, especially in South America, which had almost insu- (GILLETTE d~ RAY 1981; WHITE d~ MORGAN 2005; lar characteristics during a large part of the Cenozoic. MORGAN & WHITE 2005; CARLINI et al. 2008) (but see DOWNING & WHITE 1995). Conclusions To sum up, the evidence indicates that the Mio- Pliocene taxa from Colombia and Venezuela are not 1. The morphological evidence indicates that the Propalaehoplophorinae, but share several derived char- propalaehoplophorine Glyptodontidae as previously acters with the glyptodontine Glyptodontidae. Hence it defined (southern taxa plus taxa from northern South is possible to propose a partial reinterpretation of the America) are not a natural group. phylogenetic and biogeographic history of the Glypto- 2. Consequently, we propose the recognition of a new dontinae. Indeed, until the recognition of Boreostemma genus (Boreostemma) for the Venezuelan and Colom- n. gen. in the present contribution, the oldest records of bian species previously assigned to Asterostemma, Glyptodontinae (late Miocene - Pliocene) were re- while the latter taxon is geographically restricted to stricted to southern South America (Buenos Aires, Cat- Argentina. amarca and Jujuy provinces, Argentina) and suggested a 3. Boreostemma includes the following species: B. southern differentiation for the subfamily, probably acostae (VILLARROEL 1983) and B. gigantea (CAR- from advanced Propalaehoplophorinae (CARLINI et al. LIN! et al. 1997) from the middle Miocene of La 1997). Venta, Colombia, and B. venezolensis (SIMPSON On the contrary, the new evidence presented here 1947) and B. pliocena n. sp. from the Miocene and strongly suggests that the origin of the Glyptodontinae Pliocene of Venezuela, respectively. was geographically located in northern South America 4. The characters of this new genus and the species re- and much earlier (middle Miocene) than previously sup- ferred to it (e.g., sulci between central and peripheral posed, since the reliable records from the Honda Group, osteoderms "U"-shaped in transversal section, dorsal Colombia (Boreostemma gigantea and B. acostae), date surface of osteoderms in the dorsal carapace highly from approximately 13 Ma (CARLINI et al. 1997; KAY et punctate, greater osteoderm thickness, osteoderm ar- al. 1997). ticulation highly denticulated and with evident pro- 150 ALFREDO A. CARLINI et al.
longations, anterior molariforms complex), suggest gena Santa Marfa-Hualffn (Deptos. Santa Maria y Belfn, pro- the inclusion of these taxa among the glyptodontine vincia de Catamarca). - 12. Congreso Geol6gico Argentino y 2. Congreso de Explotaci6n de Hidrocarburos, Actas 2: 156-165. Glyptodontidae, which are rare in the fossil record of BURMEISTER, H. 1874. Monograffa de los Glyptodontes en el Museo southern South America for that period. Pdblico de Buenos Aires. - Anales del Museo Pfiblico de Bue- 5. Consequently, it is possible to hypothesize that this nos Aires 2: 367-377. genus represents the start of the cladogenesis of BUTTLER, R.F.; MARSHALL, L.G.; DRAKE, R.E. & GARNISS, H.C. glyptodontine Glyptodontidae, which seems to have 1984. Magnetic polarity stratigraphy and 4°K- 4°AR dating of taken place in much more northern regions than pre- Late Miocene and Early Pliocene continental deposits, Catamar- ca Province, NW Argentina. - Journal of Geology 92: 623- viously supposed. It is also possible that the arrival of 636. these Glyptodontinae to southern South America was CABRERA, A. 1944. Los Gliptodontoideos del Araucaniano de Cata- favored by the emergence of large open savanna en- marca. - Revista del Museo de La Plata (Nueva Serie) 3 (15): 5- vironments during the "Age of Southern Plains". 76. 6. The propalaehoplophorine Glyptodontidae are re- CARLINI, A.A. & SCILLATO-YANI~,G.J. 1999. Evolution of Quaternary stricted to the Oligo-Miocene of the southern cone of Xenarthrans (Mammalia) of Argentina. - In: RABASSA, J. & SA- LEMME, M., eds., Quaternary of South America and Antarctic South America, mainly in Patagonia. Peninsula 12: 149-175. CARLINI, A.A.; VIZCA~NO, S.F. & SCILLATO-'YANI~, G.J. 1997. Ar- mored Xenarthrans: A unique taxonomic and ecologic assem- Acknowledgments blage. - In: KAY, R.F.; MADDEN, R.H.; CIFELLI, R.L. & FLYNN, J.J., eds., Vertebrate paleontology in the Neotropics. The Mio- cene fauna of La Venta, Colombia: 213-226, Washington D.C. We wish to thank IPC (Instituto del Patrimonio Cultural de (Smithsonian Institution Press). Venezuela) for permission to study important glyptodont ma- CARRANZA-CASTAiqEDA, O. 8¢ MILLER, W.E. 2004. Late Tertiary ter- terials that are part of the national patrimony of the Repfiblica restrial mammals from Central Mexico and their relationship to Bolivariana de Venezuela; and C. Morgan for the English South American immigrants. - Revista Brasileira de Paleontolo- translation of the manuscript. AAC and AEZ especially thank gia 7 (2): 249-261. M.R. S~inchez-Villagra and O. Aguilera for the invitation to CASTELLANOS, A. 1932. Nuevos g6neros de gliptodontes en relaci6n contribute to this volume. This research was funded by grant con su filogenia. - Physis 11 (38): 92-100. PICT-R 074 (G3) and UNLP-FCNYM N-514 to AAC and by CASTELLANOS, A. 1953. Descripci6n de restos de Paraglyptodon the University of Zfirich. uquiensis n. sp. de Uqufa (Senador P6rez), Jujuy. - Memorias del Museo de Entre Rfos 32 (Paleontologia): 1-32. CLONE, A.L. & TONNI, E.P. 2005. Bioestratigrafia basada en mamfferos del Cenozoico Superior de la provincia de Buenos Aires, Argen- References tina. - 16. Congreso Geol6gico Argentino, Relatorio: 183-200. CLONE, m.; TONNI, E.P. 8¢ SOIBELZON, L. 2003. The broken Zig-Zag: AMEGHINO, F. 1887. Enumeraci6n sistemfitica de las especies de Late Cenozoic large mammal and tortoise extinction in South mamlferos f6siles coleccionados por Carlos Ameghino en los America. - Revista del Museo Argentino de Ciencias Naturales terrenos Eocenos de Patagonia austral y depositados en el Mu- 5 (1): 1-19. seo de La Plata. - Boletfn del Museo de La Plata 1:1-26 CLAPPERTON, C. 1993. Nature of environmental changes in South AMEGHINO, F. 1889. 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