,

- - - - _...._ _...

N. Jb. Geol. Palaont. Abh.

2007, vol. 243/2, p.179-189, Stuttgart, February 2007, published online 2007

Stegomastodon waringi (Mammalia, Proboscidea) from the Late Pleistocene of northeastern Uruguay Maria Teresa Alberdi, Madrid, José Luis Prado, Olavarría, Daniel Perea and Martín Ubilla, Montevideo With 6 figures and 3 tables

ALBERDI, M. T., PRADO, 1. L., PEREA, D. & UBILLA, M. (2006): Stegomastodon waringi (Marnmalia, Proboscidea) from the Late Pleistocene of northeastern Uruguay. - N. Jb. Geol. PaHiont. Abh., 243: 179-189; Stuttgart. Abstract: Fossil remains belonging to Proboscidea from the Cerro Largo Department (northeastern Uruguay), are described and taxonomically identified in this work. We compared these remains with those of Stegomastodon waringi and Stegomastodon platensis from several localíties in Brazil and Argentina using Discriminant Analysis. Based on general morphology and the resuIts of muItivariate analyses we considered this specimen as Stegomastodon waringi (HOLLAND, 1920), which along with previous reports significantly expand the southem geographic distribution of this gomphothere species in South America. Key words: Stegomastodon waringi, Marnmalia, Proboscidea, Upper Pleistocene, Uruguay, South America.

1. Introduction The knowledge of proboscidean gomphotheres from Uruguay is scarce and sorne of the recent advances from Uruguay were published by GUTIÉRREZ et al. (2005). Almost every year there are new discoveries of fossils that attempt to introduce additional discussion about recently published material. In this paper, we described a new almost complete skull from a strati­ graphic section in Cerro Largo Department (Uruguay; Fig. 1). Viewed in a geological time frame, the gompho­ theres may be considered part of the "last radiation" within "Great American Biotic Interchange" (WEBB 1991). In South America this family occurs from the Early-Middle Pleistocene (Ensenadan South American Land Mammal Age, hereafter SALMA) to the Late Pleistocene (Lujanian SALMA). Based on

001: 10.1127/0077-7749/2007/0243-0179

isolated remaÍns from Peru, CAMPBELL et al. (2000, 2001) suggested that gomphotheres arrived in South America during the Late Miocene. In our opinion, the chronology ofthese data is inconclusive (se e ALBEROI et al. 2004). The family Gomphotheriidae is considered a long­ living ancestral stock that gave origin to a succession of other groups. North America played a significant role in the history of gomphothere biogeography and diversity (LAMBERT 1996). From the Middle Miocene to the Pleístocene (late Barstovian to Rancholabrean NALMA) numerous taxa immigrated to this continent from the Old World vía Beringia. Gomphothere diver­ sity also reached its peak during this time span. There were six known genera from the late C1arendonian to the early Hemphíllian NALMA followed by a re­ duction in the number of genera during the late Hemphillian (WEBB 1983).

0077 -7749/07/0243-0179 $ 2.75 ©2007 E. Schweizerbart'sche Verlagsbuchhandlung, D-70176 Stuttgart

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Recently, PRADO et aL (2002,2003,2005) presen­ ted a simplifíed c1assification of South American gomphotheres modified after SIMPSON & PAULA COUTO (1957) and recognized one species of Cuviero­ nius: C. hyodon (FISCHER, 1814), and two species of Stegomastodon: S. waringi (HOLLAND, 1920) that was recorded from Uruguay, Brazi1 and Ecuador (GUTIÉRREZ et al. 2005; ALBERDI et al. 2002); and S. platensis (AMEGHINo, 1888) that was recorded from Argentina, Uruguay, and probably from Paraguay (PRADO et aL 2005).

2. Stratigraphic setting and age The fossils were collected from the Dolores For­ matíon. Thís unÍt was defined by Goso (1972 vide ANTÓN & Goso 1974) and 1S characterized by silt­ stones, pelites, sandy to gravely pelites and sandstones with argillaceous matrix. The prevailing colour is brown with local changes to gray and grayish green (PRECIOZZI et al. 1985; BOSSI & NAVARRO 1991). The lithologica1 characteristÍCs of this unit in the collecting site are described in Fig. 2. The Dolores Formatíon is considered to be Late Pleistocene in age, according mammalian content and radiocarbonic

Fig. 1. Geographical location of the Cam­ po Viñoles locality (Cerro Largo depart­ ment, Uruguay, South America) with Stego­ mastodon waringi in which is indicatcd an approximate location of the collecting site.

dates which range from 17,600 to 10,500 years BP (UBILLA 1999; UBILLA & PEREA 1999; GUTIÉRREZ et aL 2005). The late CenozoÍC pampean continental mamma­ lian biostratigraphy involves three chronostratigraphic units described for the Buenos Aires Province, com­ monly used to propose regional correlations (ClONE & TONNI 1999,2005): Ensenadan (Late Pliocene to early Middle Pleistocene), Bonaerian (Middle Pleistocene) and Lujanian (Late Pleistocene to Early Holocene) (Fig. 2). The Lujanian Age 1S supported by the Equus (Amerhippus) neogeus biozone. According the mammalian fauna of the Dolores Formation, it is correlated to the Lujanian Land Mammal Age of Buenos Aires Province (Fig. 2). Though a radiocar­ bonic analysis of enamel of the FC-DPV-1804 failed (Laboratory of Centrum voor Isotopen Onderzoek, Groningen, Nether1and), there are others C14 avail­ ables (GUTIÉRREZ et al. 2005; UBILLA & PEREA 1999) that along with the palaeontological content ofthe unit allow us to regard the bearing deposits as a Lujanian deposit. The fauna of the Dolores Formatíon of Uruguay includes the characteristic Equus (Amer­ hippus) neogeus and other exclusive taxa as Doedi­ curus clavicaudatus.

Stegomastodon waringi from the Late Pleistoeene of northeastern Uruguay

PAMPEAN AGES

'LATAN

------ 1-----, IONAr,.,,,"

SANANDIIlSlAN

Dolores Formatlon (Late Pleistocene) (whitish-graylsh medium to coarse sandstone with basal conglomerate)

+-

Stegomastodon waringi

Yaguari Formation (Permian)

o Fig. 2. Stratigraphie seheme showing the eorrelation of the Land Mammal Ages from South Ameriea (CIONE & TONNI 2005), and the stratigraphic sequenee at the Chacra or Campo Viñoles locality (Cerro Largo department, Uruguay, South Ameriea). Arrow indieates the location where the skull of Stegomastodon waringi was collected.

3. Taxonomic background CUVIER (1806), BOULE & THEVENIN (1920), CABRERA (1929), and HOFFsTETTER (1952) have studied South American gomphotheres in monographs. SIMPSON & PAULA COUTO (1957) considered Cuvieronius hyodon, Haplomastodon waringi, and Stegomastodon platen­ sis the only genera and species that seem to be clearly distinct among South American gomphotheres. TASSY & SHOSHANI (1996: Appendix B) and SHOSHANI & TASSY (2005) proposed a new proboscidean classifi­ cation and placed the South American gomphotheres in the Subfamily Cuvieroniinae CABRERA, 1929 (Family Gomphotheriidae HAY, 1922). In the same book, TASSY (1996) proposed a phylogeny of the

181

Order Proboscidea that includes two gomphothere groups (both paraphyletic): one for Old World gom­ photheres, and another for New World gomphotheres. He included a11 South American forms in the New World gomphotheres, without assigning them to a special family or subfamily (TASSY 1996; SHOSHANI & TASSY 2005). PRADO et al. (2002, 2003, 2005) and ALBERDI et al. (2002, 2004) presented a simplified classification of South American mastodonts, modi­ fied after SIMPSON & PAULA COUTO (1957), and reco­ gnized Cuvieronius hyodon and two Stegomastodon species: S. waringi and S. platensis. HOFFsTETTER (1952) proposed the subgenus Haplomastodon, based on the presence of transverse foramina in the atlas and axis. We found this to be a highly variable character (several specimens from Argentina and Brazil showing foramen or not); neither do tooth characters justify maintaining Haplomastodon as a different genus. Anyway, the relationships between these taxa is not yet clear.

4. Material and methods The gomphothere sku11 from Cerro Largo is stored at the Vertebrate Palaeontology Co11ection of Facultad de Ciencias, Universidad de la República, Uruguay. It consist of the complete sku11 with left and right M3 (FC-DPV-1804). We selected the more important characters and parameters in ALBERDI et al. (2002) to compare the specimens of the most important localities from the Middle and Late Pleistocene of South America (Fig. 3). For comparisons we used the remains from selected localities in Brazil (Pains, Minas Gerais), in Argentina (mainly in the Pampean Region), in La Huaca (Peru) with good stratigraphic data. These remains belong to the fo11owing co11ections: MLP: Museo de La Plata, Argentina; MMO: Museo Munici­ pal "Dámaso Arce" de Olavarría, Argentina; MACN: Museo Argentino de Ciencias Naturales "Bemardino Rivadavia", Buenos Aires, Argentina; MUCBH: Museo de la Pontifical Universidad Católica de Belo Horizonte, Brazil; MNHN: Museo de Historia Natural de la Universidad Nacional San Carlos, Lima, Peru. A11 dimensions are expressed in mi11imetres. 36 M3 were analysed using SPSS version 11.5. The authors will provide the original data underupon request. Data were analysed by Discriminant Analysis (hereafter, DA). This analysis is used when the purpose is identi­ fied of specimens of unknown affinity to a priori groups. Further details ofthis method are in REYMENT

-----------182

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------------_...

M. T. Alberdi et al.

10

Fig. 3. Nomenclature and

graphic representation of the measurements of the gomphothere skull, modí­ fied from ALBERO! et al (2002). For explanation of the numbers see Table 2.

(1991). Also, we used the robustness index of the molars defined by OSBORN (1936) because this is a good discriminant character. 5. Results of multivariate analysis Stegomastodon platensis and Stegomastodon waringi have been identified in a previous paper using teeth morphology and bivariate analysis of teeth measu­ rements (ALBERDl et al. 2002). Now we used DA to

establish a rule for differentiating among these species base on teeth measurements. DA of these groups pro­ vided identification from M3 sample (Fig. 4, Table 1). The material from Brazil and Ecuador assigned to Stegomastodon waringi is distinguished in part from the specimens in Argentina. We compared also these Uruguayan teeth with the rest of the sample, using the robustness index defined by OSBORN (1936). This index was calculated for each molar following the formula: maximum width x 100/length, and currently

- - - - - - - - - - - - - _ . _ - _ ._ _

......

Stegomastodon waringi from the Late Pleistocene of northeastern Uruguay

Discriminate analysis (M3) 2~------------------~

O

183

Table 1. Results of the c1assification of the discriminant analysis. 1: Stegomastodon platensis (late); 2: Stegomasto­ don platensis (mid); 3: Stegomastodon waringi; 4: Teeth ofthe Cerro Largo skull (ungrouped cases). Predicted Group Membership

o

O

+ Certroííls

-1

a

•

eIV

'ti In

o

Cerro Largo

O S.waríngi

,5 -2 E

El S. platensis (mííl)

%

-3 ~~_ _-".~_~~--I O S. platensis (late) -2

-1

o

2

3

3

Total

4

O

4

8

2

O

11

5

16

3

4

4

2

10

ungrouped cases

O

O

2

2

1

50

O

50

100

2

O

68,8

31,3

100

3

40

40

20

100

ungrouped cases

O

O

100

100

Count

N

e .2 Ü e

2

4

Dlscrlmlnant fundion 1

Fig. 4. Discriminant analysis (DA) ofupper M3 samples of Stegomastodon platensis from Ensenadan SALMA in Argentina (S, platensis (mid)); Stegomastodon platensis from Lujanian SALMA in Argentina (S. platensis (late)); and Stegomastodon waringi from Lujanian SALMA in Brazil and Ecuador (S. waringi). Upper M3 from Uruguay is noted as a black circ1e.

known as the molar crown index (HILLSON, 1986). The Mann-Whitney U test was applied in order to find significant differences among these values. This test is very useful in comparing the medians of small samples, We found that when measuring maximum width x 100/length the third molar (the only molars in this skull) provided a correct identifícation of both specíes of Stegomastodon (Fig. 5). Based on this analysís we assigned this skull to Stegomastodon waringi.

6. Systematic paleontology We observed that all South American gomphotheres show a generalized pattern that inc1udes the following features: a brachycephalic skull with a tendency towards an elephantoid skull, a brevirostrine jaw with a slightly curved symphysis, elongate upper tusks which vary from straight or slightly curved to very twisted, and that may or may not be present an enamel bando Within South American gomphotheres, only adult Cuvieronius hyodon conserve a true enamel bando The alternate contact between successive pre­ trite and postrite half-Ioph(id)s (anancoidy) is present

frequently in the last loph(íd)s on M3 and m3. These molars are brachyodont or subhypsodont, with simple to complex trefoíls on the occ1usal surface. M2 is trilophodont and occasionally has a developed talon. M3 varíes from tetralophodont to heptalophodont. For description of the genus Stegomastodon, see ALBERDI et al. (2004). Stegomastodon waringi (HOLLAND, 1920) Fig.6

1842 1906 1920 1920 1922 1928 1931 1948 1952

Mastodon brasiliensis LtJND...... LEssoN, p. 157, therein considered as nomen nudum. Mastodon andium. WINGE, p. 48, non CUVIER 1806: 413. Mastodon humboldti CUVIER.·-BoULE&THEVENIN, p.68. Mastodon waringi. ~ HOLLAND, p. 229. Mastodon chimborazi. ~ PROAÑO, p. 13. Bunolophodon ayora. ~ SPILLMANN, p. 70. Bunolophodonpostremus. SPILLMANN, p. 73. Notiomastodon vidalí. CASTELLANOS, p. 139. Haplomastodon (Haplomastodon) chimborazí. HOFFSTETTER, p. 192.

1952 1952

1952 1957 1995

Haplomastodon (Aleamastodon) guayasensis.

~

HOFFSTETTER, p. 208. Especies B de La Carolina (Ecuador). ~ HOFF­ STETTER,p.224. Stegomastodon brasiliensis. ~ HOFFSTETTER, p.222. Haplomastodon waringi (HOLLAND, 1920). SIMPSON & PAULA COUTO, p.17l. Haplomastodon chimborazi (PROAÑO, 1922). ~ FICCARELLl et al., p. 753-754.

--------------------_ _ _ ...

184

50 48

-

46 44

42 ~ 40 CIl

c:

ii) 38

~

36L-____r -_ _ _~------~--~ 10

S. platensís

S. waringi

Cerro Largo

Species Fig. 5. Results of the robustness index on the upper M3 of the Uruguay teeth following OSBORN (1936), compared with other samples from South America.

1996 1996

_ _...._

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_

....

M. T. Alberdi et al.

52~---------------~

i!o

.....~ ..

Haplomastodon waringi (HOLLAND, 1920). ­ CASAMIQUELA et al., p. 316. Amahuacatherium peruvium. - ROMERO-PITTMAN, p. 171-178.

Holotype: Corresponds to an incomplete mandible and bone fragment (CM-ll033) from Pedra Vermelha, Bahía, Brazil described by HOLLAND (1920: 229). SIMPSON & PAULA COUTO (1957: 172) further commented that there were three broken molars, two tusk fragments, and the distal part of a tibia - al! with the number CM-11033. Stratigraphic distribution: Middle and Late Pleisto­ cene of South America (probably only on the Late Pleisto­ cene because same localities in Brazil need more stratigra­ phic accuracy) (see ALBEROl et al. 2002; PRADO et al. 2001, 2002, 2005; GUTIÉRREZ et al. 2005). Geographic distribution: It is recorded from Ecuador (Santa Elena Península and Quebrada Pistud), possibly from Taima-Taima (Venezuela), Peru (La Huaca, Quipan, Pasamayo; ALBEROI et al. 2004), Brazil (Paíns, Lagoa Santa area in Minas Gerais, Toca dos Ossos in Bahía, Bonito in Mato Grosso do Sul, and possibly from Río Grande do Sul; ALBERO! et al. 2002), and from Uru­ guay (Dolores Formation, Montevideo, GUT!ÉRREZ et al. 2005). Diagnosis: Short and hígh skull, elephantoíd type; less elephantoid than the type species (Stegomastodon mirificus)

and less depressed than Cuvieronius. The molar occlusal drawing surfaces are in general less complicated than in Stegomastodon platensis. The double trefoil pattern (on both pretrite and postrite) varíes from simple to relativeIy complex due to the presence of secondary conules or conelets. Tusks are nearly straight to very slight1y curved, with an enamel band in some juvenile índividuals; usually simply curved and without enamel band in adults. AH these features are less marked than in S. platensis. Description of specimen from Cerro Largo: The skull corresponds to a big and old animal (FCDPV-1804), with a right and left M3, without tusks, and with a small alveolus from right M2 (Fíg. 6). The cranial vault, partially preserved, is slight1y convex, and conserved only the right zygomatic arch. The posterior or nucal area of the skull is broken and don't conserve the nucal fosse and the crest of the nucal ligament fosse. In the ventral view, the edges of the sagittal plane on the premaxilla of the Cerro Largo spe­ cimen is well marked, parallels, similar to a central fissure. It is clearly less divergent than the sagittal plane on Cuvier­ onius skulls from Tarija. The tusk alveoli diverge slight1y, a character similar to that observed on another Stegomasto­ don skuIl from the Buenos Aires province (MLP-8.2, MLP­ 8.4, and MEDA-AT-OOlO, see PRADO et al. 2002), from Pains, Brazil (ALBERO! et al. 2002), and from La Huaca, Peru (ALBEROI et al. 2004) (Table 2). This anterior part of the symphysis (alveolar part in the maxilar) is nearly straight compared with the divergent symphysis on Cuvier­ onius skulls from Bolivia. The face is incomplete and we can not illustrate. The Cerro Largo skull measurements indicate a large-sized animal (Table 3). The cross sections of the tusks are 150 x 132 mm and 150 x 123 mm, respectively from the alveolar point and slightIy ovals in shape. The base ofthe alveoli indicates that the tusks were slightly divergent. More than measures in table 2 we can take the following variables: maximal diameter of the orbit = 194 mm; maxi­ mal width ofthe mesial premaxilla plane 597 mm; maxi­ mum width at the ventral hemimaxilla view, right = 291 mm ca, and left 297 mm ca; distance between the mesial maxi­ lar plane and the palatine spine in sagittal pIane = 730 mm. The M3 have four and half lophs and a reduced talon and show signs of advanced wear. They are wide and the mid­ line is rather lost on the first, second and third lophs of the right M3 (better preserved at the left M3). There are multi­ pie internal central conules faded by wear forming the cha­ racteristic trefoil pattern very complex and with the enamel very plicate characteristic of the more evolved forms ~ ptychodonty - characteristic of S. platensis. Even though, the occIusal surface is cIoser to S. platensis than S. waringi, the robustness index indicate a very significant difference and is very close to S. waringi (Fig. 5). In this latter figure on can see the proximity between this index in this speci­ men and the index in S. waringi, very far the index in S. platensis. That is a very good character to detach between these species. They have on the edges cement residues. The mesial cingulum is lost through wear, and the lingual cingu­ lum is strongly marked in the first and second valley of left M3. They valleys on the right M3 and third and fourth valleys on left M3 are completely filled by the wear. Dental dimensions of the left and right M3 are show in Table 3.

Stegomastodon waringi from the Late Pleistocene of northeastern Uruguay

e

185

D

5cm

30cm

Fig. 6. Stegomastodon waringi (HOLLAND, 1920) (Vertebrate Paleontology Collection of Facultad de Ciencias, Universidad de la República, Montevideo, FCDPV-1804); A - ventral view of skull; B - detail of the anterior part of the maxilla; e - detail of the left M3, in occlusal view; D - detail of the right M3, in occlusal view.

7. Conclusions In the multivariate analysis we observed a pattern of increased teeth size since Ensenadan remains from Argentina to the Lujanian remains, but the morpho­ logical characters in both populations are homo­ geneous. And the DA results plus the robustness index provided a rule for differentiating among the species based on teeth measurements, concretely: 1) Stego­ mastodon platensis from Ensenadan SALMA in Argentina; (2) Stegomastodon platensis from Luja­ nian SALMA in Argentina; and (3) Stegomastodon waringi from Lujanian SALMA in Brazil, Peru, Uruguay, and Ecuador. The gomphotheres skull from Cerro Largo (Uru­ guay) is typical ofthe genus Stegomastodon.A1though,

the double trefoils pattern and the enamel very plicate (ptychodonty) link with the high variability of gom­ photheres teeth are more characteristic than S. platen­ sis, the robustness index is clearly significant to assign it to Stegomastodon waringi without doubt, thus confirming the previous report of Stegomastodon waringi to Uruguay by GUTIÉRREZ et al. (2005). Stego­ mastodon waringi is one of the largest species of South American gomphotheres, which appears to have predominated at middle latitudes (in Uruguay, Brazil, Peru and Ecuador), where it occupied savannas or xerophytic open areas, and consequently would have been better adapted to warm or temperate climatic conditions. Carbon isotope analyses for another speci­ men (Dolores Formation) reveal that Stegomastodon waringi from Uruguay was possibly a mixed-feeder

~

186

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.. _ _ _ _ _ _ _ _ _ _ _ _..JI

M. T. Alberdi et al.

Table 2. Measurements [in millimetres] of the Stegomastodon skull from the Cerro Largo (Uruguay, FCDPV-1804), and comparatively other skull from the Buenos Aires Province (MLP-8.2, MLP-8.4, and MEDA-AT-OOIO), from Pains (Brazil, MUCBH-sn), and from La Huaca (Peru, without number). The numbers indicate the following measurements; dorsal view: 1 - Maximal length of the skull in the sagittal plane; 2 Distance between the vertex and the upper plane of the nasal apertures; 3 - Maximal nasal height; 4 - Premaxilla length in the sagittal plane; 5 Minimum width between the temporal crests; 6 Maximal suborbital width; 7 - Premaxilla width at the level of the infra-orbital foramen; 8 Width of the nasal process; 9 Maximal width of the nasal aperture; ventral view: 10 - Width between the external auditive pseudo-meatus; 11 Width between external margins of the mandibular articulation fosses; 12 - Width between internal margins of the mandibular articulation fosses; 13 Width between antero-internal margins of the molars; 14 Width between postero­ internal margins of the molars; 15 Width of the choana; 16 Distance between the molar anterior plane and the palatine spine, sagittal plane; 17 - Distance between the palatine spine and the anterior margin of the foramen magnum; 18 Distance between the pterygoid process of the sphenoid and the anterior margin of the foramen magnum; 19 Maximal antero-posterior diameter of the temporal fosse - view from yugal arcade; lateral view: 20 Distance between the highest point of the temporal crest and the antero-inferior margin of the external auditive pseudo-meatus; 21 Distance between the orbital ventral point and the antero-inferior margin of the pseudo meatus auditive external; 22 Maximal orbital height; 23 - Distance between the highest point of the temporal crest and the alveolar margin; 24 Distance between the premaxilla antero-inferior margin and pterygoid process; 25 - Distance between the extreme of the frontal zygomatic process and the pterygoid process; 26 - Distance between the vertex and the pterygoid process; 27 - Distance between the posterior margin of the occipital condyles and the pterygoid process; 28 - Antero-posterior maximal diameter of the premaxilla; posterior view or external cranial vault: 29 - Maximal width of the cranial vauIt; 30 Width of the nucal ligament fosse; 31 Width between latero-external margins of the occipital condyles; 32 - Width between latero-internal margin of the occipital condyles; 33 Maximal height of the skull (from the vertex to the externo-inferior border of the occipital condyles; 34 - Distance between vertex and the top of the nucal Iigament fosse; 35 - Length of the crest of the nucalligament fosse.

measures

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

980

304

163

580

207

520

448

112

214

660

540

200

140

102

79

229

326

Tapalqué

910

267

85

430

288

540

310

89

290

425

508

214

125

82

75

307

322

MLP-8.2

1125

570

120

540

410

730

414

70

410

750

640

290

101

104

100

330

370

MLP-8.4

750

305

90

440

289

470

264

60

238

190

117

110

55

250

347

La Huaca

918

214

149

580

390ca

650

392

533

518

328

126.5

111

103

251

291

385 ca

685

476

680

570

221

167

164

85,5

340

362

Paíns

Cerro Largo

102ea 246ea

540

measures

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

35

Pajns

246

293

445

402

217

825

690

476

650

395

216

775

83

227

80

495

126

130

Tapalqué

313

250

360

280

105

415

500

426

450

370

156

580

75

209

74

330

122

129

MLP-8,2

350

310

500

340

210

760

480

460

840

420

183

920

71

254

80

440

106

176

MLP-8.4

315

290

240

333

144

410

485

390

480

360

90

440

88

195

78

110

112

La Huaca

219ca

257

350

342ca

145

425ca

655

351,5

462

311,5

150

>629

70

206,5

83

94ea

100ca

Cerro Largo

426 ca

311

658

270 ca

233

66

173

similar to Brazil samples ofthis species. The presence of Stegomastodon waringi in this quarry represents a new record for Uruguay and expands the known southern distribution of this species in South America. Acknowledgements We thank Mercedes GUTIÉRRREZ for her valuable sug­ gestions, Federico LÓPEZ, "Capincho" VIÑOLES, Gustavo

454 ca 160 ca

430

LECUONA, Felipe PÉREZ, Nora LORENZO, and Graciela PIÑEIRO for their collaboration in collecting and preparing the material. We also thank to Maria Rita PALOMBO and Pascal TASSY for their critical revision of the manuscript and valuable comments. Part of graphic materials and illustrations were created at the photographic laboratory of MNCN and A. RINDERKNECHT is thanked for the electronic management ofFig. 6. The present work was made possible by a joint Research Project from AECI, Spain Argentina (2000-2001); Projects PB94-0071, PB97-1250, BTE2001­

...

_---_..

~-~--_

..

~-_.-----_

..

~

187

fram the Late Pleistocene of northeastern

Table 3. Measurements [in míllimetres] of the right and left M3 of Stegomastodon skull from Cerro Largo Department (Uruguay). L: maximal length; W 1: width of the first loph; W2: width of fue second loph; W3: width of the third loph; W4: width of the fourth loph; W5: width of the fiffu loph; T: talon; > W: the maximum width; r = right; 1 = left. FCDPV: Facultad de Ciencias, Departamento de Paleontología de Vertebrados, Universidad de la República (Uruguay). Col. Number

Locality

kind of tooth

L

W1

W2

W3

W4

T

W>

sn-cranial

La Huaca, Piura

M3 upper I

185

87,5

89

88

75

56

89

sn-cranial

La Huaca, Piura

M3 upperr

185

92

92,5

89

83

43,5

92,5

MLP-8.2

M3 upper I

195

86

90

94

85

58

94

MLP-8.2

M3 upper r

192

83,5

85

89

81

54

89

230-235 215

97 87

96,5 90

96 82

83 68

48,7

84

97 90

W5

Ste¡;¡omastodon

sn-cranial

Pains, Brazil

M3 upper I M3 upper r

sn-cranial

Pains, Brazil

M3 upper I

218

83

89,6

92

82,5

67

FHCDPV-1804

Cerro Largo

M3 upperd

248

109

109,5

103

96

78

30

109,5

FHCDPV-1804

Cerro Lar¡;¡o

M3 ueeer I

235

106

106

104

90

73

31

106

FCDPV-1803

Canelones

92

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n

n

----------------------188

~--~

..........

----­

M. T. Alberdi et al.

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189

Addresses of the authors:

MARIA TERESA ALBERDI, Departamento de Paleobiología.

Museo Nacional de Ciencias Naturales, CSIC, José

Gutiérrez Abascal 2, 28006 Madrid, Spain;

E-mail: [email protected]

JosÉ LUIS PRADO, INCUAPA-Departamento de Arqueo­

Manuscript received: April19th, 2006.

Revised version accepted by the Stuttgart editor: November logía, Universidad Nacional del Centro. Del Valle 5737,

87400JWI Olavarria, Argentina;

8th, 2006.

E-mail: [email protected]

DANIEL PEREA, MARTIN UBILLA, Paleontología, Facultad de

Ciencias, 19uá 4225, 11400 Montevideo, Uruguay;

E-mails:[email protected]@fcien.edu.uy.