salas, stucchi y devries 2003 - palaeolama

8/3/2019 Salas, Stucchi y Devries 2003 - Palaeolama

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347PERU: PRESENCE OF PLIO-PLEISTOCENE PALAEOLAMA SP. ON THE SOUTHERN COASTBull. Inst. fr. tudes andines

2003, 32 (2): 347-359

THE PRESENCE OF PLIO-PLEISTOCENE

PALAEOLAMA SP. (ARTIODACTYLA: CAMELIDAE)ON THE SOUTHERN COAST OF PERU

Rodolfo SALAS* ** , Marcelo STUCCHI** , Thomas J. DEVRIES***

Abstract

We report the oldest record of the genus Palaeolama in Peru; its constitutes one of the

oldest records for the genus in South America. The shape and size of the fossil material iscomparable to South American forms P. (Hemiauchenia) paradoxa and the smallest forms ofP.(Palaeolama) weddellii. The bones were found within unnamed littoral marine deposits nearAtiquipa, on the south coast of Peru. The occurrence of the extinct bivalve mollusk, Chlamysvidali, at the base of the section and coquinas of exclusively extant mollusks in the highest marineterrace capping the section indicates an age of about two million years. This age is slightly olderor equal to the oldest records ofP. (Palaeolama) in North America. The present report confirmsthat the Lamini were no strangers to Andean ecosystems prior to the Holocene.Key words:Mammalia, Camelidae, Palaeolama, Plio-Pleistoceno, Atiquipa, Per.

PRESENCIA DEPALAEOLAMA SP. (ARTIODACTYLA: CAMELIDAE) DEL PLIO-PLEISTOCENO EN LA COSTA SUR DEL PER

Resumen

Se presenta el registro ms antiguo del gnero Palaeolama en el territorio peruano y unode los ms antiguos de Sudamrica. La forma y tamao del material conservado es comparablecon las formas sudamericanas P. (Hemiauchenia) paradoxa y los especmenes ms pequeos deP. (Palaeolama) weddellii. Procede de los depsitos marinos litorales innominados de Atiquipa,en la costa sur del Per. La ocurrencia del molusco bivalvo extinto Chlamysvidali en la base dela seccin y la coquina de moluscos actuales presentes en la terraza marina ms alta que terminadicha seccin, indicara una edad aproximada de 2 millones de aos. Esta edad es ligeramentemayor o igual a los registros ms antiguos de P. (Palaeolama) en Norteamrica. El presente

reporte confirma que los Lamini no fueron ajenos a los ecosistemas andinos antes del Holoceno.Palabras claves:Mammalia, Camelidae, Palaeolama, Plio-Pleistoceno, Atiquipa, Per.

* Instituto Francs de Estudios Andinos IFEA, Casilla 181217 Lima 18.** Departamento de Paleontologa de Vertebrados. Museo de Historia Natural - U.N.M.S.M.

(Lima, Per); E-mail por orden alfabtico: [email protected],[email protected].***Burke Museum of Natural History and Culture, University of Washington (Seattle, WA USA).


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348 Rodolfo SALAS, Marcelo STUCCHI, Thomas J. DEVRIES

PRSENCE DUPALAEOLAMA SP. (ARTIODACTYLA: CAMELIDAE) DU PLIO-PISTOCNE SUR LA CTE SUD DU PROU

Rsum

Nous prsentons un spcimen du genre Palaeolama qui correspond au registre le plusancien du genre sur le territoire pruvien ainsi que lun des plus anciens en Amrique du Sud. Lamorphologie et la taille des lments dcouverts sont comparables aux formes sudamricainesP. (Hemiauchenia) paradoxa ainsi quaux formes les plus petites de P. (Palaeolama) weddellii.Ce spcimen provient des dpts marins littoraux dAtiquipa (cte sud-pruvienne). La prsencedu mollusque bivalve Chlamysvidali la base de la section ainsi que de faluns coquilliers actuels,prsents au niveau des terrasses marines les plus hautes qui terminent la section, correspondrait un ge de 2 millions dannes. Cette date est lgrement suprieure ou gale aux registres lesplus anciens de P. (Palaeolama) en Amrique du Nord. Cette dcouverte confirme que les Laminitaient prsents dans les cosystmes andins avant lHolocne.

Mots cls :Mammalia, Camelidae, Palaeolama, Plio-Pleistocne, Atiquipa, Prou.

INTRODUCTION

The evolution of camelids took place principally in North America. Theirmigration to South America probably occurred during the late Pliocene (Hoffstetter,1952). The earliest material was assigned tentatively to the Plio-Pleistocene of Argentina(Cabrera, 1935; Kraglievich, 1946; Marshall et al., 1984) and Uruguay (Mones, 1988).

Descriptions of the genera Palaeolama GERVAIS, 1867, and HemiaucheniaGERVAIS & AMEGHINO, 1880, were based on material collected from the ArgentinePampas. Cabrera (1935) and Hoffstetter (1952) agreed in consideringHemiauchenia assynonymous with Palaeolama. Webb (1974) distinguished Hemiauchenia fromPalaeolama as a separate genus and tried both diagnosis. Most recently, Gurin & Faure(1999) designatedHemiauchenia as a subgenus ofPalaeolama.

The best documented and oldest account ofP. (Palaeolama) from South Americacomes from the Ensenadan of Tarija, Bolivia (Webb & Stehli, 1995). In contrast,P. (Hemiauchenia) was present in the upper Uquian, having immigrated during thesecond panamerican dispersal that occurred between 2,0 and 1,9 Ma (Marshall, 1985).In North America, P. (Hemiauchenia) is recognized from the Hemphillian (lowerPliocene), while P. (Palaeolama) is well represented in the Irvingtonian (lowerPleistocene) in shell banks from Leisey, Florida (Webb & Stehli, 1995).

There are only two reliable reports ofPalaeolama from the coast of Peru, bothupper Pleistocene. The first report describes material from deposits of La Brea, nearTalara (Lemon & Churcher, 1961), which was assigned by Churcher (1965) and Webb(1974) to P. aequatorialis HOFFSTETTER, 1952. The second refers to material identifiedby Hoffstetter collected from the Pampa de los Fsiles, La Libertad (Marshall etal., 1984).


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349PERU: PRESENCE OF PLIO-PLEISTOCENE PALAEOLAMA SP. ON THE SOUTHERN COAST

The objective of this paper is to describe material ofPalaeolama(sensuGurin& Faure, 1999) from the Plio-Pleistocene of Peru and discuss its biostratigraphic andbiogeographic implications.

The chronology of mammal ages for South America used in this study is thatproposed by McKenna & Bell (1997).

Institutional abbreviations:MUSM: Museo de Historia Natural Universidad Nacional Mayor de San

Marcos, Lima.

UF:Florida Museum of Natural History University of Florida, Florida.

1. GEOLOGY AND AGE

The camelid bones were found in an outcrop of conglomerate and bioclastic

sandstone exposed in a roadcut along the Panamerican Highway on the northwesternside of Quebrada Infiernillo (SITE 596, Fig. 1). The lower part of the section (6,6meters), which rests upon a seaward-tilting igneous platform; consists of a progradingsequence that represents shallow subtidal, lower shoreface, upper shoreface, andfluvial/alluvial environments characteristic of a high-energy beach fronting an attenuatedflood-prone alluvial fan.

The middle part of the section (7,0 meters) is a transgressive succession oflaminated bioclastic gravel, lenticular bioclastic gravel, and several meters ofmassive, poorly sorted, coarse-grained bioclastic sandstone with dispersed,transported whole fossils of mollusks, barnacles, and the camelid bones. Themassive sands may had been accumulated in a small embayment at the mouth ofQuebrada Infiernillo that captured debris washed in from the Pacific Ocean anddown from the Andean foothills.

The upper part of the section includes a regressive sequence of cross-beddedbioclastic cobbly gravel and sandstone (2,2 m) overlain by a transgressive andregressive cycle (>3,2 meters) with similar deposits. A coquina that caps the secondregressive sequence also constitutes the surface of the most elevated marine terrace,which stands 250 meters above sea level.

The oldest mollusks from the Atiquipa outcrop, disarticulated valves ofChoromytilus chorus (MOLINA, 1782) and Chlamys vidali (PHILIPPI, 1887), are

wedged between boulders at the base of the section. C. vidali is found in lower andupper Pliocene strata of Chile (Herm, 1969), lower Pliocene shoreface depositsnear Sacaco (Muizon & DeVries, 1985), and on the surface of the oldest marineterraces of northern and southern Peru (DeVries, 1986). Absent from the base ofthe Atiquipa section are mollusks that signify a lower Pliocene age, e.g.,Acanthinatriangularis DEVRIES, 1986;Herminespina mirabilis (PHILIPPI, 1887);H. saskiaeDEVRIES & VERMEIJ, 1997; Concholepas kieneri HUPE, 1859; and C. nodosaMORICKE, 1896 (Devries & Vermeij, 1997; Devries, 2000).


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Fig. 1 - Map of the area of Atiquipa showing SITE 596 where the camelidmaterial was found.

The bone-bearing sandstones contain an assemblage of mollusks that includesConcholepas camerata DEVRIES, 2000, which indicates a latest Pliocene or earlyPleistocene age. The coquina that caps the section at Atiquipa contains an assemblageof the extant mollusks Glycymeris ovata (BRODERIP, 1832),Eurhomalea lenticularis(SOWERBY, 1835), Mulina edulis (KING, 1831), and Oliva peruviana LAMARCK,1811. Elsewhere along the coast between Chala and Caman, these species occurin a comparable geological setting together with the extinct gastropod, Chorusgrandis (PHILIPPI, 1887), suggesting a pre-Pleistocene age (Devries, 1997) (Fig. 2).


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Fig. 2 - Stratigraphic column of the SITE 596 section near Atiquipa.


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2. SYSTEMATIC PALEONTOLOGY

Order ARTIODACTYLA OWEN, 1848

Family CAMELIDAE GRAY, 1821

Subfamily CAMELINAE GRAY, 1821

Tribe LAMINI WEBB, 1965

PalaeolamaGERVAIS, 1867

Material MUSM 51 - Departamento de Paleontologa de Vertebrados, Museo deHistoria Natural (UNMSM); associated distal portions of the left femur and right humeruswithout the proximal epiphysis. Collected by Thomas DeVries and Marcelo Stucchi.

Locality 154842"S, 742125"W; SITE 596, Atiquipa (Panamerican Highway,km. 596 south), 200-250 meters above sea level, Arequipa Department, Peru.

Age Late Pliocene to earliest Pleistocene (2,2 1,6 Ma).

2. 1. Description

Humerus In the humerus MUSM 51 (Fig. 3 A, B), the lateral epicondyle ismore robust than the medial epicondyle, in contrast to that ofVicugna vicugna MOLINA,1782, where the two are subequal. From a cranial perspective, the articular trochlea iscompressed proximodistally. The interlabial longitudinal distance-distal articulartransversal distance index in MUSM 51 is 39; in V. vicugna it is 48. The axial throat isdeep. In relation to the medial lip, the lateral lip extends proximally more than inV. vicugna and P. weddellii. The epiphysesof the lateral and medial epicondyles arepartially fused.

The preserved part of the MUSM 51 humerus does not exhibit significantdifferences in shape or proportion from any of the Palaeolama species. Among thesouthamerican species, it is practically equal in size to the smallest forms ofP. weddelliiGERVAIS, 1855 of the Pleistocene of Ecuador (Hoffstetter, 1952) and the average ofP. paradoxa (GERVAIS & AMEGHINO, 1880). P. majorLIAS, 1872 has only been able tobe compared in one of its dimensions. The total length of the MUSM 51 humerus isestimated to be 270-300 mm (Table 1).

Femur The distal epiphysis of the MUSM 51 femur is robust (Fig. 3 C). Thefemoral trochlea is symmetrical and perfectly parallel with the axial axis. Although thelateral condyle is only partially preserved, it can be seen to be larger than the medial condyle.

As mentioned for the humerus, the form and proportions of the femur are similarto those ofP. weddellii from Ecuador (Hoffstetter, 1952) and P. paradoxa (Cabrera,1935). Its estimated maximum transverse diameter is slightly less than the minimumvalue for P. weddellii (Table 2). The material ofP. majorof Winge (1906) has provento be that of a juvenile individual (Hoffstetter, 1952). As evidenced by the dimensionsof a femur ofP. majorfrom Brazil (Gurin & Faure, 1999), the distal epiphysis does nothave the same proportions (Fig. 3).


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Table 1 - Comparative dimensions of the humerus ofPalaeolama sp. (MUSM 51).TL total length; PTD proximal transversal diameter; DTD distal transversal

diameter; DATD distal articular transversal diameter; MTDD maximumtransversal diameter of the diaphysis (Dimensions in mm).

2. 2. Discussion

Abundant material assigned to Palaeolama and Auchenia from Miramar,Argentina, is referred to the Chapadmalalianby Cabrera (1935) and Kraglievich (1946).Nevertheless, Marshall et al. (1983) did not include the family Camelidae amongmammals of Holarctic origin assigned to the Chapadmalalian of Argentina. Later,Marshall etal. (1984) characterized the Uquian (lower Pleistocene) by the presence ofCamelidae, among other families. The occurrence of Palaeolama (sensu Gurin &Faure, 1999) in upper Pliocene-lower Pleistocene deposits of coastal southern Peru

** The specimens may belong to either P. (Palaeolama) mirifica or P. (Hemiauchenia) seymourensis.

TL-var PDT-var DTD-var DATD-var MTDD-var

Palaeolama sp. MUSM 51 - - 63,2 56 30,4

P. (Palaeolama) Hoffstetter, 1956 302-316 81-92 65,2-79 54-64,5 -weddellii Gurin & Faure, 1999 278 83,5 68,5 - 32,5

P. (Hemiauchenia) Cabrera, 1935 279-297 76-81 57-60 -paradoxa Gurin & Faure, 1999 270 74 61,5 - 30

P. (Hemiauchenia) Winge, 1906 261 79 - 47-54,5 -major

P. (Hemiauchenia) Gurin & Faure, 1999 325-348 79,5-93,570,5-82,5 - 36,5-43niedae

UF 179748 329 ~81 67 61 37

P. (Hemiauchenia) UF 45478 326 83 66 61 36

macrocephala UF179744 - - 62,5 56 33

Inglis, Florida UF 45480 - - 67 60 32

UF 176912 - - 66 60 35

UF 65323 - - 68 60 34

UF 66487 307 - 65 61 36

Palaeolama sp.** UF 66490 304 81 71 60 36

Leisey Shell Pits, UF 80400 - - 65 55 31Florida UF 80493 - - 67 57 30

UF 83661 301 84 64 61 33

UF 85018 306 83 67 61 36


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Table 2 - Comparative dimensions of the femur ofPalaeolama sp. (MUSM 51). ML medial length; LL lateral length; DTD distal transversal diameter; PTD proximaltransversal diameter; DAPD distal antero-posterior diameter (Dimensions in mm).

confirms the presence of camelids, particularly ofPalaeolama, in the Uquian, as wasalso noted by Mones (1988) based on evidence of associated fauna from Uruguay.Owing to these records, the Uquian Age (characterized by various first appearances ofmammals in Argentina) can be extended, at least with respect to the Camelidae, to Peruand Uruguay. The Peruvian material has an age similar to that of the oldest occurrenceofP. (Palaeolama) in North America, referred to the Irvingtonian of Leisey, Florida(Webb & Stehli, 1995). The existence ofP. (Hemiauchenia) is pushed back to theHemphillian (Webb & Stehli, 1995).

P. (Palaeolama) is distinguished from P. (Hemiauchenia) principally by theproportion and size of its limb bones (Webb, 1974; Webb & Stehli, 1985; Gurin &Faure, 1999). Nevertheless, a morphological analysis of the fossil material does notreveal a distinction within the genus. All Palaeolama species show important

* These measurements correspond to the maximum length of the bone.** The specimens may belong to either P. (Palaeolama) mirifica or P. (Hemiauchenia) seymourensis.

ML-var LL-var PTD-var DTD-varDAPD-var

Palaeolama sp. MUSM 51 - - - ~80 84P. (Palaeolama) Hoffstetter, 1956 369-400,5 358-394 94,5-113 83-92,5 88,5

weddellii

P. (Hemiauchenia) Cabrera, 1935 ~370 - 90 79 -paradoxa

P. (Hemiauchenia) Winge, 1906 357 - 86-95 79 -major Gurin & Faure, 1999 384* - - 89 88

P. (Hemiauchenia) Gurin & Faure, 1999 443* - 101-107 85-92 89,5-94niedae

P. (Hemiauchenia) UF 179749 - - - 93 98macrocephala UF 179750 - - - 83 89Inglis, Florida UF 45279 - - - 83 87

UF 66500 377 ~375 100 82 87UF 66602 - - - 82 83

Palaeolama sp.** UF 64331 - - - ~81 90Leisey Shell Pits, UF 66601 373 369 101 85 89

Florida UF 66603 - - - ~86 95UF 66604 - - - 82 86

UF 68884 - - - 86 87


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Fig. 3 -Palaeolama sp., MUSM 51.A-B. Right humerus:A, anterior view;B, posterior view. C. Left femur, anterior view (Scale 5 cm).

intraespecific variation in size and also are indistinguishable in humerus and femurproportions (Table 1 and 2, Figs. 4 and 5). In localities like Leisey Shell Pits, Florida,where P. (Palaeolama) and P. (Hemiauchenia) are present, complete humerus andfemur could not be assigned to infra-generic level. However, a morphological analysisof humerus and femur remains from Peru indicate that they are comparable in proportionand size with southamerican P. (Hemiauchenia) paradoxa and the smallest forms ofP. (Palaeolama) weddellii. Additionally, this can be extended to the smallest specimensof northamerican P. (Palaeolama) mirifica, P. (Hemiauchenia) seymourensis andP. (Hemiauchenia) macrocephala. Considering that the peruvian material might befrom a young adult (see below), the measurements could be slightly less than those fora completely mature individual (Figs. 4 and Fig. 5).

Other points of distinction between P. (Palaeolama) and P. (Hemiauchenia) arehabitat and diet. While P. (Palaeolama) was a browser with robust metapods that livedin montagne regions, P. (Hemiauchenia) had a mixed diet (browsing and grazing) andgracile metapods suitable to living on the open plains (Webb, 1974; Webb & Stehli,


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Fig. 4 - Scatter diagram in milimeters of MTDD (maximum transversal diameter ofthe diaphysis) versus DATD (distal articular transversal diameter) in humerus of

MUSM 51 and species ofPalaeolama from South and North America. InP.weddellii andP.paradoxa the dimension is an average.

Fig. 5 - Scatter diagram in milimeters of DAPD (distal antero-posterior diameter)versus DTD (distal transversal diameter) in femur of MUSM 51 and species of

Palaeolama from South and North America. InP. weddellii the dimension is an average.

* The specimens may belong to either P. (Palaeolama) mirifica or P. (Hemiauchenia) seymourensis;Loc. Leisey Shell Pits, Florida.

* The specimens may belong to either P. (Palaeolama) mirifica or P. (Hemiauchenia) seymourensis;Loc. Leisey Shell Pits, Florida.

82

84

86

88

90

92

94

96

98

100

75 80 85 90 95

DTD (mm)

GMUSM 51

P. weddellii

x P. niedae

P. major

P. macrocephala

I Palaeolama sp.*

DADP(mm)

DTD (mm)

25

27

29

31

33

35

37

39

54 56 58 60 62

DATD (mm)

GMUSM 51

P. weddellii

LP.paradoxa

P. macrocephala

IPalaeolama sp.*

MT

DD(mm)

DATP (mm)


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1995). The fossil record of both subgenera, the rugged relief, and the desert characterof the Peruvian coast since the middle Miocene (Sbrier etal., 1984; Alpers & Brimhall,1988) suggest that the Lamini from Atiquipa was a P. (Palaeolama)-like camelid.

MUSM 51 (portions of femur and humerus) was found associated in the samesection of the outcrop. Preceding statement and their relative size suggest that bothbelong to a single individual. However, the bone fusion secuence does not correspond.

In modern camelids, the fusion of the epiphyses of the distal epicondyles of the humerustakes place between an age of 12 and 18 months (Wheeler, 1999). The epiphyses are notcompletely fused in sample MUSM 51, which suggests it belongs to a young individual.The distal epiphysis of the femur of MUSM 51, however, is completely fused,contradicting the aforementioned suggestion. In modern camelids, the fusion ofepiphyses in the femur happens at an age of 42 to 44 months, when the animal is a youngadult (Wheeler, 1999). According to Wheeler (personal communication, 2000),exceptional cases do occur in modern camelids in which the fusion of the epiphyses ofthe distal epicondyles in the humerus never occurs due to certain environmentalconditions. In light of these comparisons, we consider MUSM 51 could belong to anindividual older than a young adult.

The nearshore marine sediments which yielded MUSM 51 were deposited nearthe mouth of a canyon and a mountainous escarpment. We propose that the bones couldhave been transported by a river from the mountains to the coast. The fragmentary natureof the bones is consistent with high-energy transport. Thus, Palaeolama sp. might haveinhabited both the Pacific coast and Andean mountains, as presently does theguanaco, Lama guanicoe (MULLER, 1776) according to Wheeler (1999). Ochsenius(1995) contends that the Lamini did not truly exploit Andean ecosystems prior to theHolocene. The occurrence of MUSM 51 in Plio-Pleistocene strata, as well as recentdiscoveries ofLama guanicoe in upper Pleistocene levels from the Peruvian coast (Salas& Stucchi, 2002) challenges that claim. We propose that the rarity of Pleistocenecamelid material in the region is due to a scarcity of outcrop, rather than a failure ofcamelids to exploit pre-Holocene Andean environments.

Acknowledgments.

The authors wish to thank Christian de Muizon (Musum National dHistoire Naturelle,Paris) for his critique revision of the manuscript and valuable suggestions; Jane Wheeler(Universidad Nacional Mayor de San Marcos, Lima) for her comments about the ontogeny ofmodern camelids; Ren Marocco (Institut de recherche pour le dveloppement, Lima) andFranois Pujos (Instituto Francs de Estudios Andinos, Lima) for their helpful insights; Bruce

MacFadden and Richard Hulbert of Florida Museum of Natural History for providing access tothe vertebrate paleontology collection. We would like to thank A. Cisneros, J. Tejada and D.Omura. We also thank Jean Vacher (Director - Instituto Francs de Estudios Andinos, Lima) andNiels Valencia (Director - Museo de Historia Natural - UNMSM, Lima) for their continuedsupport of this research.


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Institut Franais d'tudes Andines - Museo de Arte de Lima

Pedidos: IFEA, Casilla 18-1217, Lima 18 - Per, Tel. 447 60 70 -

Fax: 445 76 50 - E-mail: [email protected]

Web: http://www.ifeanet.org

salas, stucchi y devries 2003 - palaeolama

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