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Home , Alamitos Formation, Alcidedorbignya, Baurubatrachus, Casamayoran, El Molino Formation, Gasteroclupea

TIIE MESOZQIC AND OF AND THEIR STB2ATIGRAPHIC CONTEXT : A REVIEJN

MIREILLE GAYET", LARRY G. MARSHALL** & THIERRY SEMPERE*** *URA 11 du CNRS, Centre des Sciences de la Terre, Lyon I,2743 boulevard du 11 novernbre. 69622 Villeurbanne, Fraice ** Itistitute of IIunian Origins, 2453 Ridge Road, Berkeley, California 94709, USA ***Convenio YPFB-Orstom (UR IH), Sanh Cruz de la Sierra, Bolivia. Present addiess: Centre de GCologie GCnérale et Minière, Ecole des Mines, 35 rue Saint-HonorC,77305 Fontninebleau, France

~ INTRODUCTION Brazil, eastern Paraguay, northern Uruguay aiid northeastern Argentinn. In contrast, the latest Jurcissic-I'nlcocenc clcpositioiial arca Dolivia lias a rich nnd diverse record of verkbratcs for the was mainly restricted to tile Andean doniairi aiid bcloiigctl to thc al., Mcso7.oic aiid early Ceno7,oic. The majority of he 56 knowii fossil back-atc basin of the Pacific margin (Sctnpcre et 1988), localities for this time interval occur in the soutlierri Altiplano and in although soine late -Paleocene uiiits are also known in thc soutlicrii piut d the Cordillera 01ieiitnl (Figs. 1-4). parts of the iioitlicrn and cciifrril Suhnndcnii bclt (Figs. 1, 2). In Illis pnpcr we review thc iniddlc llirougli 1)nlcocciic The "Coiido cvcnt" thnt scpnriilcs tlicsc two intcrvnls ciiii be fossil vertebrote record of Bolivia in its strntigrnphic and iiiterpretcd as tlic capturc of tlic Bolivian Andean doinaiil, which palcociiviroririicntal context aiid attempt to correlate tliese rocks aid until Uien liad belonged to die crotoriic area, by tlie Pacific back-wc. faunas with those in, adjaccrit Argeiitina, Brazil, arid Pcru. As This event, acconipanied by iiiiinerous synscdimentary tectonic documented below, tlie highlights of the Bolivian vertcbrale rccord manifestations (Sempere, ¡ri press), correlates with the include: the taxonomically ricliest aiid best studied faunas of late Kirnnieridgian Araucaii event in wesicentral aiid age al., Cretaceous (from tlie marine anti continental El Molino adjacent Chile (Seinpere et 1988). Formation) aiid the spectacularly rich and only well known land maminal fauna of edy, but not earliest. Paleocene age (from the A. WfIDDLE TRIASSIC TO MIDDI,& Santa Luch Forination al Tiupninpa) ¡ti all of South Atiierica. We (Serere supersequetice) show that many or tlie taxa (especially selachians) are of clironostratigrapliic value within the Andean basin. The The rich rccord of Bolivia was traditionally thought problem of tlie K/?; bouiidary in the Andean basin of Bolivia is to lack Jurassic deposits. However, recent studies have shown that critically evaluated and based on this siudy is believed to occur certain stratigraphic units from the Subaiidean belt and from the within the upper part of tIie middle mcinber of the EI Molino Andean domain should be corTelalcd and assigned a Formation. and Jurassic age (Fig. 1) (Oller & Sempre, 1990; Senipere, 1990). The following abbreviations are used: km, kilometers; 111, nieters; The Serere supersequcnce consists of these units, and includes the Mil, nicgnniitiuin or tiiillioiis of ngo, n point in hic; Myr, Eiitrc IUos Ilnsnlt, Ipngiirv..d. Tnpecun, Cnskll6ii, lclion nnil Y nntirtri inillioiis of ycrus, a durntioii of tiine. foriiintions, as well ns lhe pnrtly cquivnlciit ßeu, Tiquina, Seynri nnd Ravelo fonnations (Fig. 1). The Tacurú Group of the central arid southern Subandcau belt is distinct froin the Scicrc supersequence bccnusc it includes tlic Intcst Crctaccous Ciijoncs Fortuntion but not Tlic "" s1r:itigr:ipIiy of Nolivia cornprises two distinct tlic Ipagunzú Forination (Scinpcrc, 1990). In gcnctal, scdiincntary tirne intervals (Sciiipcrc, 1990): tlie lirst spis tlie middle Triassic - continuity charactcl izcs tlic Sererc superscquciice (Oller & Senipere, most of Jurassic, and the secoiid latest Jurassic-l'aleoceiie (Fig. 1). 1990). As will be docuinented below, its deposits to date are sparsely The fluvio-eolian middle Triassic-Jurassic scdinietitary rocks wcre rossiliferous. dcpositcd in wh:it is now tlic castcrn liolf of tlie Andean doinairi, the The Cnstcll6ii Fortiiatioti is probably of -cnrly Jurassic age 2% Subandcari belt and a Iargc p:ut of tlic noliviali 1owl;iiids (Ollcr & (Scinpcrc & Ollcr, 1987; Ollcr Sernpcrc, 1990), althougli it Seinpere, 1990); their litliologies arid depositional enviroiitneiits lias been tentatively assigned io the early Crelaceous based on four show similarities with coeval rocks in tlie Pnranri basin of southeni new of ostracods (Pinto & Sanguiilctti, 1987). It is correlated

Y 11: SUAREZ-SORUCO, JI.(Ed.) FUSILISS FACIES DE DO1,IVIA - VOL. I VEICTEURADOS

~. - ...... 303

I v with the PirainboiQ Formation of Brazil and tlie lower part of accoinpanied by synscdinientnry tcnsioniil tectonic nnd mngiiiolic Tacuarembd Forination of Uruguay (Seinpere & Oller, 1987). rnnnifestations (Sempere et al., 1988; Seinpere, is press) and is The lower boundary of the Serere superscquemce is a diachronous termed the "Vilcapujio event" in Bolivia (Ctihvcz, 1987) or k3 in tile unconformity except in the southern Subandean belt where,tlie oldest central Aniles (Jaillard & Sempere, unpublished). Thus, the and elsewhere unknown Ipaguazú Formation overlies the late Miraflores Formation stands as a specific "hinge" unit in the - Vitiacua Formation with only a slight, non- Bolivian latest Jurassic-Paleocene stratigraphic sequence. erosional, lithologic discontinuity (Sempere et al., 1990, 1992). In Mainly red mudstones with minor sandstones and evaporites other areas of the basin, younger units of the Serere supersequence (gypsum, anhydrite, halite) were deposited between discontinuities unconformably overlie Permian or older strata, reflecting onlapping k3 and k4 in fluvial and lacustrine environments [Aroifilla .deposition on an altered, eroded and/or deformed substratum. The Formation and most of Chaunaca Formation (Sempere, in press); the upper boundary of the Serere supersequence is the surface that marks stratigraphically equivalent Torotoro Formation consists mainly of the "Condo event" (see above). This unconformity shows different red sandstones of fluvial origin]. Two greenish levels, which consist characteristics which relate to its location and position in the Iahst of very fine sandy, green to black mark, black to grey laminated Jurassic-Paleocene basin. and yellow dolomites, occur in the Chaunaca Fonnation. In suinmary, tlie middle Triassic-Jurassic geologic evolution of The foriner is more conspicuous, traditionally marks the base of the Bolivia involves: 1) an initial rifting process with deposition of red unit, and is krmed the " " of the Chaunaca Formation. beds in narrow troughs, locally with gypsum and halite (middle to This 30 m-thick sub-unit commonly yields abundant bivalves Iate Triassic); 2) termination of rifting and onlapping fluvio-eolian (Mytilidae, Brochidontes sp.) which have been interpreted to scdinicntntion (late Triassic to earliest Jurnssic'?); and 3) extensive indicate an intertidal cnvironnient (Briutisn ct al., 19GG). However, II development (early and '?) (Oller & Sempre, restricted mnrine origin for the "basal limcstonc" is now questioncd 1990). (G.Camoin, personal coinunication). Identical Bracltidoort/es sp. are recorded from a limestonc level in the Loinas Negras Formntion of U II. I,h'l'lSS'l' J It ASS IC 'IO l'A I,EO CICN IC nortlicrn Chile (Mnrinovic & Litlisen, 1984). The "bnsitl linlcstonc" (I'ucu supersequence) of the Chaunaca Formation yielded a palynological assemblage of to earliest age (Pérez, 1987), and may be Classic works on the Bolivian "Cretaceous" Puca Group are by partly equivalent to the Snntonian age shallow-marine limestones Lohmann & Branisa (1962). Russo & Rodrigo (1965), Kriz & known from the Querque and Omoye formations of southern Clierroni (1966). Lohmann (1970), Reyes (1972) and Cherroni (Jaillard & Sempere, 1989). The Aroifilla and Chaunaca formations (1977). are equivalent, respectively, to the lower and middle Vilquechico The "Cretaceous" sequence of Bolivia and Peru is Formation of southeast Peru (Jaillard et al., in press) (Fig. 1). The currently being re-evaluated and synthesized (Jaillard & Seinpere, upper greenish level of tlie Chaunaca Formation is equivalent to the 1989; Seinpere, in press). The Puca supersequence (Scmpere, 1990) uppennost green level of the middle Vilquechico (Jailllard et al., in is bound below by the kO discontinuity (;.e, the press; Sempere, in press), which has yielded the selachian Araucan-"Condo" event at about 144 Ma; Cowie & Bassett, 1989). slronieri. Thus, its age is not older than níiddle and above by tlie k5 discontinuity which records the functional onset Campanian (Jaillard el al., in press). of western Bolivia as a continental external of the The EI Molino, Santa Lucía and Impora formations occur between paleo- (Sempere et al., 1989). The age of W is close to the discontinuities k4 and k5 in Bolivia (Fig. 1). The EI Molino Paleocene- boundary (53 Ma; Cowie & Bassett, 1989) and Fonnation is equivalent to the Lecho, Yacoraite, Tunal and Olmedo could be as young as 50-51 Ma (Senipere, 1990; Marshall & formations of northwestcrn Argentina (Fig. 1); to the Areniscas de Senipere, 1991). Azúcar, Vivinn and Cnchiyacu fornialions of Subnndenn Peru In this pnper we, divide the Pucn supersequence into three (Sempere et al., 1987: Jnillard & Sempcrc, 1989); to thc upper megasequences. The Puca A megasequence includes the partly Vilquechico Formation of the Peruvian Altiplano (Jaillard et al., in equivalent Condo, Kosmina, La Puerta S.S., Sucre and Tarapaya press) (Fig. 1); and to the Estratos de Quebrada Blanca de Poquis, forinations (Fig. l), nnd consisLs of nzoic non-marine dcposits; the niid possibly parts of thc Loinns Negras. Pnjonalcs and Toncl niiuinc Miraflores Fdrntntioii constitillcs the I'ucn B nicgnscquc~icc; formations in nortlicrn Cliile (Gordcwcg 8c l

394 FOSILES Y FACIES DE BOLIVIA - VOL. I- VERTEBRADOS

ANDEAN DOMAIN SUBANDEAN BELT NORTHWEST SOUTHEAST AGE ARGENTINA PERU (Ma) (Haq el a1.,1987)

54 - - - I OW - 60 - 98 Cajones i~ 2066.5 - 70 - Maaslriclilian

- 80 -

u) Conlaclan - 90 - 3 Turonian O w Cenoinnrii:rii O Q - 100 - LuI- n O -1 10- Aptlan

-120-

-130- 131

-140-

-150- Yantata

-160- o u) u) -170- Q U 37 -180- lchoa lchoa

-190- Liassic

- 200 -

-210- 210

220- - I! Keuper u) u) 5 -230- n I-

-240- -

Figure 1. Clironostratigrapllic chart showing -bearing formations (*) of middle Triassic-Paleocene age in Bolivia, and in adjacent Argentina and Peru.

395 -I

MIREILLE GAYET. LARRY O. MARSIIALL &TIIIIiKRY SEMl'liKI!

Rased on recognition of large-scale transgressive-regressive M. Roucliy, ,personal conimunicntion), are abundant in the iippcr sequences, the EI Moho Formation is divided into lower, middle part of Uie Santa Lucía Formatioli near Potosi. Some green maris and and upper membcrs which are of stratigraphic value (Jaillard el al., light-colored limestones occur in the Impora Formation of southcrii in press; Jaillard & Sempere, unpublished; $empere, in press). Boliv,ia. Common sedimentary facies in the El Molino Formation include: The Santa Lucía and Impora formations are, respectively, mudstones and mark (green to black or purplish, reddish); grey to stratigraphic?lly equivalent to the'Paleocene age Mealla and Maíz al., light brown limestones (micritic to bioclastic and/or oolitic); Gordo formations of northwestern Argentina (Marocco ef 1987) stromatolitic boundstones referred to as Piicaliiltus by Steinmann (Fig. 1). The Mealla Formation can be correlated with the (1923) and Ahlfeld & Branisa (1960); fine to medium-grained Tiupampian (early Paleocene) Land Age based on its sandstones, usually calcareous and light-colored; some yellow equivalence with the Sai?ta Lucía Formation (see below) and the dolomites, often with algal laminations; very rare evaporites Maiz Gordo Formation with the (late Paleocene) Land (gypsum and halite casts); and a few white altered tuffs of porcelain Manimal Age based on palynology (Volkheimer ef al., 1984) and aspect. Mudstones, mark and thin limestones largely predominate in stratigraphic position (Pascua1 et al., 1981). the basin; sandstones, dolomites and evaporites are more common along its borders; and tuff levels are thicker and more frequent in the VERTEBRATE FAUNAS AND LOCALITIES IN BOLIVIA west, i.e. in the direction of the active volcanic sources located in present-day Chile and Peru. In the basinal localities, lime turbidites A. TACURU GROUP are common and where fossiliferous the paleontological materials were dcpositcd a stibstaiitial distance from their life environinent. Alilfcld & Brnnisa (1960) rcportcd rcinniiis of lurgc , Coilcordant palcocurrciit data iiidicate that the basin deepened possibly of , from the Tacurú Group in the Serranía de toward the northwest in Bolivia (Sempere er al., 1987; Seinpere, in Mandeyapecua. southeastern ßolivia. IIIthis area the Tacurú Group press), and probably connected with a marine basin in the Peruvian includes rocks of middle Triassic and Jurassic age, naiiiely tlie Subandcnii region. Aiiiiiioiiitcs tire reported 2 in bclow the top of the Tnpccun, Cnslcll6ii aiid Ichoii forlnntiolis (Fig. 1). Cachiyacu Foriiiation in the HiinIlaga aren (7"s) of iiorthern l'cru (Vargas, 1988). In Bolivia, several groups, which reputedly U. CASI'ELLON FORMAlION indicate a marine environment (see below), are known from the lower and basal middle El Molino. Dinoflagellate cysts are present in Scales and bones of semionotiform were found in the late at least some levels of the formation in Bolivia and in stratigraphic Triassic- age CastelMn Formation on the western flank equivalents in northwestern Argentina (Industry, unpublished; J. of the Charagua anticline in tlie Quebrada de Charagua, 7 km west of Oller, personal communication). Ammonite embryos have been the town of Chbagua (Wenz, in Goñi & Hoffstetter, 1964). Nothing reported from tlie lower El Molino (Sempere ef al., 1987), but their definite can be said of these bones, although a study of the scales identification needs confirmation. (Miliolidae, with Scanning Electron Microscopy (Gayet & Meunier, 1986) dnd of Discorbidae) are present in many levels aid facies (G. Tronchetti, in their transverse sections permits reference to the family Camoin et al., 1991), but no detailed study has yet been made of Semionotidae, probably Lepidofes sp. (Gayet, 1991). their or paleoenvironrnental implications. Molluscs are reported from various localities (e.g. at El Molino; Pilsbry, 1939), C.MIRAFLORES FORMATION although the systematics of these taxa is outdated and the relevance of described species has yet to be established. Echinoids are The Miraflores Formation is a marine limestone unit which, based apparently lacking and the proportion of brackish water ostracods is on knowledge of ammonites, bivalves, gastropods and echinoids, locally high (J. F. Babinot. in Cainoin et al., 1991). was regarded as middle to late Ccnoiiinnian in age (ßranisa et al., A of pctrogrnphic study the Yacoraite Forination of northwestern 19GG; Jaillard & Scmpcrc, 1089), :itthough detailed scqucncc Argentina led Marquillas (1985) to interpret its depositional sthtigrapliy now suggcsts that the entire rock unit spans Cenoinaniaii environinent as a restricted carbonate basin. An oligohalirie and Turonian time (Jaillard & Sempere, in press). Near the locality al., lacustrine environment has also been proposed (Camoin ef of Macha. 90 kni north-northwest of I'otosí, L. Branisa found minute 1991). However, paleontological data do suggest that pharyngeal teeth which are tentatively assigned to cf. communication of the basin with an open marine realm was frequent Cyprinodontiforines or Pycnodontiforrnes (Gayet, 1991). during deposition of the Ei Molino Formation, but that the basin itself was never openly marine, at least in Bolivia and Argentina I). AROIFILLA FORMA'I'ION (Gayet et al., 1992, Seinpere, in press). For reasons discussed below, we believe that most of the El Molino Formation was The continental Aroifilla Formation overlies the marine deposited principally in a restricted marine environment which Cenomanian-Turonian age Miraflores Formation and underlies the connected with a more typically open marine realm located Santonian-late Campanian age Cliaunaca Formation; it is therefore northwest of Bolivia (see also Sempere, in press), considered and possibly early Santonian in age (Fig:l). A The overlying Santa Lucía and Impora formations were deposited of four footprints of a large quadrupedal were 4 in fluvial to lacustrine environments. No marine fossils or facies are discovered by the Brigada 10 of YPFB at Calerías, in below the known from these units which consist principally of red to purplish "basal limestone" of the Chaunaca Formation. The footprints were siliciclastic sedimentary rocks. Gypsum and "gypsifred" anhydrite (J. imprinted in soft sediment and are otherwise indeterminate.

396

I F FOSILFS Y FACIES DE BoLIm - VOL. I - VERTEBRADOS

PERU 1; ,I I

,4t 15"

100 km

.LA PAZ

17" *ChuIlDa Khasa . La Cabana- *Parotan¡ -Rio Moile pa,c~s~$~ipdEst. Blanco Rancho USANTA CRUZ ORURO. Arapampap d!ilaVi'a DE LA SIERRA Lago Uru Uru Torotoro-. Tiupampa

19'

/-

*Chaup¡-Khocha echaragua ,f I / : *Calerias Serraniat , de f *Tambo *Camargo I' Salar de Uyi ni Colorado Manieyapecua I *Chocaya ,i 21" Chaupiuno i, CHILE i Rancho Hovada-. UTARIJA I Serpa\ *Villa Pacheco .í ((PARAGUI '\ \ \ \ \ ' 69" 67" 65" 63" '\,

Figure 2. Map of Bolivia stlowine: vertebrate localities of middle Triassic-Paleocene age.

397 Y MIRIIILLEGAYEI'. URRY G. hIARSIlALLLTIUERRY SEMI'EKI!

IC. CllAUNACA IWI

398 Cerro Lama Chaqui

180 Torotor

Cruz Khasa

' Ikm '

Figure 3. Detailed map of tlie Torotoro area (for location, see Fig. 2) showing vertebrale localities in the CI Molino and'Santa Lucia formations. The numbers correspond to those in Table 1. Based on Carta Nacional, Bolivia, Torotoro Quadrangle, Hoja 6439 IV, serie €1731, 1/50,000, 1968 edition, Instituto Geogrhfico Militar, La Paz VIII-68; and San Vicente Quadrangle, Hoja 6439 I, serie fi731, ,1/50,000, 1968 edition, Instituto Geogrhfico Militar, Ln Paz 111-68. siluriforin (Ariidae, Rliineastes sp.) of very large size (more than 2 in (Pycnodontiformes; Characiformes, Serrasalmidae, Myleinae; in total length). Vertebrae of Lepisosfeus sp. found in association Siluriforines, Ariidae, Rliineastes sp.) and reptiles (turtles and with the Siluriforines are also larger (more than 3 cm-long) than crocodiles). In the vicinity have also been found several levels with those of Agua Clara (about 1cm-long). As in some levels at Agua holostean Ginglyinodi (Lepisosteidne), teleostean Clupeiforrnes CI:lra, thc presence of sclaclii:ins, pycnodonti forms, tetradontiforins (Clupeidae, Castcroclicpea Invnisai), Chnraciforines (Serrasahnidae, and "saliiioniforins" (Enchodirs sp., ?Apateodirs sp.) testifies to some Myleinae). Siluriforines (Ariidae, Rliineastes sp.), and/or reptiles conmunication to a sea (Gayet, 1991; Gayet et al, 1992). (crocodiles, turtles). Ten meters below the calcareous level just described, a restricted At Sevaruyo. south of Lake Poop6. about 120 kin west-norU!west conglomeratic unii was recently found with reinaius of Siluriformes of Potosí, are reported two levels with remains of Casteroclitpea sp. (Ariidae) and Teleostean indet. A little farher, at the crossing of the (Branisa et al., 1964). The first level apparently belongs to Uie lower Quebrada Saytu JoWiu with the road, are grey-green mudstones of EI Molino. Because of the large.(double) size of the hypocoracoids the upper EI Molino wilh remains of Clupeiformes (Casteroclitpea found in the second level relative to .CasteroclupeaS holotype, branisai) and ostracods (Gayet, personal observation). Branisa ef al. (1964) questioned the age of this level. However, At Vilcapujio (=Wila Apacheta), at about hi 140 northwest of hypocoracoids of very large size have also been found at Agua Clara Potosl on Uie Potosl-Challapata-Oruro road, is a bed probably where a transitional series exists that includes all sizes recorded at belonging to the basal middle EI Molino which yielded fishes otlier localities.

399 MIREILLE CIAYET, LARRY CI. MRSIULL& 1’1 UliKKY SBMI’EKB

2. Uyuni Region of Torotoro. occur scvcrnl scts of trackways of a sinall theropod dinosaur (Coclurosaurin?). The siinic typc of trnckway occurs ncar Uic At thc locnlitics of Jay-Jay, Calnzaya and Tomave, rcspcctively top of tlic lowcr EI Molino on thc ptl~to Ulna Jetanta, about 2 north-northwest, east and northeast of Uyuni, were found remains of km west of Torotoro. Siluriformes (Ariidae, Rlrineasres sp.; Tomave only) and From the transgressive base of the middle El Molino in the Río 3 Clupeiformcs (Gasreroclupea branisai) by the$rigada 10 of YPFß Cuchira Waykho, kin south-southeast of Torotoro, a long rostrum (specimens in CTP, Santa Cruz). At Tambo Colorado, squtheast of of a “salmohiform” (family indet. 2, gen. and sp. indet.; Gayet 1991) Uyuni, was found a selachian (Pucapristis sp.) associated with is assigned to the marine Cretaceous suborder Ichtliyotringoidei gastropods (Melania potosiensis; specimens in CTP, Santa Cruz). (semu Goody, 1969). With the exception of Calazaya which is upper El Molino, the From an undetermined member at Cruz Khasa. on the west side of stratigraphic position of fossils at the other localities have not yet the cemetary south of Torotoro, remains of Prtcapristis sp., been established. crocodiles and turtles (?Ro.xoclielys sp.) have been recovered (R. CCspedes, personal communication; specimens in Museo de Historia 3. Southern Region Natural, Cochabamba). At Tiupampa, located about 95 kin southeast of Cochabamba, two In the Río Angosto, 9 kin south-southwest of Chocaya, holostean fossil levels are known (Fig. 4). The lowest, near Hera Mokho, is (Lcpisosteidae, Lepisosteus sp.), Semionotiformes about 90 m above the base of the Cretaceous section from a (Semionotidae, nov,. gen.), teleostean Clupeiformes (Clupcidae, calcareous sandstone horizon of the middle EI Molino (see Marshall ’ Gasteroclirpea ’brunisni) and Siluri formes (Ariidae, Rltineasres sp.) et al., 1985. Fig. 4); vcrtcbrntcs nrc rcprcscntcd only by a sclochinn wcrc recovered froni,the lower EI Molino. (Dasyatidae, flasyaris sc11aejJeri) and an indctcrrninatc crocodile In Quebrada Tcxisca, 1 km west of Rancho Hoyada along the.Río (Marshall et al., 1985). There are also abundant gastropods San Juan del Oro, 29 kin north-northeast of Tojo, were recovered (Melanidne, Melania potosicnsis) which unTortunulcly nrc not useful rcninins of lioloslcnii I’yciiodontiforiiics (Pycnodontidac. indct.), for ngc rcsotulion (Ahll‘cld & Ariinisn, t960). t4iglicr in lhc scction, Ginglyinodi (Lcpisostcidae, Lepisosrens sp.), teleostean along the Rio Pucarani, is an uppcr EI Molino bone bed with Osteoglossiforrnes (Osteoglossidae, indet.), Siluriformes (Ariidae, Siluriforines (Ariidae, Rhineastes sp.), indeterminate turtles and Rhineastes sp.) and Tetraodontiformes (Eotrigonodontidae, crocodiles. Sfephunodirs niinkirs) (Gayet, 1991) from the lower EI Molino. The At (=Villa Viscarra), about 90 krn southeast of saine Pycnodontidac, Osteoglossidae, Ariidae and Cochabamba and 6 kin northwest from Tiupampa, a fossil level in Eotrigonodontidae, plus two new selachians (Dusyuris nov. sp. 2 et the lower El Molino about 100 ni above the base of the Cretaceous 3; Cappetta, 1991) and a crocodile tooth were recovered from the section (see Marshall el al., 1985, Fig. 4) has yielded some of the basal middle El Molino at the same locality. saine selachians recorded by Cappetta (1975) from tht lower El Fish remains, including Pucaprisfis brunisi, have also been found Molino at Torotoro (Prtcuprislis branisi, Isclryrliiza hartenbergeri, by J. Blanco (YPFB, Brigada 10) near Serpa about 35 km south of Pircabaris hoffsfefreri);numerous actinopterygian fishes: holostean Tupiza, and dinosaur trackways were observed by Leonardi (1981) Pycnodontiformes (Pycnodontidae, indet.), Semionotiformes near Camargo, 100 kin north of Rancho Hoyada. (Semionotidae. nov. gen.); teleostean Characifornies (Serrasalmidae, Myleinae). Siluilformes (Ariidne, Rlrinensres sp.. Andinichthyidae 4. Cochabamba Region indet.), cf. Cyprinodontiformes; Brachiopterygii (=Cladistia), Polyptcriforines (Polypteridae, Dnjerelln srrdmnericnnn; Gayet & At Torotoro, aboui 95 kin south-southeast of Cochabamba, several Meunier, 1991a, b. c); Dipnoi (Lepidosirenidne, Lepidosiren cf. NC 3). fossil lcvcls known (Fig. From thc iniddlc part of thc lowcr EI paradma; Schultzc, 199 In); n turtlc (I’odocllciiiididrlc. ?I?o.toc/wI.ys Molino sonic 3 kin soulti-soutlicnst of Torotoro, Cnppctla (1975) cf. vil~~il~t~sis;fjroiii, 199 I),und two iiidctcrniinotc ciocotlilcs dcscribcd numerous sclachians [Sclerorliyncliidae, f‘itcapristis (Marshall et al., 1985). brnisi, Iscliyrhiza Irnrtenbergeri; Dasyatidae, Dasyaris branisai, D. Two new localities were discovered during the 1989 field scnson, niolinoensis, D. schnefleri; Khoinbodontidac, Pucabah Imffstelleri Estancia ßlnnco Rancho and I’iijcha Patii, niidwoy along the Clizn- (Cappetta, 1987)J and Broin -(1991) recorded turtles Anznldo road, about 40 kin southeast of Cochabanibn. At Pajchn (Podocneinididac, ?Roxoclielyssp.). Pata (lower EI Molino) were recovered reniaiils of selachians In a higher level in the same inember, on the northeast side of (Pucubaris Iroffstetreri, Pucaprisfis brunisi), “holostean“ Torotoro, numerous dinosaur trackways were found that have been Pycnodontiformes (Pycnodontidac, Coelodrcs toncoensis; 3). g n a callcd the “pista dc danzas“ (Fig. At lcast five types of dinosnurs Lep is os tc id tic, Lep isosf e II s s p. ; II II o id li o 1os te n s c ICs ) , are represented, including sauropods, tlieropods and ornithopods Clupeifornies (Gasreroclrrpea branisai), Siluriformes (Ariidae, (Branisa, 1968; Leonardi, 1981); one trackway named Rhineasfes sp.; Andinichthyidae, indet.), cf. Cyprinodontifoniies, Ligabueiclrniunr bolivianrtnr represents an ankylosaur or ceratopsian numerous bones belonging to as yet unidentified small teleosteans, (Leonardi, 1984). teeth (Lcpidosirenidae, Lepidosiren cf. L. paradoxo), Near the base of the lower EI Molino at Unia Jalanta, close to Uic Anura, one vertebra which could be an atypical Urodela, cave entrance, about 5 kin west-northwest of Torotoro, occur some two other vertebrae which have the morphology of a Urodeln but are small tridactyle (Coelurosnuria?) footprints and isolated bones of procoelic (the vertebrae of Urodela are always ophisto- or turtles (?Roxuclre[ys’sp.). At Cerro Llania Chaqui, about 4.3 km east amphicoelic), probably a , a tooth of a sninll tlieropod

400 Figure 4. Detailed map of tlie Tiupampa area (for location, see Fig. 2) showing localities (1-9)from the principal vertebrate level of Ille Santa Lucía Formation, and localities in lhe middle (A) and upper (B) El Molino Formation. Tiupampa is the local name used by Quechua indiuns for tlie urea shown on lhe mup. Bused on Cnrln Nacionul, Ihliviu, Sun Vicenle Quudrungle (see cuption lo Fig. 3).

(; Marshall,, 1989b), indeterminate turtles. various soutlieast of .Cochabamba. These are apparently the sanie foolprints crocodiles nnd indeterminate trackways. At Blanco Rancho (upper El described by Mnrshall & Molina (1990) the lower EI Molino Molino) were found Clupeirormes (Gusleroclupeu brunisui). Formntion 4 km south-soulhenst of Satitivafiez rind 10 kin east of Siluriformes (Ariidae, RltineasfPs sp.), cf. Cyprinodontilormes, Parotani: The best preserved footprints appear referable to an turtles and numerous ostracods. ornithopod, probably of the family Hadrosauridae, while Trackways of small bipedal dinosaurs (Coelurosauria?) are others may represent a small theropod, possibly a recorded from Abpampa at Cerrito de Llamachaqui, Departrnent of coelurosaur. Polosi (Leonardil 1981). From Sayari at km 87 on the road from Cochabamba to Oruro, R. Fragments of large bones, possibly of dinosaurs, and "dientes de CCspedes recovered remains of a selachian (Pucupristis brrurisi), a tipo cónico simple" were observed in a level with gastropods in tlie siluriform (Ariidae, Rhineusles sp.) and an indeterminate turtle from El Molino Formation near La Cabaña, north of Parotani and west of the lower EI Molino (specimens in the Museo de Historia Natural, the Río Rocha about 35 km southwest of Cochabamba (Ahlfeld & Cochabamba). Branisa, 1960). Chullpa Khasa, 6 km southwest of , has yielded a fish Leonardi (1981: 935) reports a series of six poorly preserved (Osteoglossiformes, Osteoglossidae, Phareodontinae) and footprints of a bipedal dinosaur in the Santa Lucía Fonnation along remains of reptiles (turtles, crocodiles) from an undetermined EI the Cochabamba-La Paz road near Parotani in the Suticollo Syncline, Molino member.

401 / I

Figure 5. Stratigrapliic sections of late Cretaceous-Paleocene rock units at Tiupampa, Pajclia Pata (+ Estancia Ulanco Rancho), Torotoro (+ Caranota),' Potosl (road to La Falca and Cayara) and, in part, Agua Clara, showing location of main fossiliferous levels. Undecompacted thicknesses. The base of the unconformable Cayara Formation, is held horizontal. Fm, Formation; I, lower; m, middle; u, upper; Pz, : 1, erosional surface; 2, conglomerates; 3, conglomeratic sandstones; 4, sandstones; 5, fining and/or thinning-upward; 6, siltstones and mudstones; 7, carbonates (mostly limestones); 8, coarsening and/or thickening-upward; 9, gypsum; 10, tuffite bed; 11, stromatolitic level; 12, dissolution breccia; 13, channels; 14, cross-bedding; 15, oiiids; 16, calcareous cement; 17, rooting; 18, coalescent rhizolith-rich levels; 19, bioturbation; 20, red color predominant; 21, dinosaur trackways; 22, vertebrate-bearing levels. All sections measured by T. Sempere, except parts of Torotoro- Caranota and Potosí (respectively by tlie Brigadas 10 and 9 of YPFU) and of Pajclia Pata and Agua Clara (by G. Camoin and J. M. Rouchy); all were checked by the authors.

402 FOSILES Y FACIES DE BOU- - VOL. I - VERIBBRMOS

1 and 2, IfofJsfertericltrltyspucai and Incaiclirhys srrarezi), Perciformes (Centropoinidnc), Polyptcriforines (I’olyptcridne, A vertebrate fauna descrilxd originally as of possible Palcocelie Dajelella sudanicrictmn) and Dipiioi (Ccratodoiitidnc, Ccralodrts sp. age was reported forti Uie northern flank of tlie Huarachani syncline, and cerntodont n. g., n. sp.; Lepidosirenidne, Lepidosiren cf. just north of tlie pueblo Huarachani along the Bolivia-Peru border paradoxa). north of Lake Titicaca (Argollo el al., 1987). The fauna includes a b) (Rage, 1986, 1991a): Anura (Leptodactylidae); fish (Osteoglossoitiorplia, Osteoglossidae, Phareodontinae; Gayet, (family indet.). 1987b, 1991) and a crocodile vertebra (Mesosuchia, , c) Turtles (Broin, 1988, 1991): Podocnemididae, Roxockelys cf. gen. arid sp. indet.). However, these fossils come from the middle vilavilensis. nieinber of the El Molino Formation (called Ococoya Formation in d) Lizards (Rage, 1991b): Lacertilia, ? and family indet. this area; Fig. 1), and are therefore of late Cretaceous age (Martinez, e) (Rage, 1991b): Aniliidae n. g.; Boidae, two species 1980: 183; Gayet, 1987b; Mourier el al., 1986, 1988: 171; see indet.; ?Maqtsoiidae; Tropidopliiidae. below). f) Crocodiles (Buffetaut, 1991; Buffetaut & Marshall, 1991): Mesosuchia, , Sebecris querejazus; Dyrosauridae, 6. Northern Subaridean belt Sokorosucltus aff. iwiwilsoni. . g) (Muizon el al., 1983; Muizon, el al., 1984a, b; About GO-70 in above the base of the late Cretaceous age EslaMn Marshall el al., 1983a, 1985. 1989; Muizon & Marshall, 1985, Formation (=Flora Formation s.1. of Perry, 1963), from levels that 198711, b, c. d, 1988, 1991. in press; Marshall, 1992: Marshall & cnii bc correlated to the lower El Molino, nloiig the Rlo Flora, 315 Muizon, 1988, 1992): Dcllotlicroidn (family indel,, Ja~kh~d~/pl~~~ kin north-northwest of La Paz, were collected rcmains of ntinufus), Peradectia (I’eradcctidae, Peradectinae, Pcradcctes Pucaprislis sp. (=anclioprisris), Gaslerocllrpea brunisai and aus~rin~n~;Caroloainegliiiiiinne, Roberllioffs~elleria charophytes (including Peckichara compressa) (Perry, 1963; DBvila tiariotialgeograpliica), (, Kliasin & Ponce de Lc611, 1971). In the saine area, Gasferoclupea branisai cordi llerensis), Did elpli i ni or p Ii ia (Did e 1pli id oc, Did el pli i ri ne, was recovered froin the overlying Flora Forination S.S., an quivalent Piicadelpltys andinris, Incudelpliys anriqrrus, Mizquedelphys of the upper El Molino (Vernet & Bokllo, 1975). pilpinensis; Eobrasiliinae, Tirilordia floresi), a (H a t h 1i acy n id a e, A Ilqo k ir 11 s us I r a 1is), Pa u citub er c u 1at a 7. Central Subandeen belt (Kollpaniidae. Kollpnnia fiupanipina), order and family indet. (Andinodelpliys cochabanibensis), (?, cf. Sanjinés-Saucedo (1982) and Mpez (1983) reported remains of Ciniofesles sp.), (Pantolambdidae, Alcidedqrbignya branisai from the Cajones Formation (partly or inopinafa), Condylarthra (, Mioclaeninae, totally equivalent to the EI Molino Formation; Fig. 1) in the Moile Tiuclaenus niinulus, Molinodus suarezi, Pucanodus gagnieri, Syncline along tlie Río Moile about 95 kin west-northwest of Santa Andinodus boliviensis), and (cf. Henricosbomiidae or Cruz. The level in which the fossils were foulid probably represents , gen, et sp. indet.). the upper El Molino. Criadero de Loro, a new locality about 1.2 km soutii-southeast of the quarry at Tiupampa and from the sanie horizon (Fig. 4, locality 7). G. SANTA LUCIA FORMATION was discovered in 1989. The taxa seem to be the scarne as those from the quarry and adjacent localities (Fig. 4, localities 1-6, 9) and The richest vertebrate fauna, in ternis of both number and quality include, Siluriformes (Ariidae. Rhineasles sp.), Perciformes of specimens, is from Tiupampa about 95 km southeast of (Centropornidae), Polypteriformes (Polypteridae, Dajetella Cochabamba (Figs. 2, 4). Two fossil levels at Tiupampa belong to sudamericana), nu iiicrous 1a rgc lu iig fis li t ce th (includi ti g the basal iniddlc and upper EI Molino (sec nbovc). A third fossil Ceratodoiitidiic and Lepidosircnidac), snnkcs (ßoidac sp. indct.), level is located about 130 in above the base of the Cretaceous section turtles (?Roxochelys vilavilensis), crocodiles and a pantodont (see Marshall el al., 1985, Fig. 4). Sempere & Marshall (in press) ( inopinafa). dcinonstratc that this innin rossil level, which represents the type At Viln Vilb, íÌom tin uppx lcvcl lociilcd nbout LGO ni above the fauna of the Tiupainpian Land Mammal Age (Marshall. 19890) base of tlie Cretaceous section (see Marshall el al., 1985, Fig. 4), belongs to the Santa Lucía Forniation.(see Figure 5). The following have come a turtle (Podocnemididae, ?Roxochelys vilavilensis; vertebrate taxa are recorded: Broin, 1971) and a crocodile (, Sebecidae, a) Fishes (Gayet. 1988b. 1990, 1991; Gayet & Meunier, 1991a. querejazus; Buffetnut & Marshall, 1991). 199 1b, 1992; Schultze, 199 1a): teleostean Clupeiforrnes (Clupeidae. At Uie two newly discovered localities of Estancia Blanco Rancho Gasteroclupeinae, Gasreroclupea branisai), Osteoglossiformes and Pajcha Pata, south of (Fig. 2), are bone beds of the lower (Osteoglossidae, Phareodontinae, Pliareodusichfltys favernei; and/or upper members of the EI Molino described above, which are Osteoglossinae. incerfae sedis) (Hiodontidae sensu Muizon el al., overlain by red sandstones of the Santa Lucia Formation similar Lo 1983 is a confusion with Osteoglossinae; Gayet, 1991), those at Tiupampa (Montaíío, 1968). A brief survey of Blanco Characiformes,(Characidae, Tetragonopterinae, indet., cf. Rancho resulted in the recovery of numerous teleostean fishes Rhoadsiinae; Serrasalmidae, Serrasalminae, Myleinae; Erythrinidae, (Osteoglossiformes, Osteoglossidae; Siluriformes, Ariidae, Hoplias nov. sp.). Siluriformes (Ariidae, Rhineasles sp.; R li inea st es s p., A n d i ni c li thy id ae, i nd e t erm i na te te leo s t s) , AndinichUiyidae, Aridiniclllliys bolivianensis; families incerrae sedis amphibians, crocodiles (Dyrosauridae, complete ), turtles

403 MIREILLE CIAYET. LARRY O. MARSIMLL & I’IUBKRY,SEMI’EKI!

TABLE 1. VERTEBRATE FOSSIL LOCALITIES OF MESOZOIC AND PALEOCENE AGE 1N BOLIVIA ARRANGED BY STRATIGRAPHIC OCCURRENCE.

TACURU GROUP UPPER MEMBER 1. Serranía de Mandeyapecua 31. Calazaya CASTELLON FORMATION 32. Cayara (Cerro Muyurina) 2. Quebrada de Charagua 33.* La Paka (Doliciwciranipsa mininia level) 34, Pajcha Pata MIRAFLORES FORMA’I’ION 35. Estancia.Blanco Rancho 3. Macha 36. RíoFlora 37. Río Moile AROIFILLA FORMATION 38. Tiupampa (Río Pucnrani) 4. Calerlas (dinosaur footprints) INDEIXI¿MINA’I‘E MEMUER CHAUNACA FORMATION 5. Aguaclara ’ 39. Arapampa (dinosaur footprints) 6. La Paka 40. Camargo (diiiosiiur footprints) 41, Chullpn Khasa EL MOLINO FORMATION 42. Jay-Jay 43. Serpa LOWER MEMBER 44. Tambo Colorado 45. Toinave 7. Aguaclara 46. Torotoro (Cruz Khasa) 8. Cayara (Cerro Muyurina) 9. Chocaya (Río Angosto) SANTA LUCIA FORMATION 10. Hotel Cordillera near Wila Khasa; Quebrada Saytu Jokhu with Gast eroclupea 47. Chaupi Khocha 11. La Palca (black shale level with Gasteroclupea, and below 48. Maragua ) 49. Pajcha Pata 50. 12. Pajcha Pata Estancia Blanco Rancho 13. Kanclio Hoyada (Quebrada Texisca) 5 1. Tiupampa (quarry and associated localilies) 14. Sayari (km 87) 52. Tiupampa (Criadero de Loro) 15. Scvaruyo 53. Torotoro 16. Torobro (Ceiro Llama Chaqui. dinosaur footprink) 54. Vila Vila 17. Torotoro (Uma Jalantn cave, dinosaur Foolprints) 18. Torotoro (Uma Jalantn road, dinosaur footprints) IMI’OKA FORMATION 19. Torotoro (“pista de danzas“. dinosaur footprints) 20. Torotoro (selachian level) 55. Cliaupiuno 21. Vila Vila (Piccaprisky icvel) 56. Villa I’achcco 22. Wila Kliasa (km 100) 23. Río Flora I 24. Parotani (=Santivaííez) (dinosaur footprints)

MIDDLE MEMBER

25. Hotel Cordillera (near Wila Khasa) 26. Huaracliani 27. Rancho Hoyada (Quebrada Texisca) 28. Tiupampa (Hera Mokho) 29. Torotoro (Río Cuchira Waykho) 30. Vilcapujio mila Apacheta)

404 TAULE 2. SYSTEMATIC LIST OF VERTEBRATES FROM THE MESOZOIC AND I'ALEOCISNE OP UOLIVlh WITH AUTHORSIIII'.

CLASS SELACHU Order Suborder Sclerorhynchoidei Family Sclerorhynchidae Pucaprisris branisi Schaeffer, 1963 Iscltyrltiza Iturtenbergeri Cappetta, 1975 Scltizorkiza un. slronwri Cappetta, 1975 ' Order Myliobatiformes Family Dasyatidae Dasyaris branisai Cappetta, 1975 Dagtis molinoensis Cappetb. 1975 Dasyatis scltucfleri Cnppeltn. I975 Dasyaris nov. sp. 1 Dasyatis nov. sp. 2 Dasyalis nov. sp. 3 Family Rhoinbodonlidnc Pucabah ltoflslelteri Cnppclln. 1975 Pucabah nov. sp.

CLASS OSTEICHTHYI Subclass Superorder "" Order F'ycnodonti€ormes Family Pycnodontidae roncoensis Benedetto & Sanchez, 1972 Pycnodontidae indet. Order Semionotiformes Family Semionotidae Lepidoles sp. nov. gen. Order Ginglymodi Family Lepisosteidae Lepisosteus sp. Superorder Tclcostci Order Clupciforincs . Family Clupeidae Gasreroclupeo branisai Signeux in Branisa et al. 1964 Order Osteoglossiformes Fnrnily Oskoglossidse Subfamily Phareodontinae Pkaerodusiclitlzys tavèrnei Gayet, 1991 Subfsinily Osteoglossinae nov. gen. Order "Salmoniformes" Family Enchodontidae Emhodis sp. Suborder ichthyotringoidei Family incertae sedis 1 ? Apdeodus sp. Family incertae sedis 2 gen. and sp. indel. Order Family incertae sedis Moliniclitliys inopinatus Gayct, 1982c Order Characiformes Family Erythrinidae Hoplias nov. sp. cf. Hoplias Family Senasalmidae Subfamily Myleinae gen. and sp. indel. cf. Subfamily Serrasalminae gen. and sp. indel. Family Characidae Subfamily Tetragonopterinae gen. and sp. indel. cf. Subfamily Rhoadsiinne gen. and sp. indel. Order Siluriformes Family Ariidae Rliineastes Superfamily Andiniclithyoidea Family indef. nov. gen. Family Andinichthyidae Andiniclttliys bolivianensis Gayet, 1988b Family incertae sedis 1 Incaichfhys suarezi Gayet, 199Ob Family incerlae sedis 2 Hoffsfettericlitkys pucai Gayet, 19Wb at least five gen. and sp. nov. Superorder Atherinomorpha cf. Order Cyprinodontiformes gen. and sp. indel. Superorder Acnnthopterygii Order Perciformes Suborder Percoidei Family Centropomidae gen and sp. indel. Order Tetraodontifonnes Family Eotrigonodontidae Slepl~anodrrsniinintus Gayet, 1991

. Subclass Order Dipnoi Family Ceratodontidae Ceratodirs sp. gen. and sp. indel. Family Lepidosirenidae Lepidosiren cf. paradoxa

CLASS CLADISTM (= BRACHIOPTERYGII) Order Polypteriformes' Family Polypteridae Dajetella sudaniericana Gayet & Meunier, 1991

406 FOSILES Y WalsDE DOLMA - VOL 1- VER’IHBRAWS

CLASS AMPIIIUIA Order Anua Family Lepldactylidae gen. and sp. indel. Order Gymnophiona Family indel. gen. and sp. indet. Order Urodela Family indel. gen. and sp. indet.

CLASS REPTILIA Order Ch,elonia Family Podocnemididae ? Roxoclielys vilavilensis Broin, 1971 ? ROXOC~ICIYSCJ vilavilcrisis Order Squaiiintn Suborder Lncertilia Family ?Iguanidae gen, and sp. indel. Family indet. gen. and sp. indel. Suborder Family Aniliidae gen. and sp. indet. Family Boidae gen. and sp. 1indel. gen. and sp. 2 indel. Family ?Madtsoiidae gen. and sp. indel. Family Tropidopheidae gen. and sp. indel. Order Crocdilia Suborder Mesosuchia Family Sebecidae Sebecus querejazus Buffetaut & Marshall, 1991 Family Dyrosauridae Sokotosukus aff. ianwilsoni Halstead, 1975 Suborder Eusuchia Family Dolichochampsidae Dolickoclmnpsa niininla Gasparini & Buffclnut, 1980 Order Saurischin Suborder Tlieropda Infraorder Coelurosauria gen. and sp. indet. Suborder Infraorder Souropoda gen, and sp. indel. Order Omithischia Suborder gen. and sp. indel. Suborder Ankylosauna or Ceratopsia Ligabueiclinium bolivianum Leonardi. 1984 Order indef. gen. and sp. indel. CLASS MAMMALIA Infraclass Order Family inder. Jaskliadelphys niinutus Marshall & Muizon, 1988 Order Peradectia Family Peradectidae Subfamily Peradectinae austrinuni (Sigé, 1971) Sublamily Caroloamegluniinae Robertlioffstetferianafionalgeograpliica Marshall et al., 1983 Order Microbiotheria Family Microbiotheriidae cordillerensis Marshall ¿k Muizon, 1988 7 Order 'Didelphimorphia Family Didelphidae Subfamily Didelphinae Pucadelpliys andinus Marshall & Muizon, 1988 Incadelpliys untiqrrrrs M~~liall& Muizon. 1988 Mizyrtedel~~liyspil~~ine~isM~~liiill & Muizoii, 1988 Subfamily Eobrasiliinae Tiulordiafloresi Marshall & Muizon, 1988 Order Sparassodonta Family Hathliacynidae Aflqokirrlsaustralis Marshall & Muimn, 1988 Order Polydolopoidea Family Pol ydo Iopídae Epidolops sp. Order Family Kollpaniidae Kollpania fiupanipinaMarshall & Muizon, 1988 Order indef. Family indef. Andinodelpliys cocliabunibensis Marshall & Muizon, 1988 Infraclass Order Lepticrida Family Pnlnporyctidae? cf. Ciniolesres sp. Order Pantodonta Family Pantolambdidae Alcidedorbignyn inopinafa Muizon & Marshall. 1987a Order Condylartlua Family Hyopsodontidae Subfamily Mioclaeninae Tiuclaenus niinirfus Muimn & Marsliail, 1987b Molinodus suarezi Muizon & Marshall, 1987c Andinodus boliviensis Muizon & Marshall, 1987c Pucanodus gagnieri Muizon &.Marshall, 1991 Order Notoungulata Family cf. Henricosbomiidae or Oldfieldtliomasiidae gen. and sp. indef.

408 ~~ ~

~

3. TABLE SYSTEMATIC LIST OF VERTEBRATES FROM THE MESOZOIC AND PALEOCENE OF BOLIVIA WITH INDICATION OF LOCALITIES AND STRATIGWHIC OCCURRENCE BASED ON INFORMATION IN TABLE 1 AND TEXT. Abbreviations: Ca. Castellón Formation; Mi, Mirdores Formation; Ar, Aroifilla Formation; Ch, Chaunaca Formation.

EI Molino Formation TAXA Ca Mi Ar Ch Santa Lucía Impora -Fm -Fm -Fm -Fm lower member middle mb upper member Fm Fm hsSclachii Order Rajiforms Suborder Sclemrhynchoidei Family Sdcmrhynchidae Puraprisrir branisì 11.12 14.20.21 2.25 lschyrhiza harrenbergen 7,11,20,21 25 Schizorhiza 4.srromcri 7.11 25 Order Myliobatifonnes Family Dasyatidae Dqaris bradsat' 20 25 Dqaris molinoemis 11.20 Dqaris Schaefferì 11.20 25.28 Dqatis nov. sp. 1 7 Dasyaris sp. nov. 2 27 Dqatis nov. sp. 3 Family Rhombodontidae n Purabarir hoffsreneri 11.20.21 Purabatis nov. sp. 7 lass Ostcichthyes ;uklassActinoptcrygii Maclass "Holostei" Order Pycnodontiformes Family Pycnodontidac Coelodur roncoqsis 12 and sp. inder. gen. 3? 6 7.12.13.21 25,27,30 Order Sanionotiforma I Family Smiqnotidac Lepidores sp. 2 and sp. gen. nov. 1.9.21 25 Order Ginglymodi Family Lepisostcidac LepisosteUr sp. 7,9,13 25.21 nfmclass Teleostei Order Clupciformes Family Qupcidae Gasreroclupea branisaÌ 25 S? 7.8.9,10,11,12,.15 3235.36, 37 51

- - - - - El Molino Formation ~ mpxa TAXA Mi Santa Lucía Fm Fm lower member middle mb upper member Fm -

.-. ..

Order Osteoglossifo~es I, 2521 49.51.52 Family Osteoglossidae 13 Subfamily Phareodontinac 26 51 Phareodurichthystavermi Subfamily Ostcoglossinae 51 gcn. and sp. nov. Ordcr "Sahoniformes" Family Enchcdontidae 25 sp. Enchodus Suborder Ichthyoiringoidei Family inccmc sedis 1 25 ?Apareo& sp. Family incenac sedis 2 29 gen. and sp. indct. MerCypriniforms Family incenae sedis 25 Mdlim'chrhys inopinafus Order Characiformes Family Erylhrinidac 51 e Hoplias nov. sp. 25 ' 51 O cf. Hoplias Family Semsahidac Subfamily Myle/íÏe 51 gen. and sp. inder. 25.30 d Subfamily Senasalminac 51 56 gen. and sp. indef. Family Characidae SubfamilyTetragonopterinac 25 51 gen. and sp. inder. cf. Subfamily Rhoadsiinae 51 gen. and sp. indef. Ordcr Siluriformes Family Ariidae 7,9, 75,21.30 41-52 56 Rhineasres sp. 12, 13, 14.21 33,35,38 Superfamily Andinichthyoidca Family indet. 51 gen. nov. Family Andinichthyidae 51 Andinichthys boiivianensis 12 34.35 49,51,52 rtndm'chrhys5p. Family incertae redis suarezi 51 Incaichrhys 51 Hoffsfetrerichrhyspucai

7 - El Molino Formation TAXA Ar Santa Lucía Fm lower member middle mb upper member Fm

Superorder Athcrinomorpha d. Order Cyprinodontiforma . .. -. gen. 25 and sp. inder. 7,12,22 35 Superorder Acanthoptcrygii Order Percifoms Suborder Raidci Family Centropomidae gen. and ider. sp. 50.51.52 Order Tetraodonrifomes Family Eotrigonodontidae I, Srephamdus m'mhu 13 25.27 'ubdas Sarwptcrygii Order Dipnoi 53.54 Family Ccratodontidae Cerotodur sp. 51.52 Family Lepidcsiddac Lepidosiren cf. paraaka 12,21 SI. 52

ass Cladistia (= Brachioptcrygii) Order Polyptcrifoms Family Polyptcridae Dajetella sudamericana 21 ass Amphibia Order hura Family Leptodactylidae gen. and sp. inder. 51 Order Gymnophiom Family inder. gen. and sp. inder. 51 Order Urodcla .Family inder. 12 gen. and sp. inder.

ES Reptilia >der Chelonia 14 30 4849.50 Family Podocncmididae 33.35.38 ? Roxochelys vilavilensir 51-54 ? Roxochelys cfi vilavilensis 20.21 Order Suborder Lacertilia Family ?Iguanidac gen. and sp. inder. Family inder. 51 gen. and sp. inder. Suborder Ophidia 51 Family Aniliidae gen. and sp. inder. 51 - Ca Mi Ar Ch El ?olino Form: Santa Lucía Impora TAXA Fm Fm Fm Fm lower member middle mb upper member Fm Fm

Family Boidac gen. and sp. inder. 1 51 gen. and sp. inder. 2 51 Family ?MávIatroiidae gen. and sp. inder. 51 Family Tmpidopheidac gen. and sp. inder. 51 Order Cmcodilia 12 21 ?.5,27,28,30 33, 38 48-50, 51 . Suborder Mesmuchia Family Scbecidae 51 Sebeeu querejazu 54

Family Dyrosauridac 26 51 Sokoiosh off.iamviLrod 51 Suborder Eusuchia Family Dolichochampsidae Doliehochampsa minima 33 Order Saurischia Suborder Thcmpoda 19 Infraorder Coeiumsauna sp. gen. and inder. 12,16,17. 18 Suborder Saumpcdomorpha &order Saumpoda gen. and sp. inder. 19 Order Omirhischia Suborder Omithopoda sp. gen. and inder. 19.3 - Suborder or Ceratopsia Ligabueichnium bolivianum ' 19 Order Inder. gen. and sp. inder. 4 lass Mammalia nfraclass Metatheria Order Deltathemida Family inder. Jmkhadelphys minufus 51 Order Peradmia Family Peradcctidae Subfamily Pcradeainae Peradectes aurrrinum 51 Subfamily Camloameghh/mae Roberrhoffsreneria nafionalgeographica 51 EMolino For ition Santa Lucía Impor: TAXA lower member middle mb upper member Fm Fm

__ . .. - ..

Order Microbiotheria Family Microbiotheriidae Khasia cordil!eremis 51 Order Didelphimorphia Family Diddphidae 50

Subfamily Didclphinae andinus 51 Incadelphys mùqm 51 Mizquedelphys pilpinensis 51 Subfamily Eobrasiliinac Tiuloraïafroren' 51 Order Spalassodonta Family Hathliacynidae -Allqokimausrrab 51 Oder Pol ydolopoidea Family Polydolopidae Epidolops sp. 50 Order Paucituberntlata Family Kollpaniidae Kollpam'a riupampina 51 Oder indct Family inder. Andinodelphys cochabambenris 51 Infmclass Euthcria Oder Lcptictida Family Palacoryctidae? cfi Cimolesressp. 51 Oder Pantadonta Family Pdntolambdidac Alcidedorbignya inopinata 51,52 Order Condylanhra Family Hyopsodontidac Subfamily Moclaeninae Tiuclaenm minurus 51 Moli~durs~lnrezi 51 hd~durboliviensis 51 Pucanadur gagnien 51

Order Notoungulata Family cf. Hennmsborniidae or Oldfieldthomasiidae gen. and sp. indct 51 MIREILLE OAYET. LARRY O. MARSIIALI. &I'IUBRRY SBMI'LKI!

(?Roxocl~elysvilavilensis, complete shells), and mammals I~adrosauricliniisarisfralis, Taponiclinrrs donoftoi, 7'elosiclirt~is (Didelphimorphia. Didelphidae, indet.; Polydolopoidea, saltensis), indeterminate reptiles, and (Yacoraifichnrrs avis) Polydolopidae. Epidolops sp.). Indeterminoted turtles and crocodiles (Alonso, 1980; Alonso & Marquillas, 1986). were observed at Pajcha Pata. The Mealla Formation (a Santa Lucia equivalent; Fig. 1) has About 3 kin south of Torotoro, remains of lungfish yielded remains of teleostean fishes (indet.), turtles (indet.) and two (Cerntodontidae, sp.; Lepidosirenidae, Lepidosiren cf. no Io un g u 1a t e inn in ni a 1s (Henric o s bo r n i id ae, Sinip son o t us paradoxa), turtles (?Roxochelys cf. vilavilensis), crocodiles and an praecursor, S. nidjor; Pascual et al., 1978), whereas the Maíz Gordo indeterminate mammal were collected from a sequence of fine to Formation (an Impora Formation equivalent; Fig. 1) has yielded medium grained red sands of the Santa Lucía Formation (Marshall et specimens of teleostean fishes (Callichthyidae, revelat us; al., 1985; Broin 1991; Schultze, 1991a) (Fig. 3). Poeciliidae, indet.), pleurodire turtles ("Podocneniis" argenfinensis; At Maragua (18 km west of Sucre) and Chaupi Khocha (11 km Cattoi & Freiberg, 1958) and possibly a notoungulate mammal west-northwest of San Lucas) were collected remains of silurifonns (Henricosborniidae, Sinipsonofus sp.) (Pascual el al., 1981). fishes (Ariidae), turtles and crocodiles by M. Gayet and L. G. In northern Chile, the following vertebrates are reported from Marshall in September 1989. Senonian age rock units: a selachian (Pucapristis brunisi) from the Tonel Formation (=lower part of Purilactis Formation S.I.) (pide H. IMPORA FORMATION Cione e/ QI., 1985) and dinosaurs (sauropods indet.) from the Estratos de Quebrada Blanca de Poquis (an EI Molino equivalent) Fossil vertebrates of possible late Paleocene age were recovered and the base of the Pajonnles Formation (Salinas ef al., 199la. b). from two localities of lhe Impora Formation in the Camargo III bru, vertebrates from the Vilqucchico Formation at its type Syncline. southern Bolivia. One is on the north side of Villa I'acheco locality near Vilquechico, include Pucaprisfis branisi and near the Río San Juan del Oro, about 14 km norlh of Tojo, which has Casteroclupeu branisai (DBvila & Ponce de Le6n, 1971; Cione CI yielded remains of siluriform fishes. Just north of this locality al., 1985: 297). A more detailed faunal list is provided by Jaillard et rcntiiins of crocodi¡cs yid turtles were collected. The other localily is 01. (it1 press). From the hiisc of llicir niidtllc Vilqucchico Forinii~ion at Chaupiuno, about 60 km norUiwest of Tarija. where indeterminate ' (a Chaunaca Formation equivalent) they report three fish fish remains were found. (P/ychofrygonsp., rhinobatid?, actinopterygian). Their upper Vilquechico Formation is divided into three sequences which are IMPORT,GNT LATE CRETACEOUS-PALEOCENE equivalent, 'respectively, to the lower, middle and upper EI Molino VERTEBRATE FAUNAS FROM ELSEWHERE IN SOUTH members in Bolivia. From the lower sequence they report an AMERICA actinopterygian , fish and name two dinosaur trackways (Coelurosauria, Ornithomimidae, Ornithoniiniipus jaillardi; A. ANDEAN BASIN Ornithopda, Hadrosauridae. Nadrosaitriclinus f iticucaensis); from the base of the middle sequence are two fish (Pucapristis brunisi and In northwestern Argentina, a sauropod (Titanosauridae, an actinopterygian); and from the upper sequence are two fish Laplatasaurus sp.) and a theropod (family incerfae sedis, (Dusyafissp. and an actinopterygian) and numerous charophytes. Unqirillosaitrus ceibalii) are known from the Los Blanquitos At Laguna Umayo, from what was called the Vilquechico Formation (a temporal equivalent of the Chaunaca and/or Aroifilla Formation and is now called the Umayo Formation (Laubaclier & formations) in the upper part of the Pirgua Subgroup (Powell, 1979) Marocco, 1990, Jaillard et al., in press) are reported charophytes, (Fig. 1). (Eokenepoidae, Eoxenepoides saltensis) are known actinopterygian fishes (Characiformes, Erythrinidae, Hoplias sp., from the Las Curtiembres Formation, which transitionally underlies Serrasalmidae, Myleinae, indet., Characidae. Tetragonopterinae, the Los Blanquitos Formation (Parodi-Bustos, 1962). indet.; Siluriformes iricertae sedis; Perciforiiics, family indet.), From the (a basal lower EI Molino equivalent; lu n g fis li (Cer a todo n t id tie, Cerato dir s s p., Lep id os ir en id ne, Flg. 1) are reported sauropods (Titanosauridae, Salfasuirrus Lepidosiren cf. paradoxa), frogs (Leptodactylidae indef.), snakes loricatus), Coelurosauria (, Noasaurus leali) and (Aniliidne), turtles (Podocnemididae, ?Roxochelys cf. vilavilemis), theropods (Carnosauria indet. and . Avisaurus crocodiles (indct.), egg shell fragments reportedly of dinosaurs, archibaldi) (Bonaparte et al., 1977; Bonaparte & Powell, 1980; (Pcrndcctidnc, Perudecles arrstrirrrinr; Didelphidac?; Brett-Surman & Paul, 1985; Bonaparte, 1986). The Yacoraite Pedioinyidae or Microbiotheriidae), and two placentals Formation (nnothcr EI Molino equivalent; Fig.1) lias yielded a (Cond y 1art lira, Didolo don t id ae, undes cribed ; No to ling u Ia ta, crocodile (Dolichocliai~ipsidae,Doliclrochunipsa niininru: Gasparini Perutlieriidae, Perutlterirrni alfiplanense) (Grambast cf al., 1967; & Duffetaut, 1980) which was collected in association with a Sigf. 1968, 1971, 1972; Bonaparte & Powell, 1980; Riige. 1981; pycnodont (Coelodits toncoensis; Benedetto & Sanchez, 1972), a Marshall ef al., 1983b, 1985; Kerourio & Sig& 1984; Marshall & clupeid (Gasleroclirpea branisai) and turitellid gastropods (see Muizon, 1988; Schultze, 1991a; Gayet, personal observation). Gasparini & Buffetaut, 1980; Cione et al., 1985: 296). Other taxa reported from various localities and levels of the Yacoraite B. SOUTHERN ARGENTINA AND SOUTHERN BRAZIL Formation include a selachian (Pucaprisfis brunisi) and a siluriform (indet.) (Cione et al., 1985; Powell, 1979: 202); the tooth of an There are four important vertebrate faunas outside the Andean indeterminate theropod (Powell, 1979: 203); and trackways of basin that warrant special consideration: they are' from the Los carnosaurs (Sa~ifichnusnienfoor), ornithopods (Hadrosauridae?. in (late Cretaceous). the Adaitiantina

114 r-.

niid Mnrllin formations in the I’nruiil bnsin (Inle Crctaccous), tile rIIdly irdef.; Coclurosnurio, lnniily nov. I, nov. 2; Ssiiropd;\, ßnnco Negro Infcrior in Pntngonin (middle I’nlcocenc?), nnd ti~e Titnnosauridnc. ?Aritarcfosnirriis brasiliensis, ‘I‘ituriosnrtritscf. fissure fillings in the limestones of São Jose de Itabornl near Rio de australis; Ornithischia). niid mainrnals (I’laccntalia. indct.) (Gayct 6t Janeiro (middle Paleocene?). ßrito, 1989; Bertini el al., in press). Both formations arc of continental, mostly fluvial. origin and arc thought to bc Scnonian in 1. Late Cretaceous age.

From the at Estancia Los Alainitos in Rlo 2. Middle Paleocene? Negro Province, southern Argentina, numerous vertebrate fossils were found in Ulis mainly brackish-lacustrine rock unit which was A local fauna is known from tlie Banco Negro Inferior in the base tentatively assigned a late Campanian-early age of the Río Chico Formation about 1km southwest of Punta Peligro (Bonaparte el al., 1987). The taxa include: selachians (Batoidei, (40 km norh-northeast of Comodoro Rivadavia along the indet.), holosteans (Semionotidae, Lepidoles sp.; Lepisosteidae, cf. coast) and another at Las Flores, 60 m above the Banco Negro Alractoslerrs sp.), teleosteans (Siluriformes, cf. Diplomystidae, cf. Inferior in the base of the Gran Barranca south of Lago Colhué- Ariidae; Percifohnes, Percoidei, inder.), indef. Huapí. Both local faunas are in the San Jorge basin, Chubut (Lepidofes or Sparidae), Dipnoi (Ptychoceratodotitidae, Cerafodirs Province, southern Argentina and were discovered in January 1979 iheringi), amphibians (, cf. Xenopus sp.; Leptodactylidae, during a National Geographic Society sponsored research program indef.),turtles (Meiolaniidae, cf. Niolamia; Chelidae, indef.),snakes under the direction of one of tlie authors (Marshall el al.. 1981). The (ßoidnc, Mndtsoniinnc. Alaniiropliis argenfiniis, Patagoniophis fossils from Punta I’cligro includc turtles (Chelidnc, at least four parviis, RionegropIris ntadtsoioides), dinosnurs (Titnnosauridnc, taxa; ßroin, 1988), crocodiles (Scbecosuchia?; Crocodylidne. Aelosarirus rionegrinris?: Hadrosauridac, Krilosauriis auslralis), Necrosirchus ionensis Simpson. 1937; Alligntoridac, Exainian sp.. and innminals including Symmetrodontn (Bondesiidne. Bondesiiis Allognatosrichris? sp.), a possiblc edentnte or niultitubcrculate fcrox; Inm. in de^., Casaniiyiielia riomgrina: ?Spalncotlicriidac, (Sudnmcricidnc, Sudrinicrica cinrcghirioi Scilleto Ynnd ¿iPnscuul. Brandonin interdiedia; B arbereniidnc, Barberenia araiijoae, 1985), ( and/or Mioclncnidae) and several Qrtirogaflteriirna niajor), Dryolestoidea (, indeterminate bone fragments (Marshall el al., 1981, n16; Pascual & Groeberllieriunr slipanicici, G. novaci, ciispidaliis; Ortiz-Jaurcguizar, 1991). Those from Las Flores include the Mesungulatidae, Mesiingulaluni Iioussayi; Reigitheriidae, polydolopoid Epidolops sp. (I’ascual C Bond, 1981). and Reigilheriuni bunodonla), Triconodonta (, other taxa which show affinities with faunas in Brazil Arislrotriconodon nickennai), (Ferugliotheriidae, (Poscunl & Ortiz-Jaureguizar, 1991). Many additional fossili of Ferugliofheriuni wiridltauseni) and two Gondwanatheria marsupials and from Las Flores were collected by L. G. (Gondwanntlieriidae, Gondwanafheririni palagoniciinr, Vucefichia Marshall and colleagues in 1979, and numerous unpublished gracilis) (Albino. 1987; Baez, 1987; Bonaparte & Soria, 1983, specimens from both Punta Pcligro niid Las Flores are now available 1985; Bonaparte, 198Ga, b, c, 1987, 1990; Bonaparte & Pascual, (Van Valen. 1988). The Banco Negro Infcrior is predominant& a 1987; Bonaparte & Rougier, 1987; Broin, 1987; Cione, 1987; black bentonitic mudstone (Andreis el al., 1975). The Salamanca/Río Mones, 1987; Powell, 1987). In northern Patagonia, some levels of Chico contact records a noteworthy marine regression which, the Los Alamitos Fonnation are probably associated with a marine according to associated geochronologic age constraints (Marshall el transgression from the west (Andreis, 1987). The Los Alarnitos al., 198l), probably corresponds to the major regression identified Formation is overlain by the transgressive marine Roca Formation of by Haq el al. (1987) in tlie early (about 58.0-58.5 Ma). Mnastriclitian-Daniail age (Bonnparte, 1990) which probably The Itaborai fauna comes from fissure fillings in thearly SEO corrclatcs with thc cocvnl nnd trnnsgrcssive lower EI Molino Pnlcoccnc (?) ngc Jose de ltnbornf Forinntion locntcd about 25 Forniution of Bolivin. ßccnusc of their chnractcristics and kin eilst of Nitcr6i. stntc of Rio de Jnnciro, Brnzil. Taxonomic lists of stratigraphic positions, we thus propose temporal correlation of the this exceptionally diverse fauna are provided by Paula Couto (1970). Los Alarnitos Formation with the Chaunaca Formation. Hence, the Palma & Brito (1976) and Marshall el al. (1983). The vertebrates Los Alamitos Formation is likely to be of Cnmpanian age. wcre recovcred from numcrous karst cavitics in a lower liniestonc The vertebrate faunas from the Adainantina and Marilia unit and predate an upper limestone unit (Brito et al., 1972). For formntions (Upper Baum Group) in. the northern part of tlie Paran6 most specimens, their association among themselves and/or with basin, southcentral Brazil, include fishes (Lepisosteiformes, specific cavities was never recorded. Although sonic nianiiiials Lcpisostcidnc, Lcpisosreus coniinatoi; Osteoglossiformes, appear referable to what has been interprcted as middle Paleocene Osteoglossidac; Characifornies; Silurifornlcs, cf. Doradidae; time (i.e. the Itaboraian Land Maniinal nge; sensu Marsliall, 1985). Perciformes. ?Percichdiyidae), lungfish (Dipnoi, Neoceratodontidae, others appear more progressive and may represent faunas of late Neocet afodris sp.), frogs (Lcptodactylitlac, Ba~irirbafracliiispricei). Paleocene and possibly cai ly Eoccne iigc (i.e. the Kiocliican and Land Mammal Ages, respectively; see Soria. 1987; turtles (Podocnemididae, Roxoclielys irarrisi, cf. R. elegam, aff. A. Podocnentis brasiliensis), lizards (Iguanidae?, Prisligiiana Van Valen, 1988; Cione, personal comunic?tion). In the brasiliensis), snakes (inder.), crocodiles (, following discussion, we regard this fauna as middle Paleocene, but Barrrrisuclius pachecoi; , forminni; caution that it may include taxa that accumulated between middle Goniopholidae, ?Goniopholis paulislanrts; Trematosuchidae, Paleocene nnd early Eoccne time. Itasrichus jesuinoi; family indel.. Brasileosaurus pacltecoi, Although a hydrothermal origin been clnssically favorcd for SEO has Splragesaiirus huenei), dinosnurs (Carnosnuria, Abelisnuridae. the Jose de Itaboraí lncustrinc limcstones (Francisco gZ Cunlin.

415 MIREILLE OAYCT. LARRY O. MARSIIALL L’I‘IUUKRY’SCMI’BKI!

1978). water level in the tectonic iicnr-shore lake where tlie Chile (Schultzc. 1981). niid existed uiitil the iiiiddlc Eocene in limestones were deposited might have been controlled by the level of marine deposits around the world (Blot, 1987). Cione (1977) the nearby through adjustment of marine and tentatively referred specimens discussed by Wenz (1969) from continental phreatic levels. In such a case, the lacustrine episode Bolivia to Coelodus loncoensis which was erected by. Benedetto & evidenced by the lower Itaboraí limestones would have been coeval SQnchez (1972) upon material from the Yacoraite Formation of with a marine highstand. Since middle (and later?, see above) northwest Argentina. Gayet (1991) cautions that the spccics identity Paleocene mammals are known from the infilling of the karstic of some Bolivian pycnodontids has still to be verified, while olhers cavities Uiat forma after lake dessication, it would be possible to from the lower and basal middle EI Molino Formation can be correlate this highstand period with the one in the late Danian, and referred with confidence to Coelodus foncoensis which was the subsequent dessication with the early Thanetian regression which apparently endemic LO the Andean basin. is also recorded near the Salamanca/Río Chico contact in southern Argentina (see above). A subsequent highstand period would have 3. Semionotiformes resulted in the deposition of the upper limestones of São Jose de Itaboraf. In Bolivia, Lepidofes sp. was collected securely from the late Triassic-early Jurassic. Elsewhere in , Lepidofes is DISCUSSION reported from the (=Portlandian) of Neuquén Province, Argentina (Aramayo, 1981). the -Albian.of Brazil (Agassiz, A. UIOSTRATIGRAPIIY 1841; Roxo & Löfgrcn, 1936; Woodward, 1895, 1908) and a possible record froin the lntc Cniiipiiiiiiiii-eiirly Munslrichtiun ngc Los The Mesozoic and I’oleoccne age vertebrate fossil localities in Alainitos Forination in Argentina (Cione, 1987). Elsewhere in the Bolivia (Fig. 2) are arranged in Table 1 according Lo their world. is recorded from the Rlietian of Germany to stratigraphic context. A systematic list of all the known fossil Cretaceous/Paleocene of India (Gayet et al., 1984). vertebrates is given in Tables 2, and in Table 3 with indication of Pillow-shape scales which rcprcscnt u new gctius of thc family locality and stratigraphic occurrence. Semionotidae are known only in the lower and basal iniddle In this section we review the known occurrence of the taxa listed members of the EI Molino Formation in Bolivia (Gayet & Meunier, in Tables 2 and 3 in order to identify their cluonostratigraphic ranges personal observation) and in the Yacoraite Formation in Argentina and hence potential usefulness for calibratin8 the Bolivian rock (A. L. Cione, pers. com.). I sequence. Of particular interest is the apparent position of the Cretaceous/Paleocene (K/T) boundary.within the 81 Molino 4. Cinglymodi Formation (see below). In Bolivia, Lepisosfeus sp. is reported from the lower and basal 1. Selachians j middle El Molino. The earliest Lepisosteidae are reported from the of Niger (Ariunbourg & Joleaud, 1943) and Congo In Bolivia, selachians were recovered from the lower and basal (Casier, 1943). Younger fossils, including Lepisosteus, are known middle members of the El Molino Formation. Rhombodontidae are from the late Cretaceous of India (Jain & Sahni, 1983; Gayet el al., present only in the lower member, whereas Sclerorhynchidae and 1984) and from the of Euiope, India and Dasyatidae are present in both. Sclerorhynchidae range from Albian (Wiley, 1976). In South America, Lepisosteidae are known in to Maastriclitian (Cappetta, 1990). Schizorhiza is known only in Argentina from the late Campanian-early .Maastrichtian age Los Maaskichtian age rocks from numerous localities around tlie world Alamitos Formation (cf. Atracfosferrssp.; Cione. 1987) and from the (A. (, Morocco, Middle-East, Niger. Nigeria and Zaire), while EI Abra Formation of uncertain age L. Cione, pers. com.) Ischyrliiza ranges from the Turonian to Maastrichtian in North (Lepisosferts; M. G.. pcrs. obs.); Lepisosteris is known froin the late America and Niger (Cappetta, 1987, 1990, 1991). Pucaprisfis Cretaceous Adainantina and Marilia formations of the Upper Bauru (Sclerorhynchidae), Pitcubatis (Rhombodontidae) and the three Group in Brazil (Santos, 1984; Gayet & Brito, 1989; Brito & Gayet, D. D. named species of Dusyuris (D. molinoensis, scltaefferi, in press), and the late Cretaceous of Colombin (Gayet, 1991). brunisui) (Dasyatidae) are currently endemic to the Andean basin Lepisosfcits cxists to Kcccnt in southenstern USA and in Cciitrul where they are proving exceptionally useful in intra-basin correlation America, (Cappetta, 1990). Pircuprisfis branisi is ‘also known from a level below another which yielded dinosaurs in the lower part of the 5. Clupeiformes Yacoraite Formation in northwest Argentina (Powell, 1979). Dasyalis is known in the Cenomanian of Texas, and from Gusferoclupeu brunisui is apparently endemic to the Andean basin Maastrichtian to Recent world-wide. where in Bolivia it is reported from the Chaunaca (possibly), Cajones, all three members of the El Molino, and the Santa Lucía 2. Pycnodontiformes formations; in Argentina it is known only from the Yacoraite Formation (Cione el al., 1985) and has never been found in Pycnodontiformes (Pycnodontidae) were collected in Bolivia from overlying Paleocene formations. The oldest known Clupeidae are the Chaunaca Fondation and the lower and basal middle members of from the Neocoinian of northern. Kyushu, Japan (Diplonlysrus the El Molino. Coèlodus was reported from the lower Cretaceous of prinzofiniis, D. kokirraensis} (Uyeno, 1979). Brazil (Santos, 1963; Wenz, 1989). (Porta, 1970) and

416 uinnyo in Perii (Gnyct. 199 I), Mioccnc Loyoln Formntion of Ecuador (Roberts, 1975) arid Mioccnc of Coloiiibin (Lundbcrg et al., In Bolivia, the subfamilies Osteoglossinae and Phareodontinae of 1986); Characidae (Tetragonopterinae) from the Umayo Formation the family Osteoglossidae are recorded from the EI Molino and at Laguna Umayo in Peru (Gayet, 1991). of Ecuador Santa Lucia formations (see Table 3). The earliest record for (Roberts, 1975) and early Mioceile of Taubaté in Brazil (Schaelfcr, Ostcoglossiformes in South America is from Uie Aptian age Areado 1947); Characidae (indef.) from the Paleocene of Morocco (Cappeta Fonnation in Brazil (Santos, 1985) where the taxon is referred to tlie ef al., 1978); Serrasalmidae (Serrasalminae) and Characidae family Arapaemidae (Laeliichtliyinae) which lias African affinities. (Rhoadsiinae) had not yet been found as fossils. The oldest known Phnreodontinae were reported from the Paleocene and Eocene of record of characiforms outside of South America is Salniinups (Cappetta, 1972) and Europe (Woodward, 1901; Taverne, ibericus from the marine late Cenomanian of Portugal (Gayet, 1978), Paleocene of southeast (Sanders, 1934), late Paleocene 1985a). Characiforms range.10 Recent in Africa and in South of Turkmenia (Danil'chenko, 1968), Paleocene of America. One recent colonized Central America. (Taverne, 1975; Gayet, 1991). and middle Eocene of North America. Thus, Lhe Phareodontinae record from the EI Molino is the earliest 10. Siluriformes known to date. Squamules (parts of scales) of Osteoglossidae (Phareodontinae In Bolivia. this order is represented by members of the families rind Osteoglossinae) were reported in the Maastrichtian of Niger, Ariidae and Andinichthyidae from both the EI Molino and Santa CrelaceouslPaleocene of India and Paleocene of Pnkistan (Gayet & Lucía formations, and at lcast by two families inccrfae scdis from Mcunicr, 1983). Ostcoglossinae wcrc rcportcd from tlic I'aleoccnc of thc Snntn Luch Forinntion nt Tiupempn (Gnyc~,199 1). Sumatra (Sanders, 1934), and range lo Rcccnt in South America, The other known siluriforliis elsewhere in Soutli America are Asia, Africa and Australia. represented by cf. Diplomystidae and cf. Ariidae in tlie Carnpanian age Los Alamitos Formation in Argentina (Cionc, 1987); 7. "Sulmoniformes" Siluriforiiics indc/. from Ilic Mnns~riclitinnrtgc Coli Toro (Cionc C Laffite, 1980) and Yacoraite formations (Cione et al., 1985) in In Bolivia, Enchodontidne (Enclrudus sp.) and Ichthyotringoidei Argentina; cf. Doradidae from the late Cretaceous age Adamantina (family indet. and 7Apafeudus sp.) were collected Gom the lower and Marííia formations in the Upper , Brazil (Gayet & and basal middle EI Molino. Encltudus is known only from marine Brito. 1989; Brito & Gayet, in press); and Ariidae from the Maastrichtian ake rocks in Congo (Dartevelle & Casier, 1949). Maastrichtian? age Umayo Formation at Laguna Umayo in Peru Morocco (Arambourg, 1952). Brazil (Rehusas & Santos, 1956), (Gayet, 1991). Europe (Goody, 1968), North America (Goody, 19691, Lebanon The only records for Cretaceous age Siluriformes outside of South (Goody, 1969) and Israel (Chalifa, 1989) to note only the most America are a single tooth referred to Arius sp. from the latest important in both preservation and quantity. Cretaceous of Central India (Jain & Salini, 1983) and doubtful Apafeudus is a member of the suborder Ichthyotringoidei which otolithes from the late Cretaceous of North America (Frizzell, 1965; ranges from early Cretaceous (?Albian) to Maastrichtian (Goody, Fitch, 1975). 1969). Siluriformes are known in the Paleocene of Argentina, [Arius? and Baclintania in Patagonia (Dolgopol de Saez. 1941), and 8. Cypriniformes Prupygidirim (Bocchino, 1964) and Coryduras (which in Recent limes is restricted to freshwater in South America) from the Maíz The cypriniform (Mulinichfltys inopinafus: Gayet, 1982b) from Gordo Formation (ßardack, 1961)J. Africa (in Congo; Dartevelle & Ihc lower and basal middlc EI Molino rcprcscnts tlic only record, Casicr, 1949; Cnsicr, 1960), Europc (ßclgium; Leriche, 1902). fossil or extnnt, of this ordcr in South Amcrica. Elscwlicrc in lhe Indoncsia (Sandcrs. 1934); nnd in tlic Eoccnc of North Anicricn world, this group is' first known from the Eocene of North America (Lundberg, 1975; Grande. 1987; Grande & Lundberg, 1988), Europe (Grande ef al., 1982). Gayet (1986b. c) described Raniallichfhys (Woodward, 1901; Casier, 1946), Africa (Peyer, 1928) and India from the marine Cenomanian in Israel which may be considcred an (Salini & Misra, 1975). Ariidae range to Recent. ancestral, if not true, cypriniform. 11. Cyprinodontiformes 9. Characiformes In Bolivia, pharyngeal teeth from the Miraflores Forniation are In Bolivia, this ordcr is rcprcscnlcd by the families Erytluinidac, tcntatively referrcd to indelcrininate cyprinodontiforms. In the lower Serrasalmidae (including Serrasalminac and Myleinae) and member of the EI Molino Forination are known several very Characidae (including Tetragonopterinae and Rhoadsiinae). All these primitive cf. Cyprinodontiformes (Gayet, 1991). The earliest groups have been found in the Santa Lucía Formation, while previous record of this group,was from the of France members of the Erythrinidae, Myleinae and Tetragonopterinae were (Gaudant. 1988). and it ranges to Recent world-wide. also found in tlie lower and basal middle EI Molino Formation. The oldest previous known Erytlirinidae (Nuplias) are reported 12. Perciformes from the Miocene of Ecuador (Roberts, 1975); Serrasalmidae (Myleinae) from the Maastrichtian? Umnyo Formation at Laguna In Bolivia, perciforms of lhe family Cencropomidae are known

417

i , MIREIUE OAYIX. LARRY O. MARSIMLL EL IIIIBKRY SEMI'IJKC from the Santa Ldcía Formation (Gayet, 1991). Indeterminate 16. hmphiblans specimens of this order nre reported in South America from the Campanian age Los Alamitos Formation of Argentina (Cione, 1987). In Bolivia, indeterminate -eptodactylidae are known only from the Maastrichtian age Marília Formation of Brazil (Gayet & Brito. the Santa Lucía Formation at Tiupampa. Elsewhere in South 1989). and the Umayo Formation at Laguna Umayo in Peru (Gayet, Americn indeterminate Leptodactylidae are recorded from the 199 1). The earliest' Centropomidae were previously recorded from Campanian nge Los Alamitos Formation in Argentina (Baez, 1987), the middle Eocene of Italy (Sorbin¡, 1970), and they range to Recent and, questionably, from the Umayo Formation at Laguna Umayo in world-wide. Peru (Sig&, 1968). A probable Leptodactylidae, Bairrubafraclius pricei has been recorded in the Maastrichtian age Marília Formation al., 13. Tetraodontiformes in Brazil (BQez, 1985; BQez & Perí, 1989; Bertini et in press). The oldest known Leptodactylidae are from the Manía Formation In Bolivia, Slepltanodus (Eotrigonodontidae) is recorded from the and the family extends to the Recent (BQez, 1987). lower and basal middle El Molino Formation. Eotrigonodontidae The oldest known Gymnophiona are represented by two vertebrae was reported from marine beds in the ?Albian-Aptian of Morocco from the Santa Lucía Formation at Tiupampa (Rage, 1986, 1991b). (Tabaste, 1964). Cenomanian of (Weiler, 1935), late Previously this group was known from the middle Paleocene (at Cretaceous of Nigeria (White, 1934), Congo (Dartevelle & Casier, Itaboraí) to the Recent. 1949), Niger (Tabaste, 1963; Cappetta, 1972). Morocco Vertebrae of urodels are known from the lower El Molino (Arambourg, 1952), Israel (Raab, 1963), Upper Cretaceous of lndia Formation at Pnjchn Pnta. This is the earliest record of this group in (Jain & Salmi, 1983) nnd levels of Europe, (Leriche, 1906; the world (Kage, pers. com.). Casier, 1946), Angola (Dartevelle & Casier, 1959) and Egypt (Peyer, 1928). Eotrigonodontidae range from ?Albian-Aptian to middle 17. Turtles Eocene, whereas Sfeplianodus was reported only from the Cretaceous (sanie rdccrcnces). Roxochelys was based on spccinicns collected [rom thc Campaninn nge Adamnntinn Formation or the Upper Bauru Group in al., 14. Polypteriformes Brazil (Broin, 1991; Bertini et in press). ?Roxoc/ielysvilavilensis Broin, (1971), from the Santa Lucía Formation is probably referable Dajefella sudamericana (Gayet & Meunier, 1991a, b, 1992) from to a genus distinct from Roxochelys (Broin, 1991). The species the lower El Molino and Santa Lucía formations is the only record vilavilensis is definitely known only from lhe Santa Lucía Formation for Polypteriformes outside of Africa, where members of this order in Bolivia and possibly frpm the Maastrichtian? Umayo Formation at are known from the Senonian of In Becetem in Niger (Gayet et al., Laguna Umayo in adjacent Peru (see above), while very fragmentary 1988), Miocene of and Kenya (Greenwood, 1951,,1973), and material from the lower EI Molino of Bolivia is referred to of Ethiopia (Arambourg, 1948). They range to Recent in ?Roxoc/ielyscf. vilavilensis. This species was apparently endemic to Africa where they occur in freshwater. the Andean basin.

15. Lungfish 18. Lizards

In Bolivia, Ceratodontidae (Cerafodus sp. and ceratodont n. sp.) An unidentified member of what appears to be the family and Lepidosirenidae (Lepidosiren cf. paradoxa) tooth plates occur Iguanidae is recorded from tlie Santa Lucía Formation at Tiupampa together in the early Paleocene of Tiupampa. In South America, the (Rage, 1991b). A possible iguanid (Prisfigrrattabrusiliensis; Estes & first ceratodonts occur in the Upper Cretnceous of Patagonia Price, 1973) is known from the Manstrichtian agc Mnrflia Formation (Ameghino, 1906; Pascual & Bondesio, 1976; Cione, 1987) and of the Upper Bauru Group in Brazil, while he earliest uncontested were also described from the Lower Cretaceous of northern Brazil members of this family are from the middle Paleocene of Brazil (Cunha & Ferreira, 1980). Ceratodonts were distributed world-wide (Estes, 1983) and North America (Sullivan, 1982). in tlie Mesozoic, still occurring during the Cretaceous in Africa, Madagascar, Australia, North and South America, whereas the 19. Snnkes Neoceratodontidae which occur in the early Cretaceous of Australia persist there until today (Schultze, 1991a). The four faitlilies of snakes listed below are known only from the Lepidosirenidae are reported from the Maastrichtinn El Santa Lucía Formation at Tiupampa in Bolivia. Molino Formation at Vila Vila and the Paleocene Santa Lucfa In South Americn, Aniliidhe are first reported from the Umayo Formation at Torotoro. Fossils are recorded from the Formation at Laguna Umayo in Peru (Rage, 1981) and middle Maastrichtian? at Laguna Umayo in Peru (Sigé, 1968). Paleocene at Itaboraí in Brazil (Rage, 1987). Eocene of Argentina (Fernandez et al., 1973; Fernandez, The earliest Boidae are from the Maastrichtian of North America 1976) and Miocene of Brazil (Santos, 1987) and Colombia (Estes & BBez, 1985), Europe (Rage, 1987) and India (Jain & Sahni, in (Bondesio & Pascual, 1977). Lepidosirenids occur today in 1983), while South America, previous to this work, they were first the Amazonian and Paraguayan regions of South America. reported in the middle Paleocene in Brazil (Rage, 1987) and early Eocene of Argentina (Albino, 1986).

418 Y 8. FOSOSILES FACIES bE noum - VOL. I - VERTEBMWS

Madtsoiidne nrc first kiiown in the Suiitoiiian/Cninpnnian of 22. M,unimuls Mndngnscnr (Rage, 1984; Roiinparte. 1986) niid Campaninn age Los Alainitos Formation in Argentina (Albino, 1986, 1987). In fact, Uie Mammals are known only from the Santa Luch Formation in oldest Madtsoiidae is probably from Niger (early Senonian) (Rage, Bolivia. Peradectes airstriniim was based on a specimen collected 1984). from tlie Umayo Formation at Laguna Uinayo in Peru (Sig&, 1971, Previous to this work, Tropidoplieidae were first reported from the 1972; Crochet, 1980) and a second specimen from Uie Santa Lucía middle Paleocene hi Brazil (Rage, 1987), middle Eocene in North Formation at Tiupampa in Bolivia was later referred to this species America (Hohnan, 1979), and Eocene-early Oligocene in Europe (Marshall & Muizon, 1988). The earliest known Perudecfes is from (Rage, 1984). the base of the early Paleocene in North America (Van Valen, 1988: 29) and is common thereafter in early Tertiary age rocks in North 20. Crocodiles America and Europe (Crochet, 1980). Robert/ioflsferteriais most closely related to Procuroloanreglrinia Three families are known in Bolivia: Dyrosauridae (basal middle from the middle Paleocene of Brazil (Marshall et al., 1983a), EI Molino and Santa Lucia formations), Dolichochampsidae (upper although other more primitive members of the subfamily EI Molino Formation) aiid Sebecidae (Santa Lucía Formation). Caroloaineghiniinae are known from the Maastrichtian in Norih Dyrosauridae are known from late Maastrichtian to late Eocene America (Marshall et al,, 1989). The genera and species of rocks around the world (Buffetaut, 1982). In South America, Mi'crobiotlieriidae, Didelphidae, Hathliacynidae and Kollpaniidae are dyrosaurids are known fTom what appears to be the Mnastrichtian in known only Erom the Snnta Lucía Formation at Tiupampa, and they ßrazil ,(Cope, 1886; Buffetnut, 1978), and a possible dyrosaurid is all represent the earliest known members of these faniilies in South kiiowii from ¡tic Manstrichtinti in Argentina (Gnspnrini, perß. coin.). Ainericri (MarslinIl & Muiimii, 1988; MiuslinIl et d.,1989). Sokotosuchus ianwilsoni (Halstend, 1975) is from the late Cf. Cintolestes sp. is the only known member of the order Maaskichtian Dukarnaje Forination in Nigeria. Leptictida in South America. Cintolestes is recorded froin the Dolichochampsidae (Doli~li~~ltanip~aniininta) is known Mnnskichtinn and cnrly Paleocene in North America (Marshall & elsewhere only from tlie upper put of the Yacornitc Forination in Muizon, 1988). northwest Argentina and the upper El Molino Formation in Bolivia The pantodont Alcidedorbignya is the only known member of this (see above). The family was apparently endemic to the Andean order in South America. Pantodonts are fist known in the middle basin. Paleocene of North America where Panfolambda batlrniodon Sebecidae were previously known from the middle Paleocene- represents a potential descendant of Alcidedorbignya (Marshall & middle Miocene (Buffetaut, 1982; Gasparini, 1984); they were Muizon, 1988). Among the New World pantodonts, Alcidedorbignya

I endemic to South America. Sebecus querejazus from the Santa is the earliest and most primitive member yet known. 1 Lucía.Formation is the earliest known member of the family The mioclaenine Hyopsodontidae are the only representatiyes of (Buffetaut & Marshall, 1991). this family in South America. In North America, members of this group are 6rst recorded in the early Paleocene (Marshall & Muizon, 21. Dinosaurs i 1988; Van Valen, 1988). The earliest known notoungulates are represented by PeruUieriimt Dinosaur trackways are known from four localities in the lower EI altiplanense (Perutheriidae) from the Umayo Formation at Laguna Molino near Torotoro and at Santivañez near Parohni, and from two Umayo in Peru (Marshall et al., 1983b), Sintpsonolus localities (Arapamba, Camargo) in an indeterminate EI Molino (Henricosborniidae) from the Mealla Formation in northwest member. The only dinosaur fossils in Bolivia are a tooth of a Argentina (Pascua1 ef al., 1979), and cf. Henricosborniidae or Coelurosnurin (Marshall, 1989b) from the lowcr El Molino at Pnjchn Oldlicldtlioinnsiidnc from lhe Siintri Luchi Forinntion nt Tiupniiipn in I'ntn, and undcscribcd boiics from two localitice of tlic Cajones Bolivia (Miusha11 & Muizori, 1988). Formation near Santa Cruz (R. Suhez, pers. com.). Although a detailed study of the trackways has yet to be B, THE CRETACEOUS/PALEOCENE (KIT) BOUNDARY IN undertaken, it appears that at least four suborders (, THE ANDEAN BASIN , Ornithopoda and Ankylosauria or Ceratopsia) are represented within the lower EI Molino at Torotoro. This Based 011 the above discussion of chronostratigraphic ranges, the demonstrates that dinosaurs were taxonomically diverse in lower EI taxa can be grouped into five categories (see Table 3): Molino time. For cluonostratigraphic purposes we follow the general mnsensus 1. Those which presently are known only from the EI Molino that dinosaurs became extinct at the end of Cretaceous time and that Formation and its stratigraphic equivalents in tlie Andean basin. there is no compelling evidence (contra Van Valen, 1988) to These are currently regarde$ as "endemics" which are potentially substantiate Uiat they may have continued into the early Paleocene. useful for intra-basin correlation' although their usefulness in Recent work by Lofgren el al. (1990), for example, negates a calibrating these rocks witti the geologic time scale has yet to be Paleocene age for dihosaurs from the upper in firmly established. Included are Pitcapristis (Sclerorhynchidae), Montana. Our position is concordabt with the biostratigraphic data in Prtcabafis (Rhombodontidae), the three species of Dasyafis the Andean basin summarized above and below. (Dasyatidae), Coelodrrs toncoensis (Pycnodon tidae). and DoliclroclIanipsa (Dolichochampsidae).

419 2. Taxa from the EI Molino and Santa Luch formations (and their migrate bctwccn fresh water and marine; nnd 4) marinc fishes. stratigraphic cquivalcnls) which are "endemics" in the Andean basin. Included are Gasteroclicpea bronisai (Clupcidae). Andinichtliys The definition of these four categories is based on living fislies bolivianensis , (Andinichthyoidea, Andinichthyidae), and is not always evident with fossil forms (see below). For this Hoffsterterichtliyspucai, Incaichthys suarezi (families incertae sedis) reason, the identification of paleoenvironment based only on one or a and ?Roxochelys vilavilensis (Podocnemididae). few taxa is generally of dubious value.

3. Taxa known only from the lower and basal middle EI Molino 1. Primary freshwater fishes Fonnation which are currently regarded as late Cretaceous "guide fossils" based on their records elsewhere in the world. Included are a. Semionotidae Schizorhiza and Ischyrhiza (Sclerorhyncliidae), apparently Lepidot es (Semionotidae) which has only one dubious record in the The fossil Semionotidae are generally considered as primary Cretaceous/Paleocene of India, Enclzodiis (Enchodontidae), freshwater fishes. A new genus is reported in Bolivia in the lower ?Apaleodris (family indet.), Ichthyotringoidei (gen. and sp. indet.), and basal middle members of the EI Molino Formation. Except for Sleplranodiis (Eotiigonodontidae) and dinosaurs. Uie questionable Cretaceous/Paleocene record from India (Gayet et al., 19841, Semionotidae are not known to survive beyond the 4. Taxa which occur in the EI Molino and/or Sant'a Lucía Cretaceous/Paleocene boundary. Their absence at Tiupampa is formations and, which based on records from elsewhere in the world. probably related to the Paleocene age of the Santa Lucía Forination. lransgresscd the K/T boundary arid arc of dubious or no value in The new genus from Aguti CI:m und Hokl Cordillern, known only defining this boundary. These include Lepisosteus (Lepisostcidae), by scales and referred to the Semionotidae, seems endemic to S Os teog I os s id ae , Cyprin i form es, Ch araci f or mes, i lur i forni es, Bolivia and northern Argentina where it is recorded in both marine Cypriiiodoiitiforrnes. Perciformes, Polypteriformes, Dipnoi. and fresh wakr environments. Leptodactylidne. Podocncniididac. Laccrtilia, Boidae, Madtsoiidae, Dyrosauridne. and Lcptictida. b. Osleoglossidue

5. Taxa recorded only in the Santa Lucía Formation which are the Living Osteoglossidae are restricted to freshwater environments in earliest or only records for these groups in South America. They do tropical South America, Africa and Asia. Some fossil remains were not occur in association with diagnostic Cretaceous taxa and reported from fresh water (Green River basin, Wyoming, Grande therefore are regakded as Paleocene. Included are Ceratodontidae n. 1984; Sumatra, Sanders, 1934; Australia, Taveme, 1979), some from gen. and sp., Gymnophiona, Aniliidae, Tropidopheidae, Sebecidae, brackish water (middle Cretaceous of Zaire; Taverne, 1976) and Peradectidae, Caroloameghiniidae, Microbiotheriidae, Didelphidae, some from marine environments (Paleocene of Zaire; Dartevelle & Hathliacynidae, Kollpaniidae, Pantodonta, Hyopsodontidae and Casier, 1959; Turkmenistan, Danil'chenko, 1968; Niger, Cappetta, Notoungulata. 1972, Gayet & Meunier, 1983; Eocene of Italy, Taverne, 1979; England, Woodward, 1901; Morocco, Arambourg, 1952; and As demonstrated by group 3 (above), apparent late Pretaceous Denmark, Bonde, 1966). In well known marine Cretaceous levels taxa occur only in tlie lower and basal middle members of the EI (Lebanon, Israel, Morocco, Italy, Portugal and Yugoslavia) where Molino Formation. It therefore appears that the K/r boundary occurs the ichthyofauna is abundant, no fossils of this order are represented. above the basal middle member, while the upper El Molino member Consequently, Osteoglossiformes are considered as primary may be regarded as early Paleocene based on palynological study of freshwater fishes following Nelson (1969) and Gayet (1987b). In a stratigraphic equivalent in norlhwestern Argentina (Quattrocchio el Bolivia, Ostcoglossinac wcrc reportcd in rill mcmbcrs of the EI al., 1986) and od the absence of diagnostic Cretaceous taxa. We Molino and in the Santa Lucia foniiat¡ons, wlicrcns' Phareodontinae therefore suggest &hatthe K/T boundary lies somewhere in the upper were reported in the Santa Lucía Formation only (Tiupampa). In the part of the middle member of the EI Molino Formation. In addition, El Molino Formation, they occur in association with marine fishes. tlie Sanla Lucla Formation, which also lacks diagnostic Cretaceous taxa and conformably overlies the early Paleocene upper member of c. Cyprinodontiformes the EI Molino Formation, may be regarded as early, but no earliest, Paleocene in age. Nothing can be said about the paleoenvironinent of the cf. Cyprinidontiforines in Bolivia, because this group is too poorly C. PALEOENVIHONMENT know II.

Pishes d. Polypteridae

Fishes have been divided into four main categories (Myers, 1938) Living Polypteridae are Ahican freshwater fishes as are all fossils based on their tolerance to fresh or salt water: 1) primary freshwater of this group known from Senoniai to Pleistocene. In Bolivia hey fishes found only in fresh water; 2) secondary freshwater fishes, were reported at Tiupampa (Santa Lucía Formation) and occur in found normally in fresh water, but capable of entering and surviving association with marine fishes at Vila Vila (lower EI Molino in marine water; 3) peripheral freshwater fishes, diadromous, which Formation).

420 e. I fresh water species. This family includes lhb gcnus Ccrr~roponrrrs from the coastal waters of the New World, a species wliicli enters Ceratodonts are generally associated with fresh water Brazilian rivers (Berra, 1981). Centropomidae are present at environments by extrapolation based on occurrence of extant forms. Tiupampa and Criadero de Loro in Bolivia. NeverUieless, ceratodonts occur in undoubtedly marine deposits in Zaire (Casier, 1961), Niger dnd Algeria (Martin, 1984), Mali d. Cypriniformes (Tabaste, 1963). Egypt (Schad, 1984) and Kansas (Schultze, 1981). According to Uiese different authors, these faunas may be interpreted Cypriniformes are living freshwater fishes and fossil remains as deah assemblages of marine, brackish and hesh water forms or attest, as far as is known, to a similar environment. Only that they were tolerant of a wide range of salinity. Ranrallicfitliys from Ein Jabrud (Israel) which may be considered as Lepidosirenids are restricted to fkesh water environments. an ancestral Cypriniformes (but is not a true Cypriniformes) is marine. Cypriniformes are reported in the lower and basal middle 2. Secondary freshwater rilies members of the EI Molino Formation in association with marine fishes. a. Lepisosteidae e. Characiformes Most of the living Lepisosteidae are found in fresh water, but one species may tolerate brackish or even marine conditions and is found Living Characiformes are freshwater fishes (only a few along Uie coast of the (Sutlkus, 1963). Bccouse of cxccptions enter brnckisli waters) while sonic fossils werc their Uiick scales covered with ganoine which are more resistan1 llinn apparcnlly adapted to brackish water (Cappetta el al., 1972; any oflier parts of thé fish, the fossil Lepisosteidae are often reported Gaudant, 1980). The oldest true Characiformes, Sulnrinops on scaks only, which may be transported far from their biotope (Gayet, 1985), was marine (Gayet, 1981). In Bolivia. without damage. In Bolivia. Lepisosteidne are always found in Characifornies are reported in the lower and basal middle association with marine fislies including selachians, pycnodonts and members of the EI Molino Formotion (Serrasalinidae, eotrigonodonts in the lower and basal middle members of the El My le in ae; C li ar aci d ae, Tetrago nop ter i n a e), i n t li e San t a Molino Formation, but they are always disassociated, indicating Lucía Formation at Tiupampa (Serrasalmidse, Myleiiiae and postmortem movement. In the only locality where no true marine S erras a I m in ae; C li ar acid a e, Tetrag o no p t er i II ae and c f. fishes were found (¡.e.Tiupampa), Lepisosteidae are absent but Ulis Rhoadsiinae) and in the Eocene age Cayara Formation absence may e the consequence of age (Paleocene) and not of (Serrasalmidae, Myleinae). In the El Molino Formation they environment. tepisosteidae had a nearly world-wide distribution occur in the same levels as marine fishes. during Cretaceous and early Tertiary times. Subsequently, they disappeared from all the continents except North America where f. Siluriformes they are now restricted to the southeastern part of the USA and Central America. I Most Siluriformes occur in fresh water. However; the Plotosidae are an advanced marine family which today occur b. Clupeidae around Auslralia and the Ariidae live along the coast or inter estuaries and coastal rivers. Tertiary Siluriformes were Gasteroclicpea branisui is a Clupeiformes endemic to the Andean reported in fresh water areas and/or in brackish water (see basin. Clupeiform,es are generally marine fishes with the exception .Gayet, 1991). In Bolivia, Andiniclithyoidea (3 families) are of some South American li.ving genera (RnnmoRnsrer, frisrignsrcr; known in the Siinln Lucln Forincilion nt Tiupiiinpn aiid Rlnnco p ol' G6ry. 1969; Cionc l'ercirii, 1985). III Uic middle Eocene of tlic Kiinclw (tlic ichtliyol'iiunii the liiter lociility is not yet well (Wyoming, USA), Kniglrtia and enough known to provide pnleoenvironmental interpretation); Diploniystus which are known to be marine, are reported in strictly they are not reported at Agua Clara or Hotel Cordillera where fresh water environments (Grande, 19820. b). Gaslerocfupea marine fishes are present and may be considered as branisui is reported at Tiupampa (Santa Lucia Formation) where no freshwater forms. Ariidae are reported everywhere in both marine fishes are found, and in some levels of the EL Molino formations with or without marine fishes. They are present at Formation (at Agua Clara) in association wih marine fishes. In some Tiupampa, but are few in number. At Hotel Cordillera they green levels at Agua Clara, Gasferoclupea is found in are very numerous and occur in association with a few association only with brackish-water ostracods. Consequently, marine taxa. Gasteroclupea may be considered, with some confidence, as a secondary JÌeshwater clupeid. 3. Marine fislies

e. Centropomidae ' a. Selachians

The family Centropomidae, as defined by Greenwood (1976). is Selachians are, considered as marine fishes. Nevertheless, one composed of marine and fresh water genera. Lares, to which the family of American batoids (Potamobygonidae) lives exclusively in Bolivian genus may be compared, is known to include marine and fresh waters. One pristid fish and a shark can also enter rivers or

421 MIKI?ILLI? tiAYE1. LARRY U. MAWIIALL&'I'IIIBKKY SBMI't!KI!

Inkcs, but it is iniportnnt to note thnt these elasinobranch fishes are More difficult is lhe interpretation of tlic ichtliyofaunn from the EI marine species which temporarily enter fresh water. According to Molino Formation. In nearly all tlie localities. niarine and fresh water Cappetta (1991), il is highly probable that the rocks containing fishes occur in association and, as noted above, bones, teeth and selachians in Bolivia were deposited in marine environments. scales are never found articulated. It is probable hat the calcareous fossiliferous levels of the lower member of tlie EI Molino Formation b. I'ycnodontiformes at Agua Clara were deposited in or near tlie sea (lagoon or coast). Other levels at the same locality (e.g. some green mark) were Pycnodonts, a purely fossil group, are considered to have been probably deposited farther from the sea (perhaps without any marine fishes (Cione 8c Pereira, 1985). They were flat and high connexion with it). Gusferoclupeu,which has been found articulated rishcs that apparently swam slowly along the coast and were linkd in these levels. has also been rcportcd at Tiupampa and is possibly a to the substratum for their alimentation. Some could probably enter freshwater genus. Nevertheless, this does not dismiss the possiblity waters of lesser salinity. Wenz (1989) noted that the Brazilian lower of a brackish water habit for this genus. Cretaceous pycnodontid Ieninnja may have lived in fresh water. Traquair (19 1O) thought that pycnodonts found at Bernissart Camoin el al. (1991) concluded that tlie EI Molino rocks were (Relgium) with thc rcptilc Igrrunodon were living in fresh water deposited in a conl~ncntalenvironment. while Gayet ef al. (1992) without any connection with the sea. Nevertheless, these examples and Sempere (in press) noted that marine fish taxa occur in these are sporadic and pycnodonts are typically found in more or less open sedimentary rocks and discuss their depositional environment., marine environments in rocks of Jurassic and Cretaceous age (Hake1 and Ht1jula in Lebanon; Ein Jabrud in Israel; Jebcl Tselfnt in IIiglier ver1el)rules Morocco; etc.). In Bolivia. pycnodonts were reported in the Chaunaca Formation (only one tooth of no paleoenvironmental Crocodiles of the family Dyrosauridae generally occur in siiallow significance) and in,the lower and basal middle EI Molino. niarine near-shore deposits, but can exist in fresh water as well (Bufretaut, 1982). The ainpiiibians. lizards. snakes. turtles. otlicr c. "Snlinoniformes" families of crocodiles and mammals all indicale terrestrial and/or fresh water paleoenvironmcnts. "Salinoniforines" are a paraphyletic group present in all kinds of e environ in n t s (Bkrr a, 19 8 1). Nevertheless, Encho don tid ae D. FALEOBIOGEOGRAPHY (Enchodus) and Ichthyotringoidei (including ?Apareodiis) are known only in marine rocks (Goody, 1969). In Bolivia, Numerous vertebrate grou'ps in the late Cretaceous and Paleocene "S almoniformes" were reported in the lower and basal middle of South America are also recorded in North America, and these members of the EI Molino Formation. faunal "links" were used by paleontologists to postulate that a trans- Caribbean land bridge existed during part or all of this time interval d. Tetraodontiformes (see Gayet el al., in press; Marshall & Sempere, in press and references therein). These concIusions have recently been One living family of tlie order Tetraodontiformes, the corroborated by geodynamic data (see Pindell ef d.,1988; Stephan Tetraodontidae with about 90 species, has a few fresh water el ul., 1990). It now appears that the began its east- representatives in South America, Africa and India (Berra, 1981). northeastward migration in Campanian time. The now submerged However, Eotrigonodontidae is a fossil family so far known only in Greater Antilles and the Aves Ridge formed a subaerial volcanic arc marine deposits. Sfephuriodrrs has only been fouiid in Iate established 'on the northe:istern edge of the Caribbean plate during Cretxeous marine scdiincnts. In Bolivia. the cotetragonopterid tlic late Crctaccous and early I'aleocciic. It is likely hat this arc was Sfephanodrts as bdcn rcportcd in the first two rneinbcrs of the EI the land bridgc that providcd the pathway for dispersal of continental Molino Forination. (and fresh water) vertebrates. However, the land bridge provided by the Aves Ridge was apparently strongly dependent on tlie inaginatic In conclusion, ye note that at Tiupampa and Criadero de Loro and tcctoniC activity of this arc. and on sen level changes. Maginatic- (Santa Lucia Formation, Paleocene), no' true marine fishes tectonic quiescence and/or marine high-stand periods would have (selachians, pycnodonts, Salmoniformes and Eotrigonodontidae) promoted relative subsidence of part of the isthmus below sea level. have been reported. All of the primary freshwater fishes reported in In view of the probable fragility of the Aves Ridge connection, Bolivia, with the exception of Semionotidae (Lepidofes) which do faunal interchange is likely to have occurred in several "pulses" not cross the Cretaceous/Paleocene boundary, are present at these beginning in the Campnninn until the land bridge finally disappeared localities. Although some Osteoglossidae may occur in marine in late Paleocene time as is suggested by paleontological data. This waters (i.e. Brychaerrrs), the subfamily Phareodontinae is known thus "pulse-IiypoUiesis" would have permitted opportunities for endemic far only in fresh water levels. Tiupampa is Uie only Bolivian locality evolution of first-wave immigrants and for subsequent dispersal where nearly complete skulls (Osteoglossiformes, Siluriformes and during a later "pulse". For this reason, the direction of dispersal of Characiformes) were found. In all other localities bones are always most groups is not certain (Marshall & Senipere, in press). separated, attesting to postmortem movement. Thus, based solely on Some vertebrate groups which participated in this interchange(s) the ichthyofauna. we can confidently conclude that the Tiupampa include dinosaurs (Avisauiidae, Ceratopsidae, Hadrosauridae, and Criadero de Loro fossiliferous sediments were deposited in a Titanosauridae), snakes (Anilioidea, Boidea). possibly lizards fresh water environment. (Teiidae) and sliore-dwelling turtles (Bohremydidae) (Gayet el al.,

b. 422 in prcss; MarslisIl & Senipere, in prcss und references thcrcili). CONCIJUSIONS Aliiong fishes arc the Lepisosteidne (Lepisosteus), Osteoglossidae (I'hareodontinae) and possibly Cypriniforines and Ariidae (Gayet el This paper is the first attempt to provide a detailed overview of he al., in press). Mesozoic and Paleocene vertebrate faunas of Bolivia within Uicir Multiple taxonomic similarities among mammals (10 examples) stratigraphic context. Much of the inforination presented here is new, "link" late Cretaceous and/or Paleocene faunas in North America and includcs data obtuiiicd as a result of IIII iiitciise palcontological with early and middle Paleocene (Tiupampian and Itaboraian, program started in 1981 (M. G., L. G. M and others) and of geologic of respectively) faunas in South America. Among marsupials, studies initiated in 1984 (T. S. and others). Most the fossils are Peradecles is known in Tiupampian faunas of Bolivia and Peru, and still being described and it will be many years before monographic is present in early Paleocene faunas of North America; publications can be completed. In addition, magnetoslratigrapliic and caroloameghiniids (Glnsbirts) are present in tlie late Cretaceous of isotopic (Ar/Ar, K-Ar) studies of the El Molino and Santa Lucía North America and Tiupampian (Robertlioffstetleria) of South formations and isotopic and strontium analyses on fishes Ne Anierica; Didelphidne are present in the early Eocene of North in progress, and cornpletion of this gcochronologic program will Anicric:i and Tiupampian of South America; and Pediornyidae (Iate hopefully perinil secure calibrntioii of these rock units witti tlie Cretaceous of North America) are a sister-group of geologic time scale. Synthetic overviews for the of Microbiotlieriidae (first recorded in the Tiupampian of South Argentinian, Chilean and Peruvian rocks in the Andean basin are Anierica). Among placentals, a Leptictida (cf. Ciniolesres) is known now urgently necdcd to permit refined correlation with those in from Tiupampa whileI Ciniofestes occurs in the late Cretaceous and Bolivia. carly Palcoceiic of North Anicrico; I'niitodontn (Alcidedor Digt~ya)at Dcspile tlic fact tlint his is by iicccssity only ii "progrcss rclwri", Tiupampa are first known in the middle Paleocene in North many advances in knowledge have been made 011 the Bolivian America; mioclaenine hyopsodontid Condylwtlua at Tiupampa are Mesozoic-Paleocene scqucnce during the last few years, and in this fist recorded in early Paleocene faunas of North America; Xenarlhra study we have been able to clarify some debated issues. The most (Dasypodidae) first appear in Itnboraian faunas in South Amcrica iinprtant of thcsc is thc apparcnt, although still ill-dcfincd, location the and in the late Paleoccne in North America; aid which of boundary and its position relative to Uic inanmal-bearing first appear in the late Paleocene of North America are regarded as a level at Tiupampa. Within the El Molino Formation, undoubted late sister-group of Xenun data which occur in the Itaboraian of South Cretaceous taxa occur only in Uie lower and basal middle members. America (Marshall & uizon, 1988; Marshall & Seinpere, in press; It therefore appears that the K/T boundary lies somewhere in the al., L Gayet ef in press and references herein). upper part of the niiddlg member, while the upper member may be Although latest Cretaceous and Paleocene interchanges are well regarded as earliest Paleocene based on the absence of diagnostic docuiiiented, it is of interest to note that several terrestrial groups of Cretaceous taxa and on Uie palynological studies of a stratigraphic vertebrates which were often rather widely distributed either in the equivalent in northwestern Argentina. The EI Molino Ihus appears to Laurasirui realm or on Gondwanian continents in the late Cretaceous range from latest Cainpanian to earliest Paleocene, while the and Paleocene, never crossed the Caribbean nor Tethyan areas. disconformably overlying Santa Lucía Formation appears 'to be Acipenserid, polyodonlid and gonorhynchoid fishes, as well as early, but not earliest. Paleocene. Thus, confra Van Valen (f988), palaeobatrachid frogs, baenid turtles, and anguimorph lizards the EI Molino dinosaurs were all late Cretaceous (apparently-early inhabited North America during the Cretaceous; hiodontid and Maastrichtian), while the land mammals from the Santa Lucía esocid fishes were present on that continent during die Paleocene. Fonnatioii at Tiupanpa were Paleocene. h None of these northern groups rdached South America. Conversely, polypterid and characifonns, as well as pipid frogs and araripeniyid ACKNOWLEDGMENTS turllcs of Gondwunirin origin. known in the Crctnccous of South Anici ¡cil. did not rcricli Noilli Anicricli. Tlic iilx~vclisted vcrlcbrnlcu ~sl)~~~sur tilis sttltiy WCIC su1)l")ltcd I)Y rillitis fro111 IIIC cclltlc rank aniong tlie most signiliciiiit ones that did not tuke part' in the late National de la Keclicrclie Scientifique (UKA 12, Institut de Cretaceous and Paleocene interchange(s). Other groups could be PalCoiitologie, Paris, 1981); ari "Action Spécifique" (1983 to 1989) added to this list but, for the time bcing, they cannot be considered to from the MusCuni National d'Histoire Naturcllc, Paris; National be very conclusive bccause their distribution is still inadcquately Geographic Society (2467-82. 2908-84, 3381-86); URlH of Orstoni; known. the Gordon Barbour Fund. Department of Geological and The failure of various groups to cross the Caribbean area would Geophysical Sciences, Princeton University; and the National apparently favour the hypothesis of a discontinuous terrestrial route Science Foundation (EAR-8804423). Logistic help was given by or even an island hopping hypothesis. But the ascertained dispersals Orstom, YPFB and VARIG. Fossil collecting was carried out under of large-size as Titanosauridae and of freshwater fishes the auspices of the Asociación Boliviana de Paleontología and (Lepisosteidae, Phareodontinae, and perhaps Ariidae) are stratigraphic work under the YPFB-Orstom research prcgram on inconsistent with such possibilities as they require a continuous land Bolivian . Thanks to L. Barrios, J. Blanco, F. de Broiii, (fluviatile for the fishes) route. These failures could result from E. Buffetaut. G. Camoin, H. Cappetta, R.' Céspedes, A. L. Cione. climatic, faunal interactions (niches occupied by ecologically J. Dejax, D. Dingerkus. E. Jaillard, M. Suárez, R. SuBrez, J. vicarious autochthonous forms) or shortness of duration of the F. J. Meunier, Oller, J. C. Rage, B. Sigé, C. de Muizon and S. unbroken nature of the land route. Wenz for help in aspects of Ulis study. Figures 1 to 4 were drawn by E. Liebmm.

423 BAEZ, A. M. 1987. The lntc Crctnccous fauna of Los Alamitos, ,' Patagonia, Argentina, Part III - Anurans. Revista del Museo AGASSIZ, L. 1841. On the fossil fishes found by Mr. Gardner in Argentino de Ciencias Naturales "Bernardino Rivadavia", the province Cearl, in the north of Brazil. Edinburgh New Paleontología, 3 (3): 121-130, Buenos Aires. Philosophical Journal, 30 (59): 82-84. Edinburgh. BAEZ A. M. & S. PERI. 1989. Baurubatrachus price;, nov. gen. et AHLFELD F. & L. BRANISA. 1960. Geología de Bolivia. Instituto sp., un Anuro del Cretlcico superior de Minas Gerais, Brasil. Anais Boliviano del Petróleo, p. 1-245, La Paz. da Academia Brasileira de Ciências, 61 (4): 447-458, Rio de ALBINO, A. M. 1986. Nuevos Boidae Madtsoiinae en el Cretlcico Janeiro. tardío de Patagonia (Formaci611 Los Alamitos, Río Negro, BARDACR, D. 1961. New Tertiary teleosts from Argentina. Argentina). IV Congreso Argentino de Paleontologia y Anierican Museum Novitales, 2041: 1-27, New York. 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G., C. de MUIZON & B. SIGE. 1983a. Late l'Académie des Sciences, sdrie 2,304 (13): 771-774, Paris. Cretaceous mammals (Marsupialia) from Bolivia. Geobios, 16 (6): MUIZON C. de & L. G. MARSHALL. 1987c. Deux nouveaux 739-745, Lyon. condylarthres (Mammalia) du Maastrichtien de Tiupampa (Bolivie). MARSHALL, L. G., C de MUIZON & B. SIGE. 1983b. Comptes Rendus Hebdomadaires des Sbances de I'Acadkmie des Perutlierium altiplanense, un notongulé du CrdtactS SupQieur du Sciences, serie 2,304 (15): 947-950, Paris. G. PBrou. Palaeovertebrata, 13: 145-155, Montpellier. MUIZON C. de & L. MARSHALL. 1987d. El alba de la edad de MARSHALL, L. G., C. de MUIZON, M. GAYET, A. LAVENU & los mamiferos en . N Congreso Latino-americano de B. SIGE. 1985. The "Rosetta Stone" for mammalian evolution in Paleontologfa,Actas 2: 757-791, Santa Cruz. South America. National Geographic Research, 1 (2):274-288, MUIZON C. de & L. G. MARSHALL. 1988. 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