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Home , Cuvieronius, Elephant, Elephantoidea, Gomphothere, Gomphotherium, Mammutidae

José Luis Prado 1, María Teresa Alberdi 2, Begoña Sánchez 2 & Beatriz Azanza 2 1 Universidad Nacional del Centro UNC, Olavarría 2 Museo Nacional de Ciencias Naturales, Madrid

Diversity of the (Gomphotheriidae, ) from

Prado, J.L., Alberdi, M.T. , Sánchez, B. & Azanza, B., 2003 - Diversity of the Pleistocene Gomphotheres (Gomphotheriidae, Proboscidea) from South America - in: Reumer, J.W.F., De Vos, J. & Mol, D. (eds.) - ADVANCES IN RESEARCH (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 347-363 [ISSN 0923-9308] Published 24 May 2003

The gomphotheres were recorded in South America from the early Middle Pleistocene (Ensenadan Land- Age) to the latest Pleistocene ( Land-mammal Age). They were descen- dants of the stock that originated in and arrived to South America during the ‘Great American Biotic Interchange’. Only two genera are recognised: with only one (Cuvieronius hyodon), and with two species (Stegomastodon waringi and Stegomastodon platensis). Two corridors would have developed during the Pleistocene in South America. These two corridors have conditioned the paleobiogeographic his- tory of most North American in South America. In fact, different models can be postu- lated for different groups depending on their capacity to produce distinct adaptive types throug- hout the duration of their dispersion process. In the case of South American gomphotheres, the small Cuvieronius utilised the Andean corridor, whereas the larger Stegomastodon dispersed through the eastern route. Cuvieronius hyodon is geographically restricted to the Andean Region in , , , and Northwestern Argentina, it inhabited an arid landscape. Cuvieronius from Tarija indicated that they were almost exclusively mixed-feeders. This species seems to have been adapted to a temperate-cold climate, since in the inter-tropical zones it has been only found at the highest levels, while in Chile it expanded to the littoral zone, that surely offered similar living conditions, in terms of temperature as the corridor. Stegomastodon waringi was recorded in the Santa Elena peninsula in Ecuador, and in . S. waringi from the Peninsula of Santa Elena, shows an adaptive trend of mixed-feeder to grazer. Stegomastodon pla- tensis was recorded in the Middle to latest Pleistocene of Argentina, especially the Pampean Region, and also during in Uruguay and Paraguay. S. platensis from the Middle Pleistocene of Argentina shows mixed-feeder to browser feeder adaptations. Stegomastodon seems to have predominated in lower latitudes, where it would occupy savannahs or xerophytic pasture areas, and consequently it would be better adapted to warm or temperate climatic conditions. Their most austral distribution does not surpass the 37th parallel in the Buenos Aires province. The fre- quency of Stegomastodon platensis diminishes in the Pampean Region by the latest Pleistocene, when environmental conditions became colder and drier.

Correspondence: José Luis Prado, INCUAPA - Departamento de Arqueología, Universidad Nacional del Centro UNC, Del Valle 5737, 7400 Olavarría (Argentina); María Teresa Alberdi *, Begoña Sánchez and Beatriz Azanza, Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal, 2, 28006 Madrid (); e-mail: malber- [email protected]; Azanza also at Area de Paleontología, Departamento de Ciencias de la Tierra, Universidad de Zaragoza, 50009 Zaragoza (Spain). [* to whom correspondence should be addres- sed]

Key words: , stable isotopes, gomphotheres, Cuvieronius, Stegomaston, early Middle and Late Pleistocene, South America.

347 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

INTRODUCTION thesised about the paleoecology and niche The family Gomphotheriidae is considered a differentiation of the three species of South long-living ancestral stock that gave origin to American gomphotheres: Cuvieronius a succession of other groups. Members of this hyodon, Stegomastodon platensis and group spread into , and North Stegomastodon waringi. America from Late to Late Pleisto- cene and arrived to South America in early MATERIAL AND METHODS Middle Pleistocene times. Casamiquela et al. We have studied the characteristics of the (1996) consider that gomphotheres arrived in , jaw, and appendicular skeleton follow- South America during the or Plio- ing Alberdi (1971), Mazo (1977), Tassy Pleistocene boundary but there are no data to (1983, 1988), Boeuf (1992) and Shoshani confirm this early arrival. The gomphotheres (1996a). From them we have selected the sensu lato existed in North America from the more important characters and parameters to Middle to the end of the Pleistocene compare the specimens from the more (Late Barstovian to the end of the Rancho- important localities of South America. Most labrean). Gomphothere diversity reached its of the material comes from the middle and peak during this span as well, with six genera late Pleistocene localities in Argentina, known from the Late Clarendonian to the Bolivia, Ecuador and Chile. We have revie- Early : , wed some remains from Museo Nacional de (in ), Historia Natural, La Paz (MNHN), Museo de , Serbelodon, , and La Plata (MLP), Museo Argentino de Torynobelodon. Gomphotheres were greatly Ciencias Naturales ‘Bernardino Rivadavia’, reduced in numbers during the late Buenos Aires (MACN), Museo de la Escuela Hemphillian with only three genera: Politécnica Nacional de Quito (MEPN), Gomphotherium, Rhynchotherium, and Museo Nacional de Historia Natural, Amebelodon. Webb (1977, 1983) based this Santiago de Chile (MNHN), and we compa- decline of the North American fauna red with the bibliographic data from Cabrera during the Miocene on the continent-wide (1929), Hoffstetter (1952), Simpson & Paula replacement of by more open forested Couto (1957), Alberdi & Prado (1995), savannahs that began in the Hemingfordian, Casamiquela (1972, 1976), and Casamiquela and the subsequent transformation of this et al. (1996). savannah into open grassland that began in The gomphotheres of South America come the Late Clarendonian/Early Hemphillian. from the Andean corridor (from Colombia at Since savannahs can support a greater the North and Chile at the South), Tarija diversity of than either closed (Bolivia), and Pampean Region (Argentina), or open grassland, this situation had and several localities in Brazil (Fig. 1). The the effect of enlarging available localities placed on the Andean corridor are niche space. During times, Rhyncho- characterised in many sites by rugged plateau therium was more abundant, and Stego- terrain at 4000 m altitude. Most of the fossil and Cuvieronius were recorded. remains were recovered from eolian deposits Stegomastodon became extinct near the assigned to the Cangahua Formation in Middle in North America and Ecuador. Sauer (1965) presented the most Cuvieronius at the end of the complete geological overview of this forma- (Kurtén & Anderson 1980). tion. Clapperton & Vera (1986) referred this The purpose of this paper is to present the formation to the last cold stage of the taxonomic scenario of the Pleistocene gom- Pleistocene. The most significant fossil photheres from South America. Based on assemblages from the coastal area are those and isotopic evidences we hypo- of La Carolina (Santa Elena Peninsula).

348 PRADO et al.: South American Pleistocene Gompotheres

Figure 1 Possible geographic distribution routes of South American gomphotheres.The small Cuvieronius utilised the Andes cor- ridor (arrow, clearer and narrower), and the large Stegomastodon dispersed through the Eastern route (arrow, darker and larger). Modified from Alberdi & Prado (1995).

349 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Hoffstetter (1952) suggests that all these fos- TAXONOMIC FRAMEWORK sils come from a single geological horizon Cuvier (1806) studied for the first time the referred to the Late Pleistocene. Most records remains found by Dombey in Perú and from Bolivia are from Tarija. This site is Humboldt in Colombia, Ecuador (Imbabura), located at an elevation of 1950 m at latitude Bolivia (Chiquitos) and Chile (La 21°33'S and longitude 64°46'W in southern Concepción). Cuvier grouped these masto- Bolivia. Oppenheim (1943) presented the dons in relation to their provenance in most complete geological overview of the "mastodonte des cordillères" and "mastodonte Tarija Formation, which is well known for its humboldien", respectively. In 1920, Boulé & classic vertebrate faunas. This formation Thevenin described the specimens collected spans a time interval from about 1 to 0.7 in Bolivia as Mastodon andium (Table 1). MyBP or perhaps younger (MacFadden et al. Later, taxonomic controversy arose about in 1983). Tarija fauna is classically assigned to which families these gomphotheres must be the Ensenadan SALMA (South American placed (Table 2). Land Mammal Age). Geographically, Tarija is Cabrera (1929) included three families in considered part of the sub-Andean physio- the : Gomphotheriidae, graphic zone. The deposits from the Pampean and , where buno- Region are considered by the palaeomagnetic dont, zygodont and stegodont and stratigraphy and were deposited during the were included respectively. All pro- last 2.5 My. The stratigraphy of the Pampean boscideans from South America are buno- typically consists of superposed 1-2 m donts and brevirostrines and are placed in the thick beds separated by either erosional dis- Gomphotheriidae family. He created the sub- continuities or palaeosols. Most of the family Cuvieroniinae differentiated from the Stegomastodon remains came from the locali- Anancinae only by the presence of an enamel ties assigned to Ensenadan and Lujanian band on the upper . Hoffstetter (1952), SALMA. The most important Chilean locali- as Cabrera, included all South American pro- ties come from central-southern and central boscideans in two subfamilies: the Chile. is located in the southern Cuvieroniinae that originated in North limit of continental Chile in the central America and spread to South America and Depression or Central Valley, at around 600 m that included remains from Ecuador altitude, between the Andean Cordillera and (Imbabura at N), Colombia, Bolivia and the Coastal Mountain Range. Monte Verde Chile; and the Anancinae that arose in are located in a depression filled by and spread into Europe and part of Asia, fluviatile, glacial, and moraine including all other genera from Ecuador, and covered by volcanic ash. The Tagua-Tagua Brazil and Argentina. Simpson & Paula locality is placed in the Laguna of Tagua- Couto (1957) indicated that these subfamilies Tagua (latitude 34º30'S), in the Central Valley in discussion, Anancinae and Cuvieroniinae, of Chile, approximately 120 km south of have a pattern similar of that of modern ele- Santiago City. The basin lies at 200 m altitu- phants in the loss of their lower tusks, elong- de on the eastern margin of the coastal cordil- ation of their molars, and deepening of their lera, and it appears to have resulted from a . Consequently, they included all South large lahar or volcanic debris avalanche event American gomphotheres in one subfamily: in the late . The majority of the Anancinae. This subfamily has priority follo- Brazilian remains have no stratigraphic data, wing the rules of nomenclature (Simpson but it can be considered that many or most of 1945; and ICZN 1985). Alberdi & Prado the Brazilian finds are from rather late in the (1995) agree with Simpson & Paula Couto Pleistocene. (1957) and included the South American gomphotheres in one subfamily: the

350 PRADO et al.: South American Pleistocene Gompotheres

Table 1 Taxonomic history of the South America gomphotheres before 1920.

Anancinae, given that the differences among distic analysis without explanation about their them are few and slight. Recently, Shoshani relationship with subfamily Anancinae. The & Tassy (1996: Appendix B), suggested a only explanation is in footnote 19, page 172, classification of the Proboscidea based on where he writes: "19. This possible ‘shuf- combined results by Tassy (in the same book) fling’ of genera within Cuvieroniinae was and Shoshani, where they included the South predicted by Simpson & Paula-Couto American gomphotheres in the Gompho- (1957)". But these authors consider that all theriidae HAY, 1922 and the Cuvieroniinae South American gomphotheres must be inclu- CABRERA, 1929, and not in the Anancinae ded in only one subfamily and "(…) the ear- subfamily. The result is the following: Tassy liest name formed according to the Rules and (1996: Cp. 6) in his cladogram indicates two available for a subfamily uniting these genera gomphothere groups: the first for the Old is Anancinae" (Simpson & Paula-Couto 1957: World trilophodont gomphotheres, and the 181). second for the New World trilophodont We found no significant differences gomphotheres that included all South between Anancinae and Cuvieroniinae. The American forms. He considers the New subfamily Anancinae is characterised by a World gomphotheres to be more closely rela- brachycephalic skull with tendency to ele- ted to tetralophodont gomphotheres than to phantoid skull, and a brevirostrine jaw with trilophodont gomphotheres from due curved symphysis. The upper tusks varied to the presence of lower tusks with a rounder from more or less elongated and straight to cross-section. He also suggested the possible curved in the distal part. There is no enamel relationship between of band in the adult specimens. Bunodont and brevirostrines from South America as molars, with angular disposition of the pretri- indicated by Tobien et al. (1986, 1988). In te and posttrite cusps, are brachydonts or sub- the same book, Shoshani (1996b: Cp. 16) hypsodonts, with single or slight complicate included all South American gomphotheres in trefoils in the occlusal surface. M2 are trilo- the subfamily Cuvieroniinae through the cla- phodont with a developed talon sometimes.

351 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Table 1 Taxonomic history of the South America gomphothe- M3 vary from tetralophodont to heptalopho- res after 1920. dont. We observed that these features are the generalised pattern for all South American gomphotheres. The subfamily Cuvieroniinae is only differentiated by the enamel band in adult upper tusks. We will try to establish an overview where we evaluate if the differences among the forms are simply individual varia- tions or represent some differences that can occur at or species level or if they reach the subfamily level.

Family Gomphotheriidae CABRERA, 1929 Genus Cuvieronius OSBORN, 1923

Synonymy: Mastodon (part) Auct., nec CUVIER, 1817: 233 (= Cuvieronius OSBORN, 1923: 1; = Cordillerion OSBORN, 1926: 15; = Teleobunomastodon REVILLIOD, 1931: 21). Haplomastodon (part) HOFFSTETTER, 1950; = Haplomastodon (Haplomastodon) HOFFSTETTER, 1952: 192. Cuvieronius from Tagua-Tagua (Casamiquela et al. 1967, 1996; Casamiquela 1972; Montané 1967, 1968). Cuvieronius from Monte Verde (Dillehay 1989, 1997; Casamiquela et al. 1996).

Type species: Cuvieronius hyodon (FISCHER, 1814: 341).

Stratigraphic distribution: The genus is known from late Pliocene-late Pleistocene localities in the (Tedford et al. 1987). The most ancient in South America corresponds to the lower-middle Pleistocene (Ensenadan SALMA) and the most recent to the late Pleistocene (Lujanian SALMA), Lujanian - boundary.

Geographic distribution: Cuvieronius occurs in as far north as NW Oaxaca (Ferrusquía-Villafranca 1978), and probably it is present during the Pliocene in Valse- quillo (Miller & Carranza-Castañeda 1984). The genus is also known in Central America from Guatemala but with doubts (Woodburne 1969; Lucas et al. 1997). In Nicaragua it is cited again, with doubts by Espinoza (1976),

352 PRADO et al.: South American Pleistocene Gompotheres

Reshetov (1982), and Lucas et al. (1997). In RAFINESQUE, 1814: 182. Mastodon Andium it is noted by Webb & Perrigo CUVIER, 1824: 527. Mastodon argentinus (1984) and it is very common during the AMEGHINO, 1888: 7. Mastodon chilensis Pleistocene in Costa Rica (Lucas et al. 1997) PHILIPPI, 1893: 88. Mastodon bolivianus on the Cordilleras. In El Salvador it is obser- PHILIPPI, 1893: 89. Mastodon tarijensis ved during the Late Pleistocene, and also in AMEGHINO, 1902: 2. Cuvieronius hyodon Panama (Gazin 1957). The most important (FISCHER, 1814), in Cabrera (1929: 82-90). record for this genus in South America comes Cuvieronius hyodon (FISCHER, 1814), in from the Andean Cordillera. In Colombia Hoffstetter, 1952: 186. Species A from Rio specimens are cited in Tibitó and Mosquera, Chiche, in Hoffstetter, 1952: 223-224. close to Bogotá at 3.800m altitude Cuvieronius tarijensis FICCARELLI et al. 1995: (Hoffstetter 1952; Simpson & Paula-Couto 754. 1957; Correal Urrego 1981). In Ecuador spe- cimens are recorded at Imbabura volcano, Type: M2 figured by Cuvier (1806: pl. II, Rio Chiche and Andean localities (Hoffstetter fig.1), from Imbabura (Northern Ecuador). 1952). There are also remains in Ulloma and Tarija in Bolivia (Boule & Thevenin 1920; Stratigraphic distribution: The more Hoffstetter 1952; Simpson & Paula-Couto ancient remains known in South America 1957; Alberdi & Prado, 1995). In Chile come from the early Middle Pleistocene in remains are cited in Tagua-Tagua, La Ligua the Andean region from Imbabura. The most and Chillán in the Central Chile and Monte recent come from the Late Pleistocene from Verde, Southern Chile (Casamiquela 1972; Tibitó in Colombia, and Tagua-Tagua and Dillehay 1984, 1989, 1997; Alberdi & Prado Monte Verde in Chile (Fig. 1). 1995). See Figure 1. Geographic distribution: Cuvieronius Diagnosis: The skull is comparatively low hyodon entered in South America extending and elongate. They are brevirostrine, buno- from Colombia in the North to Monte Verde, lophodont, trilophodont mastodons with single Chile in the South. The most important data trefoils or with posttrite cusps poorly develo- come from Colombia (Tibitó and Mosquera ped (M2 trilophodont). Pretrite and posttrite localities), Ecuador (Imbabura volcano, Rio cusps opposite of slightly alternating, on pos- Chiche), Bolivia (Ulloma and Tarija), and terior lophs of M3 (with 4 1/2 to 5 lophs Perú (Cabrera 1929, Boule & Thevenin 1920, having a certain angular inclination). Tusks Hoffstetter 1952, Simpson & Paula-Couto twisted in a long, cross-section sub-circular, 1957, Correal Urrego 1981, Alberdi & Prado open spiral and with a spiral band of enamel 1995, Ficcarelli et al. 1995). The record in persisting in the adults, and tusks alveolus Chile comes from Tagua-Tagua, La Ligua and more or less divergent. Chillán in Central Chile and Monte Verde in Southern Chile (Casamiquela 1972, 1976; Discussion: It is important to note that all Casamiquela & Dillehay 1989; Dillehay authors agree in recognise the presence of the 1989, 1997; Casamiquela et al. 1996). genus Cuvieronius in South America with Burmeister (1867) and Siroli (1954) cited only one species: C. hyodon. some localities in NW Argentina, but we found no reliable data about them. Cuvieronius hyodon (FISCHER, 1814) Diagnosis: The same as for the genus. Synonymy: Mastotherium hyodon FISCHER, 1814: 341. Mastotherium Humboldtii Discussion: We accept the assignment to this FISCHER, 1814: 341. Mastodon rhomboides species of material from most of the localities

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cited above, as Ulloma and Tarija in Bolivia, Venezuela. Imbabura in the North of Ecuador and the Andean Ecuadorian localities, Tagua-Tagua Geographic distribution: The most signifi- and Monte Verde in Chile. Ficcarelli et al. cant record of Stegomastodon in South (1995: 747) identified the gomphothere of America comes from the Taima-Taima site in Tarija as Cuvieronius tarijensis nov. sp. We Venezuela (Bryan et al. 1978, Bryan 1986, do not understand why they revalidated this Casamiquela et al. 1996), and Santa Elena specific name created by Ameghino (1902) peninsula in Ecuador (Hoffstetter 1952). for the Tarija site, without any explanation Also, it was widely dispersed through the about it. Hoffstetter (1952) considered it as Eastern route into the tropical region of South nomem nudum which opinion met general America (Proaño 1922, Hoffstetter 1952). It consensus among the specialists. arrived as far as Argentina, the most austral The genus Haplomastodon has not been well distribution, especially the Pampean Region described. Their separation from Cuvieronius and in Uruguay, and possibly in Paraguay and Stegomastodon is not clear and someti- (Cabrera 1929; Simpson & Paula Couto mes arduous. The only characteristic feature 1957; Mones & Francis 1973). Simpson & is the presence of a transverse foramen in the Paula Couto (1955, 1957) indicated their pre- atlas (Hoffstetter 1950, 1952). Simpson & sence in Brazil as well as in tropical and Paula Couto (1957) indicated that this charac- warm zones (Figure 1). ter is variable. The differential characters between these genera are not very clear Diagnosis: Gomphotheres with short and because of their great variability related to high elephantoid skull, less depressed than age or to the allometric state. Cuvieronius. Brevirostrine, jaw with bunolo- phodont and trilophodont teeth (M2). Less Genus Stegomastodon POHLIG, 1912 bunodont or choerodont than the most primi- tive Old World Anancinae but more than Synonyms: CABRERA, 1929: Cuvieronius. Moderate alternance of posterior 90-96, figs. 2 and 4; = Stegomastodon lophs of M3 (have a certain angular inclina- (Haplomastodon) HOFFSTETTER, 1950: 22, tion, pretrite and posttrite cusps opposite, and figs. 2 and 3; = Haplomastodon lophs 5 - 5 1/2 principally). With double tre- (Aleamastodon) HOFFSTETTER, 1952: 208). foils but relatively simple pattern. Tusks Haplomastodon sensu Ficcarelli et al. 1993: simply curved to nearly straight, without 233, 1995: 753. Species actually placed in enamel, only in some juvenile individuals. this genus have also been referred to Mastodon, Gomphotherium, , Stegomastodon waringi (HOLLAND, 1920) Cuvieronius and probably to other genera (see Cabrera 1929, Osborn 1936). Synonymy: Mastodon brasiliensis LUND, 1842; in Lesson (1842: 157), is considered a Type species: Stegomastodon mirificus nomen nudum. Mastodon andium WINGE, LEIDY, 1858: 10. 1906: 48, nec CUVIER, 1806: 413. Mastodon waringi HOLLAND, 1920: 229. Mastodon Stratigraphic distribution: The genus was chimborazi PROAÑO, 1922. Bunolophodon recorded in the Late Pliocene and the Early Ayora SPILLMANN, 1928: 70. Bunolophodon Pleistocene in central and western regions of postremus SPILLMANN, 1931: 73. North America. In South America it is present Notiomastodon vidali CASTELLANOS, 1948: in the Middle - Late Pleistocene of Brazil, 139. Haplomastodon (Aleamastodon) guaya- Argentina, Paraguay and Uruguay, and the sensis HOFFSTETTER, 1952: 208. Species B late Pleistocene of Ecuador, Colombia and from La Carolina, in Hoffstetter, 1952: 224.

354 PRADO et al.: South American Pleistocene Gompotheres

Haplomastodon (Haplomastodon) chimborazi consider that the skull and other remains (PROAÑO, 1922), in Hoffstetter 1952: 192. from Barranca Pistud, near Bolivar province Stegomastodon brasiliensis HOFFSTETTER of Carchi, Northern Ecuador, described by 1952: 222. Haplomastodon waringi Ficcarelli et al. (1995:753) as Haplomastodon (HOLLAND, 1920), in Simpson & Paula Couto chimborazi, must be included in Stego- 1957: 171. Haplomastodon chimborazi mastodon waringi. The differences noted by (PROAÑO, 1922), in Ficcarelli et al. 1995: these authors (p. 754), are in our opinion not 753. Haplomastodon waringi (HOLLAND, consistent with this taxonomic separation 1920), in Casamiquela et al. 1996: 316. because they correspond to an allometric variability. In general we consider that there Type: CM-11033 described by Holland are few differences among the gomphotheres (1920: 229) as a partial lower jaw and other species from South America. We think that fragments from Pedra Vermelha, Bahia, the smallest differences between Stego- Brazil. Simpson & Paula Couto (1957:172) mastodon species are more related to dif- notified that in the Carnegie Museum collec- ferences in the environmental conditions tions there are only three fragments, than to genetic conditions. the tip of a , a fragment of tusk dentine, and part of the distal end of a tibia, under Stegomastodon platensis (AMEGHINO, number CM-11033. 1888)

Stratigraphic distribution: Stegomastodon Synonymy: Humboldtii BLAINVILLE, waringi is characteristic from the Middle to 1845: 249, nec Cuvier, 1824: 527. Mastodon Late Pleistocene (Ensenadan and Lujanian platensis AMEGHINO, 1888: 7. Mastodon SALMAs). superbus AMEGHINO, 1888: 7; also in Casamiquela et al. 1996: 316. Mastodon Geographic distribution: Stegomastodon bonaerensis MORENO, 1888: 17-18 (nomen waringi is known from Lagoa Santa (southern nudum). Mastodon rectus AMEGHINO, 1889: Brazil) and Santa Elena Peninsula in Ecuador. 643. Mastodon maderianus AMEGHINO, 1891: In Brazil they were found on the so-called 243. Notiomastodon ornatus CABRERA, 1929: ‘East route’ or ‘ corridor’ (Webb 1978, 91; also in Casamiquela et al. 1996: 316. See 1985). Possibly, it also exists in the Taima- Cabrera (1929) and Simpson & Paula Couto Taima locality in Venezuela (Fig. 1). (1957) to complete the synonymies.

Diagnosis: The skull is somewhat less Type: MLP-8-63, an isolated tusk, from the elephantoid than in the type species but Pampean Region. Signalled by Ameghino somewhat less depressed than in Cuvieronius. (1888: 7), and figured in Ameghino (1889). The occlusal surfaces have a relatively simple trefoil pattern. The pretrite and posttrite cups Stratigraphic distribution: Stegomastodon with stronger mean tendency towards alterna- platensis is characteristically from the middle tion that in Cuvieronius. Tusks simply curved Late Pleistocene (Ensenadan and Lujanian to nearly straight or very slightly spiral, with SALMAs) from the Pampean Region. enamel band in some juvenile individuals; and usually simply upcurved and without Geographic distribution: Stegomastodon enamel in adults. platensis was the more austral gomphothere, Discussion: We understand that the species typical of the Pampean Region, specially waringi is closer to Stegomastodon than to from Buenos Aires, Córdoba, Santa Fé and Haplomastodon because, as we explained Entre Ríos provinces (Argentina). It was also above, Haplomastodon is poorly defined. We found in Uruguay (Mones & Francis 1973)

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and possibly in Paraguay (Cabrera 1929; conditions. Sánchez et al. (in press) analysed Simpson & Paula Couto 1957). See Fig. 1. the diet from different species of gomphotheres from Pleistocene deposits of South America Diagnosis: The skull is similar to the type through isotopic analysis and reconstructed the species, short and high, elephantoid. The diets of Cuvieronius hyodon, Stegomastodon teeth are tetralophodont or pentalophodont platensis, and Stegomastodon waringi. The upper and lower M3; and the occlusal pattern carbon isotopic results for Cuvieronius and with secondary conules or conelets that give Stegomastodon from the Middle Pleistocene it a morphology that varies from simple to indicate significant ecological patterns. On the complex. The tusks are long and relatively one hand, Cuvieronius from Tarija have more thin, without enamel band, curved in males homogeneous values, in agreement with those and straight in females. previously analysed for MacFadden & Sockey (1997), and indicating that they were almost Discussion: Ameghino (1888) created four exclusively mixed-feeders. On the other hand, species from the Pampean remains. Cabrera Stegomastodon from Argentina have more (1929) argued that three of these are syno- negative values showing a trend from mixed to nyms and retained Stegomastodon superbus. browser feeders. The statistical test confirms However, following the International Code of significant differences between the two gom- Zoological Nomenclature (ICZN 1985), the phothere groups from the Middle Pleistocene. priority lies with S. platensis. The S. platensis Stegomastodon waringi from the Peninsula of in both papers are referred firstly, and have a Santa Elena, Ecuador shows an adaptational wide geographical and stratigraphical distri- trend of mixed-feeder to grazer, clearly defined bution. Stegomastodon platensis (AMEGHINO, by the altitudinal and latitudinal distribution. 1888) show the characters of Stegomastodon Stegomastodon platensis from the Late in skull and jaw. We have observed that Pleistocene of Argentina seem to be the contra- thickness and curvature of tusks is a very ry, indicating a trend from mixed to browser variable character and related to sexual feeding (Sánchez et al. in press). The ecologi- dimorphism, not only in the South American cal pattern of the latter is in agreement with forms but also in other parts of the world the results obtained from the modern , (Tobien 1973; Mazo 1977; Boeuf 1992). We Loxodonta africana, from Amboseli Park also include in this genus and species the (Bocherens et al. 1996). From an ecological specimen from the late Pleistocene of Playa point of view, we can conclude that there is a del Barco (a tusk number 5451-MACN, feeder evolution since the Middle to the Late Montehermoso, Buenos Aires) referred by Pleistocene Stegomastodon group from Cabrera (1929) to Notiomastodon ornatus. He Buenos Aires that implies a change of the created this genus and species based on a dif- diet from C3 to mixed , and from this ferent stratigraphical provenance (the prove- last to exclusively mixed-feeders. nance is unclear), and the absence of an enamel band. In this sense, the discussion in , GOMPHOTHERES AND Simpson & Paula Couto (1957: 177) is very THE GREAT AMERICAN BIOTIC interesting. INTERCHANGE (GABI) Around 2.5 My ago, tectonic activities along FEEDING AND HABITATS the Pacific margin caused the American con- Generally, it is considered that the Cuvie- tinents to be sutured. One habitat corridor ronius habitat could be high grassland with opened, facilitating the dispersion of terres- cold to temperate climatic conditions, and trial plants and into and out of South Stegomastodon would seem to be adapted to America, triggering an event known as the more open grassland with warm to temperate GABI (Webb 1976, 1991). This land bridge

356 PRADO et al.: South American Pleistocene Gompotheres

functioned as an ecologically selective dis- elephants and extinct Cuvieronius that was a persal corridor (Webb 1978; Simpson 1980). mixed-feeder. These different preferences Biogeographic data indicate an alternation among the mastodons, mammoths and gom- among three major types of Plio-Pleistocene photheres could explain why only the bunod- habitat corridors on the Panamanian land ont forms arrived to South America when all bridge: mesic tropical forest, mesic , types lived together in North America during and xeric scrub savanna (Webb 1978). During the Pleistocene. the humid phase, the tropics were dominated by rain forests, and the principal GOMPHOTHERE IN biotic movement was from Amazonia to SOUTH AMERICA Central America and southern Mexico. Causes for the Pleistocene extinction are During the more arid glacial phase, savannah many, making it difficult to consider and eva- habitats extended broadly right through tropi- luate all of the complex phenomena that pro- cal latitudes (Webb 1991). Simpson & Paula duced the disappearance of an important part Couto (1957) proposed that all known gom- of the Pleistocene fauna. Thus, to test the photheres from South America derived from theories, analysis of certain factors are favou- an independent radiation in Central America. red over others. Two types of theories have The early Cuvieronius and Stegomastodon been offered for this extinction. Some authors apparently entered South America at the attribute the extinction to the direct impact of Early or Middle Pleistocene during the gla- man through activities. This hypothe- cial phase. sis has been mainly held by Martin (1984) No common biological feature appears to who proposed that the extinction of large explain why Mammuthus (Elephantidae) and mammals from North America, South Mammut (Mammutidae), that were recorded America and are related to sudden in and Honduras during the and several impacts on these conti- Pleistocene and which might be expected to nents. It seems that human activities, through have crossed the Panamanian land bridge, did hunting pressure or disturbing the habitat, not reach South America. The reasons for this affected the Pleistocene population of gom- were perhaps individualistic. As cited before, photheres (Politis et al. 1995). Human activi- the Late Pleistocene mammoth and mastodon ties that would have disturbed the environ- had distinct diets and perhaps different habi- ment in such a way as to greatly affect gom- tat preference. Mastodons had relatively low- photheres include human-made fires on gras- crowned molars with cusps arrayed in widely slands. The overkill hypothesis is supported spaced lophs and high relief on the occlusal by the synchrony of extinction with the arri- surface prior to heavy wear. This dental val of large numbers of in these con- morphology led to recognise mastodons as tinents. In contrast, the data from South browsers (Webb et al. 1992). Mammoths had America do not support the overkill hypothe- high-crowned molars with closely spaced sis because several data of coexistence with enamel lophs coated with cement. This mor- hunter-gatherer bands were recorded. The phology led to the classification of mam- archaeological record from South America moths as grazers (Davis et al. 1985). Isotopic shows that gomphotheres are frequent in analyses confirm this hypothesis (MacFadden Paleo-Indian Sites. Evidence that gomphothe- & Cerling 1996). The gomphotheres from res were a human food resource come from West Palm Beach, Florida and from South Monte Verde in the southern Chile (Dillehay America had a d13 C values intermediate & Collins 1988), Tagua-Tagua in Central between browsers and grazers. In sum, both Chile (Montané 1968), Tibitó in the Bogotá extinct mammoths and mastodon species Sabana of Colombia (Correal Urrego 1981), were more specialised feeders than modern and Taima-Taima in the Coro Peninsula of

357 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Venezuela (Bryan et al. 1978). These sites are and Mammuthus have higher and more varia- dated between c. 13,000 to 11,000 yBP. ble values, indicating a diet variably enriched Gomphotheres are also present in the in C4 grass. These data seem to support the Pampean Region during this time but no Graham & Lundelius model. association with humans has been recorded yet. CONCLUDING REMARKS On the other hand, climatic and ecological The South American gomphotheres are changes, particularly to nutritional stress included in the family Gomphotheriidae. induced by rapid change in communi- We recognise two genera: Cuvieronius and ties, are identified as the main cause of Stegomastodon, and three species: extinction (Graham & Lundelius 1984; King Cuvieronius hyodon, Stegomastodon waringi & Saunders 1984). These models imply that and S. platensis. The genus Cuvieronius gomphothere taxa were feeder specialists, arrived to South America during the Great adapted to a kind of plant that disappeared American Biotic Interchange, and it has been during the Holocene. The hypothesis propo- recorded from the to the sed by Guthrie (1984) was based on the Late Pleistocene. The genus Stegomastodon assertions that plant diversity was higher and appeared later during the Middle Pleistocene. that the growing season was longer in the Two corridors would have developed Pleistocene than in the Holocene. With during the Pleistocene in South America. changes in seasons and diversity during the These two corridors have shaped the paleo- Holocene, these herbivores could not obtain biogeographic history of most North the diverse mixture of plants needed for sur- American mammals in South America. The vival. This model implies a different type of model postulated most viable for gomphothe- specificity, namely that extinct res dispersion process seems to indicate that were obligate mixed-feeders. the small Cuvieronius utilised the Andes cor- African and Asian elephants live in a diverse ridor, whereas the large Stegomastodon dis- habitat. They are opportunists, capable of persed through the Eastern route. Cuvieronius living on nearly any dietary mixture and Stegomastodon are bunodont mastodons. (Bocherens et al. 1996, Koch et al. 1995). If In general, the bunodont forms lived in open gomphotheres showed a similar level of and drier areas, like woodland savannahs. opportunism, they seem unlikely victims of The several preferences between the different nutritional stress due to floral change. The kinds of proboscideans could explain why South American gomphotheres from the Early only the bunodont forms arrived to South to Middle Pleistocene exhibit similar feeding America when both types, lophodont and adaptation as modern elephants. Only popula- bunodont, lived together in North America, tions from the late Pleistocene show more previous to formation of the Panama isthmus focused feeding, but in general all exhibit connection. greater variability in the consumption of C3 Cuvieronius hyodon habitat could be high versus C4 plants. These observations match grassland with cold to temperate climatic the predictions of the Guthrie hypothesis and conditions, since in the inter-tropical zones it do not support the Graham & Lundelius has been only found at the highest levels, model. while in Chile it expanded to the littoral However, recent data from mastodons and zone, that surely offered similar living condi- mammoths from Florida (Koch et al. 1998) tions, in terms of temperature as the Andes show different patterns. Data suggest that corridor. This species would feed mainly on Mammut and Mammuthus partitioned resour- hard grasses, , and seeds from bushy ces and perhaps habitat. Mammut shows species. values that indicates a diet rich in C3 plants, Stegomastodon would seem to be adapted

358 PRADO et al.: South American Pleistocene Gompotheres

to more open grassland with warm to tempe- Ameghino, F., 1891 - Mamíferos y aves fósiles argenti- rate conditions, because their most austral nos. Especies nuevas, adiciones y correcciones - distribution does not surpass the 37th parallel Revista Argentina de Historia Natural 1: 240-259 in the Buenos Aires province (Tonni 1987). Ameghino, F., 1902 - Cuadro sinóptico de las formacio- The frequency of Stegomastodon platensis nes sedimentarias, terciarias y cretáceas de la diminishes in the Pampean Region by the Argentina en relación con el desarrollo y descenden- latest Pleistocene, when environmental condi- cia de los mamíferos - Anales del Museo Nacional, tions became colder and drier. In lower latitu- Buenos Aires 8: 1-12 des, where Stegomastodon is more frequent, Blainville, H.M.D. de, 1845 - Ostéographie ou descrip- it would occupy savannahs or xerophytic tion iconographique comparée du squelette et du pastures (Alberdi & Prado 1995). système dentaire des mammifères. III. Quaternatès. Des éléphants. - Paris, J. B. Baillière et Fils, pp. 1- ACKNOWLEDGEMENTS 367 We wish to thank the curators of the Museo Bocherens, H., Koch, P.L., Mariotti, A., Geraads, D. & de La Plata and the Museo Argentino de Jaeger, J.-J., 1996 - Isotopic biogeochemistry (13C, Ciencias Naturales ´Bernardino Ribadavia´, 18O) of mammalian enamel from African Pleistocene Argentina; the Museo de la Escuela hominid sites - Palaios 11: 306-318 Politécnica Nacional de Quito, Ecuador; also, Boeuf, O., 1992 - Anancus arvernensis chilhiacensis nov. the Museo Nacional de Historia Natural de subsp. (Proboscidea, Mammalia), un Mastodonte du Santiago, Chile. The present work was made Plio-Pléistocène de Haute-Loire, - Geobios possible through a joint Research Project MS 14: 179-188 from AECI, Spain - Argentina (1994-96); Boule, M., & Thevenin, A., 1920 - Mammifères fossiles projects PB94-0071 and PB97-1250 from de Tarija - Paris Imp.Nat., 256 pp. DGICYT, Spain; grants from the Universidad Bryan, A., 1986 - Paleoamerican Prehistory as seen from Nacional del Centro; project PIP-502-98 South America - in: Bryan, A. (ed.) - New evidence (CONICET); PICT-03-00000-00192 BID for the Pleistocene Peopling of the - 14 pp, 802/OC-AR, Argentina. University of Maine at Orono Bryan, A., Casamiquela, J., Cruxent, J., Gruhn, R. & REFERENCES Ochsenius, C., 1978 - An El Jobo Mastodon Kill at Alberdi, M.T., 1971 - Primer ejemplar completo de un Taima-Taima, Venezuela - Science 200: 1275-1277 longirostris KAUP, 1835, encontrado Burmeister, G., 1867 - Fauna Argentina, primera parte: en España - Estudios geológicos 17: 181-196 Mamíferos fósiles - Anales del Museo Público de Alberdi, M.T., & Prado, J.L., 1995 - Los mastodontes de Buenos Aires 1(2): 87-300 América del Sur - in: Alberdi, M.T., Leone, G. & Cabrera, A., 1929 - Una revisión de los Mastodontes Tonni. E.P. (eds.) - Evolución biológica y climática Argentinos - Revista del Museo de La Plata 32: 61- de la Región Pampeana durante los últimos 5 millo 144 nes de años. Un ensayo de correlación con el Casamiquela, R.M., 1972 - Catalogación crítica de Mediterráneo occidental - pp.277-292, Monografías, algunos vertebrados fósiles chilenos. II Los Museo Nacional de Ciencias Naturales, CSIC, Mastodontes - Ameghiniana 9(3): 193-208 Madrid Casamiquela, R.M., 1976 - Los vertebrados fósiles de Ameghino, F., 1888 - Rápidas diagnosis de algunos Tagua-Tagua, Chile - Primer Congreso Geológico mamíferos fósiles nuevos de la República Chileno, Santiago 88-102 Argentina - Obras Completas, Buenos Aires V: 469- Casamiquela, R.M. & Dillehay, T.D., 1989 - Vertebrate 480 and invertebrate faunal analysis - in: Ameghino, F., 1889 - Contribución al conocimiento de Dillehay, T.D. (ed.) - Monte Verde: a late Pleistocene los mamíferos fósiles de la República Argentina. settlement in Chile. Vol. I. Palaeoenvironment and Actas Academia Nacional Ciencias, Córdoba 6: 1- site context - pp. 205-210, Smithsonian Institution 1027 Press, Washington DC

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