gayet+marshall+sempere 1991 mz-pc vertebrates of bolivia

8/10/2019 Gayet+Marshall+Sempere 1991 Mz-Pc vertebrates of Bolivia

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with the PirainboiQ Formation of Brazil and tlie lower part ofTacuarembd Forinationof Uruguay (Seinpere Oller, 1987).

The lower boundary of the Serere superscquemce isa diachronousunconformity excepti n the southern S ubandean belt w here,tlie oldestand elsewhere unknown Ipaguaz Formation overlies the latePermian-early Triassic Vitiacua Formation with only a slight, non-erosional, lithologic discontinuity (Sempereet al., 1990, 1992). In

other areas of the basin, younger units of the Serere supersequenceunconformably overlie Permianor olde r strata, reflecting onlapping.deposition on an altered, eroded and/or deformed substratum. Theupper boundary of the Sere re supersequenc e is the surface that marksthe "Condo event" (see above). This unconformity shows differentcharacteristics which relate to its location and position in the IahstJurassic-Paleocene basin.

In suinmary, tlie middle Triassic-Jurassic geologic evolution ofBolivia involves: 1) an initial rifting process with deposition ofr e dbeds in narrow troughs, locally with gypsum and halite (middletoIate Triassic); 2) termination of rifting and onlapping fluvio-eolianscdinicntntion(late Triassicto earliest Jurnssic'?); and3) extensiveerg development (early and middle Jurassic'?) (Oller Se m pr e,1990).

II. ,h l lSS l J U t ASS I 'IO l'A I,EO CICN C(I'ucu supersequence)

Classic works on the Bolivian "Cretaceous" Puca Group are byLohmann Branisa (1962).Russo Rodrigo (1965), KrizClierroni (1966). Lohmann (1970), Reyes (1972) and Cherroni(1977).

The "Cretaceous" sequence stratigraphy of Bolivia and Peru iscurrently being re-evaluated and synthesized (Jaillard Seinpere,1989; Seinpere,in press). The Puca supersequence (Scmpere, 1990)is bound below by the kO discontinuity( ; .e , he KimmeridgianAraucan-"Condo" event at about144 Ma; Cow ie Bassett, 1989).

and a bove by tlie k5 discontinuity which records the functional onsetof western Boliviaas a continental external foreland basin of thepaleo-Andes (Sempereet al., 1989). The age ofW is close to thePaleocene-Eocene boundary(53 Ma; Cow ie Bassett, 1989) andcould be as young as 50-51 Ma (Senipere, 1990; MarshallSenipere, 1991).

In this pnper w e, divid e the Pucn supersequence into threemegasequences. The Puca A megasequence includes the partlyequivalent Condo, Kosmina,La PuertaS.S. Sucre and Tarapayaforinations (Fig. l), nnd consisLs of nzoic non-marinedcposits; theniiuinc Miraflores Fdrntntioii constitillcsthe I'ucn B nicgnscquc~icc;and the PucaC megasequence include's the Aroifilla, Chaunaca, EIMolino, Santa Luca and Impora formations, and their stratigraphicand teniporal equivalents (Fig. 1). Unlike the Puca A, the Puca B andC have yielded many marine (Puca B and C) and/or continental(Puca C ) fossils.

The marine Miraflores Formation consists of fossil iferouslimestones and minor mudstones, up to28 m in total thickness; itisof Cenomanian and Turonian age (Jaillard Sempere, in press).Withinh e Puca supersequence, the Miraflores Formation is the onlyunit of undisputed marine origin because it contains ammonites andechinoids (B ranisa, 1968; Reyes, 1972). It is separated from theoverlying Aroifilla Formation byan erosional discontinuity whichis

accoinpanied by synscdinientnry tcnsioniil tectonic nnd mngiiiornnnifestations (Sempereet al., 1988; Seinpere, is press) and istermedt he "Vilcapujio event"in Bolivia (Ctihvcz, 1987)or k3 in tilecentral Aniles (Jail lard Se mp ere , unpublished). Thu s, thMiraflores Formation stands as a specific "hinge" unit in thBolivian latest Jurassic-Paleocene stratigraphic sequence.

Mainly red mudstones with minor sandstones and evaporite

(gypsum, anhydrite, halite) were deposited between discontinuitk3 andk 4 i n f luv ia l and l acus t r ine env i ronm ents [Aro i f i l laFormation and m ost of Chaunaca Formation (Sem pere,in press); thestratigraphically equivalent Torotoro Formation consists mainly red sandstones of fluvial origin]. Two greenish levels, which consof very fine sandy, green to blackmark, black to grey laminatedlimestones and yellow dolomites, occur in the Chaunaca FonnatiThe foriner is moreconspicuous, traditionally marks the base of theunit, and is kr m ed the "basal limestone" of the Chaunaca FormatioThis30 m-thick sub-unit comm only yields abundant bivalves(Mytilidae,Brochidontess p . ) which have been interpreted toindicatean intertidal cnvironnient(Briutisn ct al., 19GG). How ever,IIrestricted mnrineorigin for the "basal limcstonc" is now questioncd(G. amoin, personal coinunication). IdenticalBracltidoort/essp. arerecorded froma limestonc level in the LoinasNegras Formntion ofnortlicrn Chile (Mnrinovic Litlisen, 1984).The "bnsitl linlcstonc"of the Chaunaca Formation yielded a palynological assemblage oSantonian to earliest Campanian age (Prez, 1987), and may partly equivalent to the Snntonian age shallow-marine limestonknown from the Querque and Omoye formations of southernPeru(Jaillard Sem pere , 1989). The Aroifilla and Chaunaca formatioare equivalent, respectively, to the lower and middle VilquechiFormation of southeast Peru (Jaillardet al., in press) (Fig. 1). Theupper greenish level of tlie Chaunaca Formation is equivalentto theuppennost green level of the middle Vilquechico (Jailllardet al., i npress ; Sem pere , in press) , which has yie lded the se lachiaSchizorhizaslronieri .Thus, i ts age is not older than niddle

Campanian (Jaillardel al., i n press).The EI Molino, Santa Luca and Impora formations occur betweediscontinuit ies k4 and k5i n Bolivia (Fig.1).The EI MolinoFonnation is equivalent to the Lecho, Yacoraite, Tunal and Olmeformationsof northwestcrn Argentina (Fig. 1); to the Areniscas deAzcar, Vivinn and Cnchiyacu fornialions of SubnndennPeru(Sempereet al., 1987: Jnillard Sem pcrc , 1989);to thc upperVilquechico Formationof the Peruvian Altiplano (Jaillardet al., i npress) (Fig. 1); and to the Estratosde Quebrada Blanca de Poquis,niid possibly parts of thcLoinns Negras.Pnjonalcs and Tonclformationsi n nortlicrn Cliile (Gordcwcg8c l


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FOSILES YFACIES DE BOLIVIA - VOL. I VERTEBRADOS

AGE

(Ma)

4 -

0 -

0 -

- 0 -

- 0 -

00 -

- 1 10-

-120-

-130-

-140-

-150-

-160-

-170-

-180-

-190-

- 00 -

-210-

- 20-

-230-

-240-

GEOLOGIC TIME SCALE

Haq e l a1.,1987)

O W9 82066.5

u3

w

QI- un

131

ouu

QU

3

2 1 0

I

5

uu

nI-

Danian

Maaslriclilian

ConlaclanTuronian

Cenoinnrii:rii

A l b i a n

Aptlan

Liass ic

Keuper

ANDEAN DOMAIN SUBANDEAN BELTNORTHWEST SOUTHEAST

ARGENTINA PERU

Cajones ~

Yantata

lchoa lchoa

Figure 1. Clironostratigrapllic chart show ing vertebrate-bearing formations (*) of middle Triassic-Paleocene age in Bolivia,and inadjacent Argentina and Peru.

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

Rased on recognitionof large-scale transgressive-regressivesequences, theEI M o h o Formation is divided into lower, middleand upper membcrs which areof stratigraphic value (Jaillardel al.,in press; Jaillard Sem pere, unpublished; $empe re, in press).Common sedimentary facies in the El Molino Formation include:mudstones and mark (green to blackor purplish, reddish); grey tolight brown limestones (micrit ic to bioclastic and/or ooli t ic);

stromatolitic boundstones referred to asPiicaliiltusby Steinmann(1923) and Ahlfeld Branisa (1960); f ine to medium-grainedsandstones, usually calcareous and l ight-colored; some yellowdolom ites, ofte n with algal laminations; very rare evaporites(gypsum and halite casts); anda few white altered tuffs of porcelainaspect. Mudstones, mark and thin limestones largely predominate inthe basin; sandstones, dolomites and evaporites are more commonalongits borders; an d tuff levels a re thicker and more frequent inthewest,i.e.in the directionof the active volcanic sources located inpresent-day C hile a nd Peru. In the basinal localities, lime turbiditesare comm on and wher e fossiliferous the paleontological materialswere dcpositcd a stibstaiitial distancefrom their life environinent.

Coilcordant palcocurrciit data iiidicate that the basin deepenedtoward the northwest in Bolivia (Sempereer al., 1987; Seinpere, inpress), and probably connected with a marine basin in the PeruvianSubandcnii region. Aiiiiiioiiitcs tire reported2 in bclow thetop of theCachiyacu Foriiiationi n the HiinIlaga aren (7"s)of iiorthern l'cru(Vargas, 1988).In Bolivia, several fish groups, which reputedlyindicate a marine environment (see below), are known from thelower and basal middle El Molino. Dinoflagellate cysts are present inat least so me levelsof the formation in Bolivia and in stratigraphicequivalents in northwestern Argentina (Industry, unpublished; J.Oller, personal communication). Ammonite embryos have beenreported from tlie lower El Molino (Sempereef al. 1987), but theiridenti f ica tion needs conf i rma t ion. Foraminifera (Mil io lidae ,Discorbidae) are presentin many levels aid facies (G. Tronchetti,inCamoinet al., 1991), but no detailed study has yet been made oftheir taxonomyor paleoenvironrnental implications. Molluscs arereported from various localities(e.g. at El Molino; Pilsbry, 1939),although the systematicsof these taxa is outdated and the relevanceof described species has yet to be established. Echinoids areapparently lacking and the proportion of brackish water ostracods islocally high (J.F. Babinot.in Cainoinet al., 1991).

A pctrogrnphic studyo f the Yacoraite Forinationof northwesternArgentina led Marquillas (1985) to interpret i ts deposit ionalenvironinen t as a res t r ic ted ca rbonate bas in. An ol igohal ir ielacustrine environm enthas also been proposed (Camoinef al.,1991) . However, pa leon to log ica l da ta do sugges t tha tcommunication of the basin with an open marine realm was frequentduring deposition of theEi Molino Formation, but that the basin

itself was never openly marine, at least in Bolivia and Argentina(Gayetet al., 1992, Seinpere,i n press). For reasons discussedbelow, we believe that most of the El Molino Formation wasdeposited principally ina restricted marine environment whichconnected wi th a more typical ly open marine realm locatednorthwe st of Bo livia (see also Se mpere, in press),

The overlying Santa Lu ca and Impora formations were depositedin fluvial to lacustrine environments. N o marine fossils or facies areknown from these units which consist principally of red to purplishsiliciclastic sedimentary rocks. Gypsum and "gypsifred" anhydrite (J.

M. Roucliy, ,personal conimunicntion), are abunda nti n the iippcrpart ofUie Santa Luca Formatioli near Potosi. So me green maris anlight-colored limestones occur in theImporaFormationof southcriiBoliv,ia.

The Santa Luca and Impora format ions are , respect ivelystratigraphic?lly equivalentto the'Paleocene age Mealla and MazGordo formations of northwestern Argentina (Maroccoef al., 1987)

(F ig . 1 ) . The M ea l l a Format ion can b e cor re la t ed wi th thTiupampian (early Paleocene) Land Mammal Age based on iequivalence w ith the Sai?ta Luca F orma tion (s ee below) and Maiz Gordo Formation with the Riochican (late Paleocene) LaManimal Age based on palynology (Volkheimeref al., 1984) andstratigraphic position (Pascua1et al., 1981).

VERTEBRATE FAUNAS AND LO CALITIES IN BOLIVIA

A. TACURU GROUP

Alilfcld Brnnisa(1960) rcportcd rcinniiisof lurgc bones,possibly of dinosaurs,from the Tacur Groupin the Serrana deMandeyapecua. southeastern olivia.III his area the Tacur Groupincludes rocks of middle Triassic and Jurassic age, naiiiely tlTnpccun, Cnslcll6iiaiid Ichoii forlnntiolis (Fig.1).

U . CASI 'ELLON FO RM AlIO N

Scales and bones of semionotiform fishes were found inthe lateTriassic-early Jurassic age CastelMn Form ationon the western flankof the Charagua anticline in tlie Quebrada de Charagua,7 km west ofthe townof Chbagu a (Wenz,in Go i Hoffstetter, 1964). Nothingdefinite can be said of these bones, althougha study of the scaleswith Scanning Electron Microscopy (Gayet Meunier, 1986) dndoft he i r t r ansverse sec t ions pe rmi t s r e fe rence to the f ami lSemionotidae, probablyLepidofes sp. (Gayet, 1991).

C . MIRAFLORES FORMATION

The Miraflores Formationis a marine lim estone unit which, basedon knowledge of ammonites, bivalves, gastropods and echinoiwas regarded as middle to late Ccnoiiinnianin age (ranisaet al.,19GG; Jail lard Scmpcrc,1089), :itthough detailed scqucnccsthtigrapliy now suggcsts that the entire rock unit spans Cenoinaand Turonian time (Jaillard Sem pere , in press).Nearthe localityof Macha.90 kni north-northwestof I'otos,L. Branisa found minutepharyngeal tee thwhich are tenta t ively ass igned to cf .Cyprinodontiforines or Pycnodontiforrnes (Gay et, 1991).

I). AROIFILLA FORMA'I'IONTh e con tinen ta l Aro i f i ll a Format ion over l i e st h e mar ine

Cenomanian-Turonian age Miraflores Formation and underlies Santonian-late Campanian age Cliaunaca Formation; it is thereconsidered Coniacian and possibly early Santon ian in age (Fig:l).series of four footprintsof a large quadrupedal dinosaur werediscovered bythe Brigada10 of YPFB at Caleras,4 n below the"basal limestone" of the Chaunaca Formation. The footprints wimprinted in soft sediment and are otherwise indeterminate.

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FOSILFSY FACIESDE B o L I m- VOL.I - VERTEBRADOS

P E R U ;I

15

17

,I4

t

.LA PAZ

*ChuIlDa Khasa

100k m

.COCHABAMBALa Cabana-

-Rio MoileParotanp a , c ~ s ~ ~ i p dst. Blanco Rancho USANTA CRUZ

ORURO. Arapampap d i l a Vi ' a DE LA SIERRALag o Uru Uru Torotoro-. Tiupampa

19'

*Chaup-Khocha

: *Cale r ias

*Tambo *CamargoSalar de Uyi ni Color adoI

21

C H I L E

*Chocaya

Chaupiuno

Rancho Hovada-. UTARIJA

Serpa \ *Villa Pacheco

/-

e c h a r a g u a ,f/

tSerrania ,

eManieyapecua I

,

i.

I

PARAGUI'\

\

\\

\

69" 67 65" 63

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

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Y

MIRIIILLEGAYEI'. U R R Y G. hIARSIlALLL TIUERRY SEMI'EKI

IC C l l A U N A C AIWI


7/42

CerroL a m a C h a q u i

180 To r o t o r

Cruz Khasa

' Ikm

Figure 3. Detailed map of tlie Torotoro area (for location, see Fig. 2) showing vertebrale localities in the CI Mol ino and'Sa nta Luciaformations. The number s 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, serief i731 , 1/50,000, 1968 edition, Instituto Geogrhfico Militar, Ln Paz 111-68.

siluriforin (Ariidae,Rliineastessp.) of verylarge size (morethan 2 inin total length). Vertebrae ofL e p i s o s f e u s sp. found in associationwith the Siluriforinesare also larger (morethan 3 cm-long) thanthoseof Agua Clara(about 1 m-long). As insome levelsa t AguaCI:lra, thc presenceof sclaclii:ins, pycnodonti forms, tetradontiforinsand "saliiioniforins"Enchodirs sp., ?Apateodirssp.) testifies to someconmu nication to asea (Gayet, 1991; Gayetet al,1992).

Ten meters below the calcareous leveljust described, a restrictedconglomeratic unii was recently found with reinaiusof Siluriformes(Ariidae) and Teleostean indet.A little farhe r, atthe crossingof theQuebrada SaytuJoWiu with the road, are grey-green mudstones ofthe upper EI Molino wilh remains of Clupeiformes(Casteroclitpeabranisai)andostracods (G ayet, personal observation).

At Vilcapujio (=Wila Apacheta), at abouti 140 northwestofPotosl on Uie Potosl-Challapata-Oruro road, is abone bed probablybelongingto the basal middleEI Molino which yielded fishes

(Pycnodontiformes; Characiformes, Serrasalmidae, MyleSiluriforines, Ariidae,Rliineastessp . ) and reptiles (turtlesandcrocodiles).In the vicinityhave also been found several levelswithholostean Ginglyinodi (Lepisosteidne), teleostean Clupeifo(Clupeidae,Castcroclicpea Invnisai),Chnraciforines (Serrasahnidae,Myleinae). Siluriforines (Ariidae,Rliineastes sp.), and/or reptiles(crocodiles, turtles).

At Sevaruyo.south of LakePoop6. about 120 kin west-norU westof Potos, are reportedtwo levels with remains ofCasteroclitpeasp.(Branisaet al.,1964).The first level apparently belongs toUie lowerEI Molino. Because of the large.(double) sizeof the hypocoracoidsfound in the second level relativeto .CasteroclupeaSholotype,Branisaef a l .(1964) questioned theageof this level. However,hypocoracoidsof very large size have also been found at Agua Clwherea transitional series exists that includes all sizes recordeatotlier localities.

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MIREILLE CIAYET, LARRY CI. M R S I U L L 11 UliKKY SBMIEKB

2. Uyuni Region

At thc locnlitics of Jay-Jay, Calnzayaand Tomave, rcspcctivelynorth-northwest, e ast and northeast of Uyuni, were found remains ofS i lu r i fo rmes (Ar i idae ,Rlrineasressp . ; Tomave on ly ) andClupeiformcs(Gasreroclupea branisai)by the$rigada10 of YPF(specimens inCTP, Santa Cruz). At Tambo Colorado, squtheast of

Uyuni, was founda selachian(Pucapristissp.) associated withgastropods(Melania potosiensis;specimens in CTP, Santa Cruz).With the exception of Calazaya which is upperEl Molino, thestratigraphic position of fossils at the other localities have not yetbeen established.

3. Southern Region

In theRoAngosto, 9 kin south-southwest of Chocaya, holosteanGinglymod i (Lcpisosteidae,Lepisosteussp . ) , Semionotiformes(Sem ionotidae, nov,. gen.), teleostean Clupeiformes (C lupcidae,Gasteroclirpea brunisni)and Siluriformes (Ariidae,Rltineasressp.)wcrc recovered froni,the owerEI Molino.

In Quebrada Tcxisca,1 km west of Rancho Hoyada along the.RoSan Juan delOro, 29 kin north-northeastof Tojo, were recoveredrcninins of l ioloslcnii Iyciiodontiforiiics (Pycnodontidac.indct.),Ging ly inod i (Lcp i sos tc idae ,Lepisosrenss p . ) , te leosteanOsteoglossiforrnes (Osteoglossidae, indet.), Siluriformes (Ariidae,Rhineastessp . ) and Te t raodon t i fo rmes (Eo t r igonodon tidae ,Sfephunodirs nii nk irs )(Gayet, 19 91) from the lowerEIMolino. Thesa ine Pycnodon t idac , Os teog loss idae , Ar i idae andEotrigonodontidae, plus two new selachians(Dusyurisnov. sp. 2 et3; Cappetta, 1991) anda crocodile toothwere recovered from thebasal middleEl Molino at the sam e locality.

Fish remains, includingPucapr isfis brunisi,have also been foundby J. Blanco (YPFB, Brigada 10)nearSerpa about35km south ofTupiza, and dinosaur trackwayswereobserved by Leonardi (1981)near Camargo,100 kin north of Ra ncho Hoyada.

4. Cochabamba Region

At Torotoro, aboui 95 kin south-southeast of Cochabam ba, severalfossil lcvcls NC known(Fig. 3). rom thc iniddlc partof thc lowcrEIMolinosonic 3 k in soulti-soutlicnst of Torotoro, Cnppctla (1975)dcscribcd numerous sclachians [Sclerorliyncliidae,fitcapristisb rn is i, Iscliyrhiza Irnrtenbergeri;Dasyatidae,Dasyaris branisai,D.niolinoensis,D. schnefleri;Khoinbodontidac,Pu ca ba h Imffstelleri(Cappe t t a , 1987)J and Bro in- ( 991) recorded tur t les(Podocneinididac,?Roxoclielysp.).

In a higher leveli n the same inember, on the northeast sideofTorotoro, numerous dinosaur trackways were found that have beencallcdthe pistadc danzas (Fig.3). t lcast five types of dinosnursare represented, includingsauropods, tlieropods and ornithopods(Bran i sa , 196 8 ; Leonard i , 1981) ; one t r ackway namedLigabueiclrniunr bolivianrtnrrepresents an ankylosauror ceratopsian(Leonardi, 1984).

Nearthe baseof the lower EI Molino at Unia Jalanta, close to Uiccave entrance, about5 kin west-northwest of Torotoro, occur somesmall tridactyle (Coelurosnuria?) footprints and isolatedbones ofturtles(?Roxuclre[ysp.). At Cerro Llania Chaqui, about4.3 kmeast

of Torotoro. occurscvcrnl scts of trackways ofa sinalltheropoddinosaur (Coclurosaurin?). Thesiinic typc of trnckway occurs ncarUic top of t l ic lowcr EI Molino on thcp t l ~ o Ulna Jetanta, about2km west of Torotoro.

From the transgressive base of the middle El Molino in the RCuchira Waykho,3 kin south-southeast of Torotoro, along rostrumof a salmohiform (family indet.2, gen. andsp. indet.; Gayet 1991)

is assigned to the marine Cretaceous suborder Ichtliyotringoi(semu Goody, 1969).

From an undetermined member at Cru z Khasa.on the west sideofthe cemetarysouth of Torotoro , remainsof Prtcapristissp.,crocodiles and turtles(?Ro.xoclielyssp.) have been recovered(R.CCspedes, personal com munication; spec imens in Museo d e HistorNatural, Cochabam ba).

At Tiupampa, located about 95 kin southeast of Cochabamba, fossil levels are known(Fig.4). The lowest,near Hera Mokho, isabout 90 m above the base of th e Cretaceous section fromcalcareous sandstone horizon ofthe middleEI Molino(see Marshallet al.,1985. Fig. 4); vcrtcbrntcsnrc rcprcscntcd only bya sclochinn(Dasyatidae,flasyarissc11aejJeri) and an indctcrrninatc crocodile(Marshallet al . , 1985) . Therea r e also abundantgastropods(Melanidne,Melania potosicnsis)which unTortunulclynrc not usefulfor ngc rcsotulion (Ahllcld A r i i n i s n , t960). t4iglicr i n lhc scction,along the R i o Pucarani, isan uppcr EI Molino bone bed withSiluriforines (Ariidae,Rhineastessp.), indeterminate turtles andcrocodiles.

At Vila Vila (=Villa Visc arra ), abo ut 90 krn southeast Cochabamba and6 kin northwestfrom Tiupampa, a fossil level inthe lowerEl Molino about100 ni abovethe base of the Cretaceoussection(see Marshallel al., 1985, Fig. 4)has yielded some of thesaine selachians recorded by Cappetta (1975) from tht lowerMolino at Torotoro(Prtcuprislis branisi, Isclryrliiza hartenbergeriPircabaris hoffsfefreri);umerous actinopterygian fishes: holosteanPycnodontiformes (Pycnodontidae, indet.) , Semionotiform(Semionotidae. nov. gen.); teleostean Characifornies (SerrasalmMyleinae). Siluilformes (Ariidne,Rlrinensressp..Andinichthyidaeindet.),cf. Cyprinodontiformes; Brachiopterygii (=CladistiaPolyptcriforines (Polypteridae,Dnjerelln srrdmnericnnn;GayetMeunier, 1991a, b. c); Dipnoi (Lepidosirenidne,Lepidosirencf.paradma;Schultzc,199 n); n turtlc(Iodocllciiiididrlc. ?I?o.toc/wI.yscf. v i l ~ ~ i l ~ t ~ s i s ;j ro i i i , 199 I), nd two iiidctcrniinotc ciocotlilcs(Marshallet al., 1985).

Two new localities werediscovered duringthe 1989fieldscnson,Estancia lnncoRancho and Iiijcha Pa tii, niidwoyalong theClizn-Anznldoroad, about40 kin southeast of Cochabanibn. At PajchnPata (lowerEI Molino) were recovered reniaii lsof selachians(Pucubaris Iroffstetreri , Pucaprisfis brunisi) ,ho los teanPycnodon t i fo rmes (Pycnodon t idac ,Coelodrcs toncoensis;L e p is os c id t i c , Lep s o s e s sp. ; g I II o id li o 1 s e n sc a C ) ,Clupeifornies(Gasreroclrrpea branisai),Siluriformes (Ariidae,Rhineasfes sp.; Andinichthyidae, indet.), cf. Cyprinodontifoniienumerous bones belonging toas yet unidentified small teleosteans,lungfishteeth (Lcpidosirenidae,Lepidosirencf. L .paradoxo),amphibian Anura,one vertebra which could bean atypicalUrodela,two other vertebrae which havethe morphologyo f a Urodelnb u t areprocoe l i c (the ve r t eb rae o f Uro de laare a lways oph i s to - o ramphicoelic), probablya snake , a tooth ofa sninll tlieropod

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Figure4. Detailed ma p oftlie Tiupa mpa ar ea (for location, see Fig.2) showing ocalities(1-9) rom the principal ve rtebrate level oIlleSan ta Luca Formation, an d localit iesin lhe middle(A)and upper(B) El Molino Formation. Tiupampais the local name usedbyQuechua indiunsfor tlie ureashown on lhe mup. Bused on Cnrln Nacionul, Ihliviu, Sun Vicenle Q uudrungle (see cuption lFig.3).

(Coelurosauria; Marshall,, 1989b), indeterminate turtles. variouscrocodiles nnd indeterminate trackways. At Blanco Rancho(upperElMolino) were found Clupeirormes(Gusleroclupeubrunisui) .Siluriformes (Ariidae,RltineasfPssp.), cf. Cyprinodontilormes,turtles and numerous ostracods.

Tra ckw ays of small bipedal dinosaurs (Coelurosauria?) arerecorded from Abpampa at Cerrito de Llamachaqui, Departrnento fPolosi (Leonardil1981).

Fragments of large bones, possibly of dinosaurs, and "dientes detipo cnico simple" were observed ina level with gastropods in tlieEl Molino Formation nearLa Caba a, north of P arotani and westofthe Ro Rocha about35 km southwest of Cochabamba (AhlfeldBranisa,1960).

Leonardi (1981: 935) reportsa series of six poorly preservedfootprints ofa bipedal dinosaur in the Santa Luca Fonnation alongthe Cochabamba-LaPaz road near Parotani in the Sutico llo Syncline,

soutlieastof .Cochabamba. These are apparently thesaniefoolprintsdescribed by Mnrshall Molina(1990) the lowerEI MolinoFormntion4 km south-soulhenst of Satitivafiezrind 10 kin east ofParotani:The best preserved footprints appear referable tanorn i thopod , p robab ly o f the f ami ly Hadrosaur idae ,

o t h e r s m a y r e p r e s e n ta s m a l l t h e r o p o d , p o s s i b l yacoelurosaur.From Sayari at km87 on the road from Cochabamba toOruro, R.

CCspedes recovered rem ains ofa selachian(Pucupristis brrurisi),asiluriform (Ariidae,Rhineuslessp.) andan indeterminate turtle fromthe lowerEI Molino (specimens in the Museo de Historia NaCochabamba).

Chullpa Khasa,6 km southwest of Morochata,has yieldeda fishmandible (Osteoglossiformes, Osteoglossidae, Phareodontinremains of reptiles (turtles, crocodiles) from an undetermEIMolino member.

401


10/42

/Figure5. Stratigrapliic sections of late Cretaceous-Paleocene rock units at Tiupampa, PajcliaPata (+ Estancia

Ulanco Rancho), Torotoro(+ Caranota),' Potosl (road toLa Falca and Cayar a) and, in part , Agua C lara, showinglocation of main fossil iferous levels. Undecompacted thicknesses. The base of the unconformable CayarFormation,is held horizontal.Fm,Formation;I, lower;m,middle;u, upper;Pz, aleozoic:1 erosional surface;2,conglomerates;3, conglomeratic sandstones;4, sandstones;5, fining and/or thinning-upward;6, siltstones andmudstones;7, car bon ate s (mostly limestones);8, coarsening and/or thickening-upward;9, gypsum;10, tuffitebed; 11 strom atolitic level;12, dissolution breccia;13, channels;14, cross-bedding;15, oiiids;16, calcareouscement;17, rooting;18, coalescent rhizolith-rich levels;19, bioturbation;20, red color predominant;21, dinosaurt rackways;22, verte brate -bea ring levels.All sections measured by T. Sempere, except part s of Torotoro-Caranota and Potos (respectively by tlie Brigadas10 and9 of YPFU) and of PajcliaPata and Agua Clara (byG .Camoinand J. M. Rouchy);all were checked by the authors.

402


11/42

FOSILESYFACIES DEBOU- - VOL. I - VERIBBRMOS

A vertebrate fauna descrilxd originallyas of possible Palcocelieage was reported fo rtiUie northern flank of tlie Huarach ani syncline,just northof tlie pueblo Huarachani along the Bolivia-Peru bordernorth of Lake Titicaca (Argolloel al., 1987). The fauna includesa

fish (Osteoglossoitiorplia, Osteoglossidae, Phareodontinae; Gayet,1987b, 1991) anda crocodile vertebra (Mesosuchia, Dyrosauridae,gen. arid sp. indet.). However, these fossils come from the middlenieinber of the El Molino Formation (called Ococoya Formationinthis area; Fig. 1), and are therefore of late Cretaceous age (Martinez,1980:183; Gayet, 1987b; Mourierel al., 1986, 1988:171; seebelow).

6 .Nor thern Subaridean belt

AboutGO-70 in abovethe base of the late Cretaceous age EslaMnFormation (=FloraFormation s.1. of Perry, 1963), from levels thatcniibc correlatedto the lowerEl Molino,nloiig the Rlo Flora, 315kin nor th -nor thwes tof L a P a z , were co l l ec t ed r cmainsofPucapris l issp. (=an cliop risris) , Gaslerocllrpea brunisaiandcharophytes (includingPeckichara compressa)(Perry, 1963; DBvila

Poncede Lc611, 1971).I n the sainearea, Gasferoclupea branisaiwas recoveredfroin the overlying Flora ForinationS.S., an quivalentof theupper El Molino (Vernet Bok llo, 1975).

7. Cen tral Subandeen belt

Sanjins-Saucedo (1982) and Mpez (1983) reported remainsofGasteroclupea branisaifrom the Cajones Formation (partly ortotally equivalentto the EI Molino Fo rmation ; Fig. 1) in the MoileSynclin e alongtlie Ro Moile about 95 kin west-northwest ofSantaCruz. The level in which the fossils were foulid probably represents

the upper E l Molino.G. SANTA LUCIA FORMATION

Th e richest vertebrate fauna, in ternis ofboth number and qualityo f spec imens , i s f rom Tiupampa abou t 95 km sou theas t ofCochabamba (Figs.2, 4). Two fossil levels at Tiupampa belongtothe basal iniddlcand upper EI Molino(sec nbovc). Athird fossillevel is locatedabout130in above the base of t he Cretaceous section(see Marshallel al., 1985, Fig. 4). Sempere Marshall (in press)dcinonstratc that this i n n i n rossil level, which representsthe typefaunaof the TiupainpianLand Mammal Age (Marshall. 19890)belongsto the Santa Luca F orniation.(see Figure5). The followingvertebrate taxa are recorded:

a) Fishes (Gayet. 1988b. 1990, 1991; Gayet Meunier, 1991a.1991b, 1992; Schultze, 1991 ): teleostean Clupeifo rrnes (Clupeidae.Gasteroclupeinae,Gasreroclupea bra nisai),Osteoglossiformes(Osteoglossidae, Phareodontinae,Pliareodusichfltys favernei;Osteoglossinae.incerfae sedis)(HiodontidaesensuMuizonel al.,1 9 8 3 i sa confus ion with Os teog loss inae ; Gaye t , 1991) ,Cha rac i fo rme s , Charac idae , Te t ragonop te r inae , i ndet ., c f.Rhoad siinae; Serrasalm idae, Serrasalminae, Myleinae; Erythrinidae,Hoplias nov. sp . ) . Si lu r i fo rmes (Ar i idae,Rhineasless p . ;AndinichUiyidae,Aridiniclllliys bolivianen sis;familiesincerraesedis

1 and 2 , IfofJsfertericltrltyspucai and Incaiclirhys srrarezi),Perciformes (Centropoinidnc), Polyptcriforines (IolyptcriDajelella sudanicrictmn)and Dipiioi (Ccratodoiitidnc,Ccralodrtssp.and cerntodont n.g., n. sp.; Lepidosirenidne,Lepidosirencf.paradoxa).

b) Amphibians (Rage, 1986, 1991a): Anura (Leptodactylid

Gym nophion a (family indet.).c) Turtles (Broin, 1988, 1991): Podocnemididae,Roxockelyscf.vilavilensis.

d) Lizards (Rage, 1991b): Lacertilia, ?Iguanidae and family ie) Snakes (Rage, 1991b): Aniliidaen. g.; Boidae, two species

indet.; ?Maqtsoiidae; Tropidopliiidae.f) Crocodiles (Buffetaut, 1991; Buffetaut Marshall, 19

Mesosuchia , Sebecidae ,Sebecris querejazus;Dyrosaur idae ,Sokorosucltusaff.iwiwilsoni . .

g) Mammals (Muizonel al., 1983; Muizon,el al., 1984a, b;Marshallel al. , 1983a,1985. 1989; Muizon Marshall, 1985,198711, b, c. d, 198 8, 199 1.i n press;Marshall, 1992: MarshallMuizon, 1988, 1992):Dcllotlicroidn (family indel,,J a ~ k h ~ d ~ / pntinufus), Peradectia (Ieradcctidae, Peradectinae,Pcradcctesa u s ~ r i n ~ n ~ ;aroloainegli i i i i inne, Roberl l ioffs~el ler iat iar io t ia lgeograpl i ica) , Microbiotheria (Microbiotheriidae,Kliasincordi llerens is),Dide l pli ini or pIi ia (Did e 1 li id oc , Didel pli i ri ne,Piicadelpltys andinris, Incudelpliys anriqrrus, Mizquedepi lpinensis ;Eobrasil i inae,Tirilordia flore si) ,Sparassodonta(Ha t h1 acy n ida e , AI lqokir 11s a s r a1 s ) , Pauc i u be rc u1 ta(Kollpaniidae.Kollpnniafiupanipina), order and family indet.(Andinodelpliys cochabanibensis),Leptictida (Palaeoryctidae?, cf.Ciniofeslessp.), Pantodonta (Pantolambdidae,Alcidedqrbignyainopinafa ) ,Condy la r th ra (Hyopsodon t idae , Mioc laen inTiuclaenusniinulus,Molinodus suare zi, Pucanodus gagn ierAndinodusbol iv iens is ) , and Notoungulata (cf. HenricosbomiidaeorOldfieldthomasiidae, gen, et sp. indet.).

Criadero de Loro,a new locality about1.2kmsoutii-southeast ofthe quarry at Tiupampa and fromthe sanie horizon (Fig.4, locality7). was discovered in 1989. Thetaxa seemto be the scarne as thosefrom the quarry and adjacent localities (Fig.4, ocalities 1-6, 9)andinclude, Siluriformes (Ariidae.Rhineasles sp.) , Perciformes(Centropornidae) , Polypter i formes (Polypter idae ,Dajetel lasudamericana) ,nu iiicrous1 rgc lu iig f is i t ce t h ( inc luditi gCeratodoiitidiic and Lepidosircnidac), snnkcs(oidac sp. indct.),turtles(?Roxochelys vilavilensis),crocodiles anda pantodont(Alcidedorbignya nopinafa).

At VilnVilb, om tin u p p x lcvcl lociilcd nboutLGOni above thebase oftlie Cretaceous section (see Marshallel al., 1985,Fig. 4),have come a turtle (Podocnemididae,?Roxochelys vilavilensis;Broin, 1971) anda crocodile (Sebecosuchia, Sebecidae,Sebecus

querejazus;Buffetnut Marshall, 1991).At Uie two newly discovered localities ofEstancia Blanco Ranchoand PajchaPata, south of Cliza (Fig. 2), are bone beds ofthe lowerand/or upper members of the EI Molino described above, whareoverlain by red sandstones of the Santa Lucia Formation simLothose at Tiupampa(Montao, 1968). A brief surveyof BlancoRancho resulted in the recovery of numerous teleostean (Osteogloss i formes , Osteogloss idae; Si lur i formes , Ar iR li nea st es sp., A n dini cli thy id ae, i nd ete rm na te teleo s s ,amphibians, crocodiles (Dyrosauridae, comp lete skulls),

403


12/42

MIREILLECIAYET. LARRY O . MARSIMLL IIUBKRY,SEMIEKI

TA B L E 1. VERTEBRA TE FOSSIL LOCALITIES OF MESOZOIC AND PALEOCENE AGE 1N BOLIVIA ARRANG ED BYSTRATIGRAPHIC OCCURRENCE.

TACURU GROUP1. Serranade Mandeyapecua

CASTELLON FORMATION2. Quebrada de Charagua

MIRA FLO RES FORMAIION3. Macha

AROIFILLA FORMATION4. Calerlas (dinosaurfootprints)

CHAUNACA FORMATION

5. Aguaclara 6. LaPaka

EL MOLINO FORMATION

LOWER MEMBER

7. Aguaclara8. Cayara (Cerro Muyu rina)9. Cho caya (Ro Angosto)10. HotelCordillera near W ila Khasa; Quebrada Saytu Jokhu with

11. LaPalca (blackshalelevel withGasteroclupea,and below

12. Pajcha Pata13. Kanclio Hoyada (Q uebr ada Texisca)14. Sayari(km87)15. Scvaruyo16. Torobro (Ceiro Llama Chaqui. dinosaurfootprink)17. Torotoro (Uma Jalantn cave, dinosaur Foolprints)18. Torotoro(UmaJalantnroad, dinosaur footprints)19. Torotoro (pista de danzas. din osaur footprints)20. Torotoro (selach ian level)21. VilaVila(Piccapriskyicvel)22. WilaKliasa (km 100)23. RoFlora24. Parotani (=Santivaez) (dinosaur footprints)

Gast eroclupea

stromatolites)

MIDDLE MEMBER

UPPER MEMBER

31. Calazaya32. Cayara (Cerro Muyu rina)33.*LaPaka(Doliciwciranipsamininialevel)34, Pajcha Pata35. Estancia.Blanco Rancho36. RoFlora37. Ro Moile38. Tiupampa (Ro Pucnrani)

INDEIXIMINAIE MEMUER

39. Arapampa (dinosaur footprints)40. Camargo (diiiosiiur fo otprints)41, ChullpnKhasa42. Jay-Jay43. Serpa44. Tambo Colorado45. Toinave46. Torotoro (CruzKhasa)

SANTA LUCIA FORMATION

47. Chaupi Khocha48. Maragua49. Pajcha Pata50. Estancia Blanco Rancho5 1. Tiupampa (quarry and associated localilies)52. Tiupampa (Criaderode Loro)53. Torotoro54. VilaVila

IMIOKA FORMATION

55. Cliaupiuno56. VillaIachcco

25. Hotel Cordillera (near Wila Khasa)26. Huaracliani27. Rancho Hoyada (Quebrada Texisca)28. Tiupamp a (HeraMokho)29. Torotoro(Ro Cuchira Wayk ho)30. Vilcapujio m i l a Apacheta)

404


13/42

TAULE2. SYSTEMATICLIST OF VERTEBRATES FROMTHE MESOZOICA N D I'ALEOCISNE OPUOL IVlh WIT H AUTHORSIIII'.

CLASS SELACHUOrder RajiformesSuborder Sclerorhynchoidei

Family SclerorhynchidaePucaprisris branisiSchaeffer, 1963Iscltyrltiza IturtenbergeriCappetta, 1975Scltizorkizaun.slronwriCappetta, 1975

' Order MyliobatiformesFamily DasyatidaeDasyarisbranisaiCappetta, 1975Da gt is molinoensisCappetb. 1975Dasyatis scltucfleriCnppeltn. I975Dasyarisnov.sp. 1Dasyatisnov.sp. 2Dasyalis nov.sp. 3

Pucabah ltoflslelteriCnppclln. 1975Pucabahnov. sp.

Family Rhoinbodonlidnc

CLASS OSTEICHTHYISubclass ActinopterygiiSupero rder "Holostei"Order F'ycnodontiormes

Family PycnodontidaeCoelodus roncoensisBenedetto Sanchez, 1972Pycnodontidae indet.

Order Semionotiformes

Family SemionotidaeLepidolessp.nov. gen.

Order Ginglymodi

Lepisosteussp.Family Lepisosteidae

Superorder T clcostciOrder Clupciforincs

. Family ClupeidaeGasreroclupeo branisaiSigneux in Branisaet al. 1964

Order Osteoglossi ormesFnrnily Osk oglossidseSubfamily PhareodontinaePkaerodusiclitlzys tavrneiGayet, 1991Subfsinily Osteoglossinaenov. gen.

Order "Salmoniformes"Family Enchodontidae

Suborder ichthyotringoideiFamily incertae sedis1

Emhodissp.

? Apdeodussp.


14/42

Family incertae sedis2gen. andsp.indel.

Order CypriniformesFamily incertae sedisMoliniclitliys inopinatusGayct, 1982c

Order CharaciformesFamily ErythrinidaeHopliasnov. sp.cf.Hoplias

Family SenasalmidaeSubfamily Myleinaegen. andsp. indel.cf. Subfamily Serrasalminaegen. andsp. indel.

Family CharacidaeSubfamily Tetragonopterinaegen. andsp. indel.cf. Subfamily Rhoadsiinnegen. andsp. indel.

Order SiluriformesFamily Ariidae

RliineastesSuperfamily AndiniclithyoideaFamilyindef.

nov. gen.Family AndinichthyidaeAndiniclttliys bolivianensisGayet, 1988b

Family incertae sedis1Incaichfhys suareziGayet, 199Ob

Familyincerlae sedis2Hoffsfettericlitkys pucaiGayet, 19Wbat least five gen. and sp. nov.

Superorder Atherinomorphacf. Order Cyprinodontiformesgen. and sp.indel.

Supero rder AcnnthopterygiiOrder PerciformesSuborder Percoidei

Family Centropomidaegenandsp. indel.

Order TetraodontifonnesFamily Eotrigonodontidae

Slepl~anodrrs iinintusGayet, 1991

. SubclassSarcopterygiiOrder Dipnoi

Family CeratodontidaeCeratodirssp.gen. andsp. indel.

Lepidosirencf. paradoxaFamily L epidosirenidae

CLASS CLADISTM (= BRACHIOPTERYGII)Order Polypteriformes'

Family PolypteridaeDajetella sudaniericanaGayet Meunier, 1991

406


15/42

FOSILESYW a l s DEDOLMA - V O L 1 VERIHBRAWS

CLASS AMPIIIUIAOrder Anu a

Family Lepldactylidae

gen. and sp.indel.Order GymnophionaFamilyindel.

gen. and sp.indet.Order Urodela

Familyindel .gen. and sp.indet.

CLASS REPTILIAOrder Ch,elonia

Family Podocnemididae? Roxoclielys vilavilensisBroin, 1971? ROXOC~ICIYSJ vilavilcrisis

OrderSquaiiintn

Suborder LncertiliaFamily ?Iguanidaegen, and sp.indel.

Familyindet.gen. and sp.indel.

Suborder OphidiaFamily Aniliidae

Family Boidaegen. and sp.indet.

gen. and sp.1 ndel.gen. and sp.2 indel.

Family ?Madtsoiidaegen. and sp.indel.

Family Tropidopheidae

gen. and sp.indel.Suborder MesosuchiaFamily S ebecidae

Family D yrosauridae

Order Crocd il ia

Sebecus quer ejazus Buffetaut Marshall,1991

Sokotosukusaff. ianwilsoni Halstead,1975SuborderEusuchia

Family D olichochampsidaeDolickoclmnpsa n i in in la Gasparini Buffclnut, 1980

Order SaurischinSuborder TlieropdaInfraorder Coelurosauria

gen. andsp. indet.Suborder SauropodomorphaInfraorder Souropoda

gen, and sp.indel.Order O mithischiaSuborder Ornithopoda

Suborder AnkylosaunaorCeratopsia

Orderindef.

gen. and sp.indel.

Ligabueic l in ium bol iv ianumLeonardi.1984

gen. and sp.indel.


16/42

CLASS MAMMALIAInfraclass M etatheriaOrder Deltatheroida

Familyinder.Order Peradectia

Jaskliadelphysniinutus Marshall Muizon, 1988

Family PeradectidaeSubfamily PeradectinaePeradectes austrinuni(Sig, 1971)Sublamily CaroloamegluniinaeRobertlioffstetferia afionalgeograpliicaMarshallet al.,1983

Order MicrobiotheriaFamily MicrobiotheriidaeKhasiacordillerensisMarshallk Muizon, 1988

7 Orde r DidelphimorphiaFam ily DidelphidaeSubfamily D idelphinae

Pucadelpliysandinus Marsha ll Muizon, 1988Incadelpliys untiqrrrrsM ~ ~ l i a l lMuizon. 1988M i z y r t e d e l ~ ~ l i y s p i l ~ ~ i n e ~ i s~ ~ l i i i l lMuizoii, 1988Subfamily EobrasiliinaeTiulordiafloresiMarshall Muizon,1988

Order SparassodontaFamily HathliacynidaeAflqokirrlsaustralisMarshall Muim n, 1988

Order PolydolopoideaFamily Pol ydo IopdaeEpidolopssp.

Order PaucituberculataFamily Kollpaniidae

Kollpania fiupanipinaMarsha ll Muizon, 1988

Family indef.Orderindef.Andinodelpliys cocliabunibensisMarshall Muizon, 1988

Infraclass EutheriaOrderLepticrida

Family Pnlnporyctidae?cf.Ciniolesres sp.

Order PantodontaFamily PantolambdidaeAlcidedorbignyn inopinafaMuizon Marshall. 1987a

Order CondylartluaFamily H yopsodontidaeSubfamily MioclaeninaeTiuclaenusniinirfus Mu imn Marsliail, 1987b

Molinodus suareziMuizon Marshall, 1987cAndinodus boliviensisMuizon Marshall, 1987cPucanodus gagnieriMuizon &.Marshall,1991

OrderNotoungulataFamily cf. HenricosbomiidaeorOldfieldtliomasiidaegen. andsp. indef.

408


17/42

~~ ~

~

TABLE 3. YSTEMATIC LIST OFVERTEBRATESFROM THEMESOZOICA N DPALEOCENEOFBOLIVIAWITH INDICATIONOFLOCALITIESANDSTR AT IGW HIC OCCURRENCE BASEDON INFORMATION INTABLE1 ANDTEXT . Abbreviations: Ca. Castelln Formation;Mi, M ird or es Formation;

Ar, Aroifilla Formation; Ch, Chaunaca Formation.

TAXA

h s clachiiOrder RajiformsSuborder Sclemrhynchoidei

Family SdcmrhynchidaePuraprisrirbranislschyrhiza harrenbergenSchizorhiza4 rromcri

Order MyliobatifonnesFamily DasyatidaeDq ar is bradsat'

Dq ar is SchaefferD q a t i snov. p. 1Dasyarisnov. p.2D q a t i snov. sp. 3

Family RhombodontidaePurabarir hoffsreneriPurabatisnov. sp.

D q a r i smolinoemis

lass Ostcichthyes;uk lassActinoptcrygiiMaclass "Holostei"Order Pycnodontiformes

Family PycnodontidacCoelodurroncoqsisgen. and sp.inder.

Family SmiqnotidacOrder Sanionotiforma

Lepidoressp.gen. andsp.nov.

Order GinglymodiFamily LepisostcidacLepisosteUrsp.

nfmclass TeleosteiOrder Clupciformes

Family QupcidaeGasreroclupea branisa

CaFm

2

MiFm

3?

ArFm

ChFm

6

S?

EI Molino Formation

lower member

11.12 14.20.217,11,20,21

7.11

2011.2011.207

11.20.217

12

7.12.13.21

1.9.21

7,9,13

7.8.9,10,11,12,.15

middle mb

2.252525

25

25.28

27n

25,27,30

25

upper member

25

25.21

3235.36, 37

Santa LucaFm

51

ImporaFm


18/42

ee

TAXA

OrderOsteoglossifo~esFamily OsteoglossidaeSubfamily PhareodontinacPhareodurichthys avermiSubfamily Ostcoglossinaegcn. andsp. nov.

Ordcr SahoniformesFamily Enchcdontidae

Suborder chthyoiringoideiFamily inccmcsedis 1

Family incenacsedis 2

Enchodus sp.

?Apareo& sp.

gen. andsp. indct.M e r Cypriniforms

Family incenaesedisMdlim'chrhys nopinafus

Order CharaciformesFamily ErylhrinidacHopliasnov.sp.cf.Hoplias

Family Sems ahid acSubfamily Myle/egen. andsp. inder.d Subfamily Senasalmina cgen. andsp. indef.

Family CharacidaeSubfamilyTetragonopterinacgen. andsp. inder.cf. Subfamily Rhoadsiinaegen. andsp.indef.

Ordcr SiluriformesFamilyAriidaeRhineasressp.

Superfamily AndinichthyoidcaFamily indet.gen.nov.

Family AndinichthyidaeAndinichthys boiivianensisrtndm'chrhys p.

Family incertae redisIncaichrhys uareziHoffsfetrerichrhys ucai

M iFm lower member

. - .

I, 13

7,9, 12,13, 14.21

12

ElMolinoFormation

middle mb

2521

26

25

25

29

25

25

25.30

25

75,21.30

~

upper member

33,35,38

34.35

Santa Luca

Fm

. .

49.51.52

51

51

5151

51

51

51

51

41-52

51

5149,51,52

5151

m p x aFm

56

56


19/42

TAXA

Superorder Athcrinomorphad. rder Cyprinodontiformagen. and sp. inder.

Superorder AcanthoptcrygiiOrder PercifomsSuborder R a i d c iFamily Centropomidae

gen. and sp der.Order Tetraodon rifomes

Srephamdusm m h uFamily Eotrigonodontidae

'ubdas SarwptcrygiiOrder Dipnoi

Family CcratodontidaeCerotodursp.

Family Lep idcsidd acLepidosirencf. paraaka

ass Cladistia (= Brachioptcrygii)Order Polyptcrifoms

Family PolyptcridaeDajetella sudamericana

ass AmphibiaOrder h u r a

Family Leptodactylidaegen. and sp. inder.

Order Gymnophiom

gen. and sp. inder.Family inder.

Order Urodcla.Family nder.

gen.and sp. inder.

ES Reptilia

>de r CheloniaFamily Podocncmididae? Roxochelys vilavilensir? Roxochelys cfi vilavilensis

Order SquamataSuborder LacertiliaFamily ?Iguanidac

gen. and sp nder.Familynder.gen. nd sp. nder.

Suborder OphidiaFamily A niliidaegen. and sp inder.

ArFm

El MolinoFormation

lower member

7,12,22

I, 13

12,21

21

12

14

middle mb

.

25

25.27

30

20.21

upper member

.35

33.35.38

Santa LucaFm

50.51.52

53.54

51.52

SI. 52

51

51

4849.5051-54

51

51

51


20/42

TAXA

Family Boidacgen. and sp.inder. 1gen. and sp.inder. 2

Family ?MvIatroiidaegen. andsp. nder.

Family T mpidopheidacgen. andsp. nder

Order CmcodiliaSuborder Mesmuchia

FamilyScbecidaeSebeeuquerejazu

Family DyrosauridacS o k o i o s h off amviLrod

Suborder EusuchiaFamily DolichochampsidaeDoliehochampsaminima

Order SaurischiaSuborder Thcmpoda

Infraorder Coeiumsauna

Suborder Saumpcdomorphaorder Saumpoda

gen. and sp.inder.Order Omirhischia

Suborder Omithopodagen. andsp. nder.

Suborder Ankylosauria or CeratopsiaLigabueichniumbolivianum

Order Inder.gen. and sp.inder.

gen. andsp. nder.

lass Mammalianfraclass MetatheriaOrder Deltathemida

Familyinder.

Order PeradmiaJmkhadelphysminu fus

Family PeradcctidaeSubfamily PcradeainaePeradectesaurrrinumSubfamily Camloameghh/maeRoberrhoffsreneria nafio nalgeographica

C aFm

MiFm

rFm

4

ChFm

Ellower member

12 21

19

12,16,17.18

19

1 9 . 3 -

19

?olinoForm:middle mb

?.5,27,28,30

26

upper member

33, 38

33

Santa LucaFm

5151

51

5148-50, 51 .

5154

5151

51

51

51

ImporaFm


21/42

TAXA

Order MicrobiotheriaFamily MicrobiotheriidaeKhasia cordil eremis

Order DidelphimorphiaFamily Diddphidae

Subfamily DidclphinaePucadelphysandinusIncadelphysm q mMizquedelphys pilpinensisSubfamily EobrasiliinacTiuloraafroren

Order SpalassodontaFamily Hathliacynidae-Allqokimausrrab

Family PolydolopidaeOder Polydolopoidea

Epidolopssp.Order Paucituberntlata

Family KollpaniidaeKollpam'a riupampina

Oder indctFamilyinder.

Infmclass EuthcriaOd er Lcptictida

Andinodelphys cochabambenris

Family Palacoryctidae?cfi Cimolesressp.

OderPantadonta

Family PdntolambdidacAlcidedorbignya nopinataOrder Condylanhra

Family HyopsodontidacSubfamily M oclaeninaeTiuclaenm minurusMol i~durs~lnrez ih ~ u r oliviensisPucanadur gagnien

Order NotoungulataFamily cf. Hennmsborniidaeor Oldfieldthomasiidaegen. andsp. indct

EMolino Forlower member middle mb

itionupper member

_ _ . .. -

Santa LucaFm

. .

51

50

515151

51

51

50

51

51

51

51,52

51515151

51

Impor:Fm


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MIREILLE OAYET. LARRY O. MARSIIALI. &I'IUBRRY SBMI'LKI

( ? R o x o c l ~ e l y si l a v i l e n s i s , comple te she l l s ) , and mammals(Dide lph imorph ia . Dide lph idae , inde t . ; Po lydo lopo idea ,Polydolopidae.Epidolopssp.). Indeterminoted turtles and crocodileswere observed a t Pajcha Pata.

About 3 kin south of Torotoro , remains of lungf ish(Cerntodontidae,Ceratodussp.; Lepidosirenidae,Lepidosirencf.paradoxa),turtles(?Roxochelyscf. vilavilensis), crocodiles and an

indeterminate mammal were collected froma sequenceof fine tomedium grained red sands of the Santa Luca Formation (Marshalletal., 1985; Broin 1991; Schultze, 199 1a) (Fig.3).

At Maragua (18 km west of Sucre) and Chaupi Khocha (11kmwest-northwest of San Lucas) were collected remains of silurifonnsfishes (Ariidae), turtles and crocodiles by M. Gayet andL. G .Marshall in September1989.

H. IMPORA FORMATION

Fossil vertebrates of possible late Paleocene age were recoveredfrom two locali t iesof lhe Impora Formation inthe CamargoSyncline.southern Bolivia.One is on he north side of Villa I'acheconear theRoSanJuandelOro, about 14 kmnorlhof Tojo, which hasyielded remains of siluriform fishes. Just north of this localityrcntiiinsof crocodicsy i d turtles were collected. The other localilyisat Chaupiuno, about 60 km norUiwestof Tarija. where indeterminatefish remains were found.

IMPORT,GNT L AT E CRETACEOUS-PALEOCENEVERTEBRATE FAUNAS FROM ELSEWHERE IN SOUTH

AMERICA

A. ANDEAN BASIN

In nor thwes te rn Argen t ina ,a sauropod (Ti t anosaur idae ,Laplatasaurussp. ) and a theropod ( familyincerfaes e d i s ,

Unqirillosaitrusceibalii) are known from the Los BlanquitosFormation (a temporal equivalent ofthe Chaunacaand/orAroifillaformations) in theupperpart ofthe Pirgua Subgroup (Powell, 1979)(Fig. 1).Frogs (Eokenepoidae,Eoxenepoidessaltensis) are knownfrom the Las Curtiembres Formation, which transitionally underliestheLos Blanquitos Form ation (Parodi-Bustos, 1962).

From the Lecho Formation (a basal lowerEI Molino equivalent;Flg . 1) are repor ted sauropods (Ti tanosaur idae,Salfasuirrusloricatus),Coelurosauria (Noasauridae,Noasaurusl e a l i ) andtheropods (Carnosaur ia indet . and Avisaur idae .Avisaurusarchibaldi)(Bonaparteet al., 1977; Bonaparte Powell, 1980;Brett-Surm an Paul, 1985; Bonaparte, 1986).The YacoraiteFormation (nnothcrEI Molino equivalent; Fig.1)lias yieldedacrocodile (Dolichocliai~ipsidae,Doliclrochunipsaniininru:Gasparini

Duffetaut, 1980) which was collected in association with apycnodont(Coelodits toncoensis; Benedetto Sanchez, 1972),aclupeidGasleroclirpea branisai)and turitellid gastropods (seeGasparini Buffetaut, 1980;Cione et al., 1985: 296).Other taxarepor ted f rom v ar ious locali t ies an d levels of the Yacorai teFormation includea selachian(Pucaprisfisbrunisi)anda siluriform(indet.) (Cioneet al., 1985; Powell, 1979: 202); the tooth of anindeterminate theropod (Powell, 1979: 203); and trackways ofcarnosaurs(Sa~ifichnus ienfoor),ornithopods (Hadrosauridae?.

114

I~adrosauricliniisarisfralis,Taponiclinrrs d o n o f t o i , 7'elosiclirt~issa l tens i s ) , indeterminate reptiles, and birds(Yacoraifichnrrsavis)(Alonso, 1980; Alonso Marquillas, 1986).

The Mealla Formation(a Santa Lucia equivalent; Fig. 1) hasyielded remains of teleostean fishes (indet.), turtles (indet.) andnoIo ungu1 te inni n nia1 (Henr ico s bor n iid ae, S i ipson o u spraecursor,S. nidjor;Pascualet al.,1978), whereas the Maz Gordo

Formation (an Impora Formation equivalent; Fig. 1)has yieldedspecimens of teleosteanfishes (Callichthyidae,Corydorasrevelat us;Poeciliidae, indet.), pleurodire turtles( Podocneniis argenfinensis;Cattoi Freiberg, 1958) and possibly a notoungulate mam(Henricosborniidae,Sinipsonofussp.) (Pascualel al., 1981).

In northern Chile, the following vertebrates are reported fSenonianage rock units: a selachian(Pucapristisbrunisi)from theTonel Formation (=lower part of Purilactis FormationS. I . ) (pideCionee / Q I . 1985)and dinosaurs(sauropods indet.) from theEstratos de Quebrada Blanca d e Poquis (anEI Molino equivalent)andthe baseof the Pajonnles Formation (Salinasef al., 199la. b).

IIIbru, vertebratesfrom the Vilqucchico Formation at its typelocal i ty near Vi lquechico, includePuc apr i s f i s b ran is iandCasteroclupeu branisai(DBvila Ponce de Le6n, 1971; CioneCal., 1985: 297). Amoredetailed faunallist is provided by Jaillardet01. (it1 press). Fromthe hiisc o f llicir niidtllcVilqucchico Forinii~ion(a Chaunaca Format ion equivalent) they repor t three f(P/ychofrygonsp., rhinobatid?, actinopterygian). Their upperVilquechico Formation is divided into three sequences whichequivalent, respectively, to the lower, middleand upperEI Molinomembers in Bolivia. From the lower sequence they reporanact inopterygian, f i sh and name two d inosaur t r ackways(Coelurosaur ia , O rni thomimida e,Ornithoniiniipusj a i l l a rd i ;Ornithopda, Hadrosauridae.Nadrosaitriclinusf iticucaensis); fromthe base ofthe middle sequence are two fish(Pucapristis brunisiandan actinopterygian); and from the upper sequence are two (Dusyafissp. and an actinopterygian) and numerous charophytes.

At Laguna Uma yo, f rom what w as cal led the V i lquechFormation and is now called the Umayo Formation (LaubacliMarocco, 1990, Jaillardet al., in press) are reported charophytes,actinopterygian fishes (Characiformes, Erythrinidae,Hoplias sp.,Serrasalmidae, Myleinae, indet., Characidae. Tetragonopterindet.; Siluriformesiricertaesedis ; Perciforiiics, family indet.),lung fis i (Cera todon id tie C e r ao dir s s p., Lep dosirenidn e ,Lepidosirencf. paradoxa),frogs (Leptodactylidaeindef . ) , snakes(Aniliidne), turtles (Podocnemididae,?Roxochelyscf. vilavilemis),crocodiles(indct.), egg shell fragments reportedly of dinosaurs,marsu pials (Pcrndcctidnc,Perudecles arrstrirrrinr;Didelphidac?;Pedioinyidaeo r Microbiother i idae) ,and two placenta ls(Cond y1 rt ira, Didoo don idae , undes cr bed; No to linguIa ta,Perutlieriidae,Perutlterirrnial f ip lanense) (Grambastcf al., 1967;Sigf . 1968, 1971, 1972; BonaparteP o w e l l , 1980;Riige. 1981;Marshallef al. 1983b, 1985; Kerourio Sig& 1984; MarshallMuizon, 1988; Schultze, 1 991a; Gayet, personal observation).

B. SOUTHERN ARGENTINA AND SOUTHERN BRAZIL

There are four important vertebrate faunas outside the Anbasin t h a t warrant special consideration: theyare' fromthe LosAlamitos Formation in Patagonia (late Cretaceous). the Adaiti


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MIREILLE OAYCT. LARRY O. MARSIIALL LIIUUKRYSCMIBKI

1978). waterlevel in the tectonic iicnr-shore lake wheretlielimestones were deposited might have been controlled by the levelofthe nearby Atlantic O cean through ad justment of marine andcontinental phreatic levels. In sucha case, the lacustrine episodeevidenced by the lower Itabora limestones would havebeencoevalwith a marine highstand. Since middle (and later?, see above)Paleocene mammals are known from the infilling of the karstic

cavities Uiat fo r m a after lake dessication,it

wouldbe

possible tocorrelate this highstand period withtheone in thelate Danian, andthe sub sequent dessication with the early Thanetian regression whichis also recorded near the Salamanca/Ro Chico contact in southernArgentina (see above).A subsequent highstand period wouldhaveresulted in the deposition oftheupper limestones ofSo Jose deItaboraf.

DISCUSSION

A. UIOSTRATIGRAPIIY

The Mesozoicand Ioleoccne age vertebrate fossil localities inBol iv ia (Fig .2) are ar ranged in Table1 accordingLo their

stratigraphic context. A systematic list of allthe known fossilvertebrates is givenin Tables2, and in Table3 with indication oflocality and stratigraphic occurrence.

In this section we review the k nown occurrence of the taxa listedin Tables2 and3 in orderto identify their cluonostratigraphic rangesand hence potential usefulness for calibratin8 the Bolivian rocksequence. Of particular interest is the apparent positionof theCretaceous/Paleocene(K/T)boundary.within the81 MolinoForm ation (see below).

1. Selachians

In Bolivia, selachians were recovered from the lower and basalmiddle members of the El Molino Formation. Rhombodontidae arepresent only in the lower member, whereas Sclerorhynchidae andDasyatidae are present in both. Sclerorhynchidae range from Albianto Maastriclitian (Cappetta, 1990).Schizorhizais known only inMaaskichtian age rocks from numerous localities around tlie world(Texas, Morocco, Middle-East, Niger. Nigeria and Zaire), whileIschyrliizaranges from the Turonian to Maastrichtian inNorthAmerica and Niger (Cappetta, 1987, 1990, 1991).Pucaprisfis(Sclerorhynchidae),Pitcubatis(Rhombodontidae) and the threenamed species ofD u s y u r i s (D. molinoensis,D. cltaefferi,D.b r u n i s u i ) (Dasyatidae) are currently endem icto the Andean basinwhere they are proving excep tionally useful in intra-basin correlation(Cappetta, 1990).Pircuprisfis branisiis also known froma levelbelow another which yielded dinosaurs in the lowerpart of theYacorai te Format ionin northwest Argentina (Powell , 1979).Dasya l i si s known in the Cenomanian of Texas , and f romMaastrichtiantoRecen t world-wide.

2. Pycnodontiformes

Pycnodon tiformes (Pycnodo ntidae) were collected in Bolivia fromthe Chau naca Fondation and the lower and basal middle members ofthe El Molino.Coloduswas reported from the lower Cretaceouso fBrazil (Santos, 1963; Wen z, 1989). Colombia (Porta, 19 70) and

Chile (Schultzc.1981). niid existeduiit i l the iiiiddlc Eocenei nmarine deposits around the world (Blot , 1987). Cione (197tentatively referred specimens discussed by Wenz (1969) frBoliviato Coelodusloncoensiswhich was erected by. BenedettoSQnch ez (1972) upon material from the Yacoraite Form ationofnorthwest Argentina. Gaye t (1991) cautionsthatthe spccics identityof some Bolivian pycnodontids has stillto be verified, while olhersfrom the lower and basal middleEI Molino Formation can bereferred with confidence toCoelodus foncoensiswhich wasapparently endemicLOthe Andean basin.

3. Semionotiformes

In Bolivia,Lepidofessp. was collected securely from thel a t eTriassic-early Jurassic. Elsewhere in South America,Lepidofes isreported from the Tithonian (=Portlandian) of Neuqun ProviArgentina (Aramayo, 1981). the Aptian-Albian.of Brazil (Aga1841;Roxo Lfgrcn, 1936; Woodw ard, 1895, 1908)a n d apossiblerecord frointhe lntc Cniiipiiiiiiiii-eiirly Munslrichtiunngc LosAlainitos Forination in Argentina (Cione, 1987). Elsewherein theworld.Lepidotesis recorded fromthe Rlietianof Germanyto

Cretaceous/Paleoceneof India (Gayetet al., 1984).Pillow-shape scales which rcprcscntu new gctius of thc familySemionotidae are known only in thelower and basal iniddlemembers of theEIMolino Formation in Bolivia (Gayet Meunierpersonal observation) and inthe Yacoraite Formation in Argentina(A. L. Cione, pers. com.).

4. Cinglymodi

I n Bolivia,Lepisosfeussp. is reported from the lower and basalmiddle El M olino. The earliest Lepisosteidae are reported fromearly Cretaceous of Niger (Ariunbourg Joleaud, 1943)andCongo(Casier, 1943). Younger fossils, includingLepisosteus,areknownfrom the late Cretaceous of India (Jain Sahni, 1983; Gaye tel al.,1984) and from the Cenozoic of Euiope, India and North Am(Wiley, 1976). In South America, Lepisosteidae are knowninArgentina from the late Campanian -early .Maastrichtian ageAlamitos Formation (cf.Atracfosferrsp.; Cione. 1987)andfrom theEI Abra Formation of uncertain age(A. . Cione, pers. com.)(Lepisosferts;M. G.. pcrs. obs.);Lepisosterisis known frointhe lateCretaceous Adainantina and Marilia formationso f the Upper BauruGrou p in Brazil (Santos, 1984; Gaye t Brito, 1989; Brito Gin press), and thelate Cretaceousof Colombin (Gayet, 1991).Lepisosfcits cxiststo Kcccntin southenstern USA and in CciitrulAmerica,

5. Clupeiformes

Gusferoclupeu brunisuiis apparently endemicto theAndeanbasinwhere in Bolivia it is reported from the Chaunaca (possibCajones,all three members of the El Molino, and the Santa Luformations; in Argentina it is known only from the YacorFormation (Cioneel al., 1985) and has never been found inoverlying Paleocene formations. The oldest known Clupeidafrom the Neocoinian of northern. Kyushu, Ja pan(Diplonlysrusprinzofiniis,D .kokirraensis}(Uyeno, 1979).

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n Bolivia, the subfam ilies Osteoglossinae and Phareodontinae ofthe family Osteoglossidae are recorded from the EI Molino andSanta Lucia formations (s ee Table3). Th e earl iest record fo rOstcoglossiformes in South Americais fromUie Aptian age Areado

Fonnation in Brazil (Santos, 1985) where the taxon is referredto

tliefamily Arapaemidae (L aeliichtliyinae) whichlias African affinities.Phnreodontinae were reported from the Paleocene and Eocene ofAfrica (Cappetta, 1972) and Europe (Woodward, 1901; Taverne,1978), Paleocene of southeast Asia (Sanders,1934), late Paleoceneof Turkmenia (Danil 'chenko, 1968), Paleo ceneof Austral ia(Taverne, 1975; Gay et, 1991). and middle Eocene of North America.Thus, Lhe Phareodontinae record from the EI Molino is the earliestknowntodate.

Squamules (partsof scales)of Osteoglossidae (Phareodontinaerind Osteoglossinae) were reported in the Maastrichtian of Niger,CrelaceouslPaleoceneof India and Paleoceneof Pnkistan (GayetMcunicr, 1983). Ostcoglossinae wcrc rcportcd from tlic I'aleoccncofSumatra (Sanders, 1 934), and rangelo Rcccnt inSouth America,

Asia, Africa and Australia.7. "Sulmoniformes"

In Bolivia, Enchodontidne(Enclrudus sp.) and Ichthyotringoidei(family indet. and7Apafeudussp.) were collected Gom the lowerandbasal middle EI Molino.Encltudus is known only from marineMaastrichtian ak e rocks in Co ngo (Dartevelle Casier, 1949).Morocco (Arambourg,1952). Brazil (R eh us as Santos, 1956),Europe (Goody, 1968), North America (Goody, 19691, Lebanon(Goody, 1969) and Israel (Chalifa, 1989)to note only the mostimportant in both preservation and quantity.

Apafeudus is a member of the suborder Ichthyotringoidei whichranges from early Cretaceous (?Albian)to Maastrichtian (Goody,

1969).

8. Cypriniformes

The cypriniform(Mul in ichf l tys inopinafus : Gayet, 1982b) fromIhc lower and basal middlc EI Molino rcprcscntst l ic only record,fossil orextnnt, of this ordcri n South Amcrica. Elscwlicrcin lheworld, this group i s' first know n from the Eocene ofNorthAmerica(Grandeef a l . , 1982). Gayet (1986b. c) describedRania l l ichfhysfrom the marine Cenomanian in Israel which maybe considcred anancestral, if not true, cypriniform.

9. Characiformes

In Bolivia, thisordcr is rcprcscnlcd bythe families Erytluinidac,Serrasa lmidae ( inc luding Serrasa lminac and Myle inae) andCharacidae (including Tetragonopterinae and Rhoadsiinae). All thesegroups have been foun d in the Santa Luca Formation, whilemembers ofthe Erythrinidae, Myleinae and Tetragonopterinae werealso found in tlie lowerand basal middleEIMolino Formation.

The oldest previous known Erytlirinidae(Nupl ias) are reportedfrom the Miocene of Ecuador (Roberts, 1975); Serrasalmidae(Myleinae) fromthe Maastrichtian? Umnyo Formationat Laguna

uinnyoin Perii (Gnyct. 199 I), M ioccnc L oyoln Formntion ofEcuador(Roberts, 1975) arid Mioccncof Coloiiibin (Lundbcrget al.,1986); C haracidae (Tetragonop terinae) fromthe Umay o Formationa t Laguna Umayo in Peru (Gayet, 1991). Miocene of Ecuado(Roberts, 1975) and early Mioceile of Taubat in Brazil (Schaelfcr,1947); Characidae(indef.) from the Paleocene ofMorocco(Cappetaef

a l . , 1978); Serrasalmidae (Serrasalminae) and Characidae(Rhoadsiinae) had not y et been foundas fossils. T he oldest knownrecord of characiform s outside ofSouth Americais Salni inupsibe r i cu s from the marine late Cenomanian ofPortugal (Gayet,1985a). Characiforms range.10 Recent in Africa and inSouthAmerica. One recent genus colonized Cen tral America.

10. Siluriformes

In Bolivia. this orderis represented by members of the familiesAriidae and Andinichthyidae from both theEI Molino and SantaLuca formations, andat lcast by two familiesinccrfae scdisfromthc Snntn Lu ch Forinntionnt Tiupempn (Gnyc~,991).

Theother known siluriforliis elsewhere in Soutli America are

represented by cf. Diplomystidae and cf. Ariidae in tlie Carnpaniage L o s Alamitos Format ion in Argent ina (Cion c , 1987) ;Siluriforiiicsindc/. fromIlic Mnns~riclitinntgc ColiToro (CioncCLaffite, 1980) and Yacoraite formations (Cioneet a l . , 1985) inArgentina; cf. Doradidae from the late Cretaceous age Adamantiand Maria formations in the UpperBauru Group, Brazil (GayetBrito. 1989; Brito Gayet, in press); and Ariidae from thMaastrichtian? age Umayo Formationa t Laguna Umayo in Peru(Gayet, 1991).

The onlyrecords for Cretaceous age Siluriformes outside of SouthAmerica area single tooth referredto A r i u s sp. from the latestCretaceous of Central India (Jain Salini, 1983) and doubtfotolithes from the late Cretaceous of North America (Frizzell, 19Fitch, 1975).

Siluriformes are known in the Paleocene of Argentina,[Arius?andB a c l i n t a n i a in Patagonia(Dolgopol de Saez. 1941), andPrupyg i d i r im (Bocchino, 1964) andC o r y d u r a s (which in Recentlimes is restricted to freshwater in South America) from the MaGordo Formation (ardack, 1961)J. Africa (in Congo; DartevelleCasicr, 1949; Cnsicr, 1960), Europc (clgium; Leriche, 1902Indoncsia (Sandcrs. 1934);nnd in tlic Eoccnc of North Anicricn(Lundberg, 1975; Grande. 1987; Gran de Lundberg, 1988), Euro(Woodward, 1901; Casier, 1946), Africa (Peyer, 1928) and Ind(Salini Misra, 1975). Ariidaerangeto Recent.

11 Cyprinodontiformes

In Bolivia, pharyngealteeth from the Miraflores Forniation aretcntatively referrcdto indelcrininate cyprinodontiforms.In the lowermember of theEI Molino Forination are know n several veryprimitive cf. Cyprinodontiformes (Gayet, 1991). The earliestprevious record of this gro up,w as from the Oligocene of Fran(Gaudant. 1988). and it rangesto Recent world-wide.

12. Perciformes

In Bolivia, perciforms of lhe family Cencropomidae are know

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MIREIUE OAYIX. LARRY O. MARSIMLL ELIIIIB KRY SEMI'IJKC

from the Santa Ldca Formation (Gayet, 1991). Indeterminatespecimens of this orde r nre reported in South America from theCampanian ageLos Alamitos Formation of Argentina (Cione, 1987).the M aastrichtianage Marlia Formation of Brazil (Gayet Brito.1989). and the Umayo Formation at Laguna Umayo in Peru (Gayet,199 1). The earliest' Centropomidae were previously recorded fromthe middle Eoc ene of Italy (Sorbin, 1970), and they range to Rec ent

world-wide.13. Tetraodontiformes

In Bolivia,Slepltanodus(Eotrigonodontidae) is recorded from thelower and basal middle El Molino Formation. Eotrigonodontidaewas reported from marine beds in the ?Albian-Aptian of Morocco(Tabas te , 1964) . Cenom anianof Egypt (Wei ler, 1935) , la teCretaceousof Nigeria (White, 1934), Congo (Dartevelle Casier,1949) , Nige r (Tabas te , 1963 ; Cappe tt a , 1972) . Morocco(Arambourg, 1952), Israel (Raab, 1963), Upper Cretaceous of lndia(Jain Salmi, 1983) nnd Tertiary levels of Europe, (Leriche , 1906;Casier, 1946), Angola (Dartevelle Casier, 1959) and Egypt (Peyer,1928). Eotrigonodontidae range from ?Albian-Aptian to middleEocene , whereasSfepl ianoduswa s reported on ly f rom theCretaceous (sanie rdccrcnces).

14. Polypteriformes

Dajefella sudamericana(Gayet Meunier, 1991a, b, 1992) fromthe lower El Molino and Santa Luca formations is the only recordfor Polypteriformes outside of Africa, where members of this orderare known from the Senon ian of In Becetem in Niger (Gayetet al.,1988), Miocene of Tunisia and Kenya (Greenwood, 1951,,1973), andPleistocene of Ethiopia (Aram bourg, 1948). They rangeto Recent inAfrica where they occ ur in freshwater.

15. Lungfish

In Bolivia, Ceratodontidae(Cerafodussp. and ceratodont n.sp.)and Lepidosirenidae(Lepidosirencf.paradoxa)tooth plates occurtogether in the early Paleocene of Tiupampa. In South America, thefirst ceratodonts occur in the Upper Cretnceous of Patagonia(Am eghino, 1906; Pascua l Bonde sio, 1976; Cione, 1987) andwerealso described from the Lower Cretaceous of northern Brazil(Cunha Ferreira, 1980). Ceratodonts were distributed world-widein tlie Mesozoic, still occurring during the Cretaceous in Africa,Madagascar, Australia, North and South America, whereas theNeoceratodontidae which occurin the early Cretaceous of Australiapersist there until today (Schultze, 1991a).

Lep idos i ren idae a re r epor ted f rom the Maas t r ich t inn E lMolino Formation at Vila Vila and the Paleocene Santa LucfaF o r m a t i o n a t To r o t o r o . F o s s i l s a r e r e c o rd e d f r o m t h eM a a s t r i c ht i a n ? a t L a g u n a U m a y oi n P e r u (Sig,1968).E o c e n e o f A rg e n t i n a ( F e r n a n d e zet a l . , 1973; Fernandez ,1976) and Miocene o f B raz i l(Santos, 1987) and Colombia(Bondesio Pasc ual, 1977). Lepidosirenids occu r today inthe Amazonian and Paraguayan re gions of South America.

16. hmphiblans

,

In Bolivia, indeterminate -eptodactylidae are known only fthe Santa Luca Format ion a t Tiupampa. Elsewhere in SoAmericn indeterminate Leptodactylidae a re recorded fromCampaniannge Los Alamitos Formation in Argentina (Baez, 1987and, questionably, from the Umayo Formation at Laguna Uma

Peru (Sig &, 1968). A probable L eptodacty lidae,Bairrubafracliuspriceihas been recorded in the Maastrichtian age Marlia Formain Brazil (BQez, 1985; BQez Per, 1989; Bertiniet al. in press).The oldest known Leptodactylidae are from the M a n a Formand the family extendsto the Recent (BQez, 1987).

Th e oldest known G ymnophiona are represented by two vertfrom the Santa Luca Formation at Tiupampa (Rage, 1986, 19Previously this group was known from the middle Paleocen(atItabora) to the Recen t.

Vertebrae of urodels are known from the lower El MolFormation at Pnjchn Pnta. This is the earliest recordof this groupinthe world(Kage, pers. com.).

17. Turtles

Roxochelyswas basedon spccinicns collected [rom thcCampaninn nge Adamnntinn Formationor the Upper Bauru Group inBrazil (Bro in, 1991; Bertiniet al., n press).?Roxoc/ielysvilavilensisBroin, (1971), from the Santa Luc a Form ation is probably refto a genus distinct fromRoxochelys(Bro in, 1991). The speciesvilavilensisis definitely known only fromlhe Santa Luca Formationin Bolivia and possibly frpm the Maastrichtian? Umayo FormatLaguna Umayo in adjacent Peru (see above), w hile very fragmmaterial from the lower EI Molino of Bolivia is referreto?Roxoc/ielysf. vilavilensis.This species w as apparently endemic tothe Andean basin.

18. LizardsAn unidentified member of what appears to be the fam

Iguanidae is recordedfrom tlieSanta Luca Formation at Tiupampa(Rage, 1991b). A possible iguanid(Prisfigrrattabrusiliensis;EstesPrice, 1973)is known from the Manstrichtianagc Mnrflia Formationof the Upper Bauru Group in Brazil, whileh e earliest uncontestedmembers of this family are from the middle Paleocene of B(Estes, 1983) and North America (Sullivan, 1982).

19. Snnkes

Th e four faitlilies of snakes listed below are known only froSanta Luca Formation at Tiupampa in Bolivia.

In South Americn, Aniliidhe are first reported from the UFormation at Laguna Umayo in Peru (Rage, 1981) and mPaleocene at Itabora in Brazil (Rage, 1987).

The earliest Boidae are from the Maastrichtian of North Am(Estes BBez, 1985), Euro pe (Rage , 1987) and India (Jain S1983), whilein South America, previous to this work, they were reported in the middle Paleocene in Brazil(Rage, 1987)andearlyEocene of Argentina (Albino, 1986).

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Madtsoiidnenrc first kiiown inthe Suiitoiiian/CninpnnianofMndngnscnr(Rage, 1984;Roiinparte. 1986) ni id Campaninnage LosAlainitos Formation in Argentina (Albino, 1986, 1987). In fact,Uieoldest Madtsoiidae is probably from Niger (early Senonian) (Rage,1984).

Previousto this work, Tropidoplieidae were first reported from the

middle Paleocene hi Brazil (Rage, 1987), middle Eocene inNorthAmerica(Hohnan, 1979), and Eocene-early O ligocene in E urope(Rage, 1984).

20. Crocodiles

Three families are known in Bolivia: Dyrosauridae (basal middleEIMolino and Santa Lucia formations), Dolichochampsidae(upperEIMolino Formation)aiid Sebecidae (Santa Luca Formation).

Dyrosauridaeare known from late Maastrichtian to late Eocenerocks around the world (Buffetaut, 1982). In South America,dyrosaurids are knownfTom what appears tobe the Mnastrichtian inrazil ,(Cope,1886; Buffetnut, 1978), anda possible dyrosauridiskiiowiifrom tic Manstrichtinti i n Argentina(Gnspnrini, per . coin.).Sokotosuchus ianwilsoni(Hals tend, 1 975) i s f rom the la teMaaskichtian Dukarnaje Forina tion in Nigeria.

Do l i chochamps idae( D o l i ~ l i ~ ~ l t a n i p ~ aiininta)i s knownelsewhereonly from tlie upper p u t of the Yacornitc Forinationinnorthwest Argentina and the upper El Molino Formation in Bolivia(see above). Th e family was apparently endemic to the Andeanbasin.

Sebecidae were previously knownfrom the middle Paleocene-middle Miocene (B uffetaut, 1982; Gasparini, 1984); they wereendemicto South America.Sebecus querejazusfrom the SantaLuca.Formationis the earliest known member o f the family(Buffetaut Marshall, 1991).

21. Dinosaurs i

Dinosaur trackways are knownfrom four localities in the lower EIMolino near Torotoro andat Santivaez nearParohni, and from twolocalities (Arapamba, Camargo) in an indeterminateEI Molinomember. The only dinosaur fossils in Bolivia area toothof aCoelurosnurin(Marshall, 1989b)from the lowcrEl Molinoat PnjchnI'ntn, and undcscribcd boiics from two localiticeof tlic CajonesFormationnear Santa Cruz(R. uh ez, pers. com.).

A l thougha deta i led s tudy of the t rackways has ye tto b eundertaken, it appears that at leastfour suborders (Theropoda,Sauropoda, Ornithopoda and Ankylosauriaor Ceratopsia) arerepresented wi th in the lower EI Mol ino a t Torotoro . Thisdemonstratesthat dinosaurs were taxonomically diverse in lowerEIMolino time.

For cluonostratigraphic purposes we followthe general mnsensusthat dinosaurs became extinct at the endof Cretaceous time and thatthere isn o compelling evidence(contraVan Valen, 1988) tosubstantiate Uiat they may have continued into the early Paleocene.Recent work by Lofgrenel al. (1990),for example, negatesaPaleocene age for dihosaurs from the upper Hell Creek Formation inMontana.Our position is concordabt with the biostratigraphic data inthe Andean basin summarized above and below.

8 FOSOSILES Y FACIES bE noum - VOL. I VERTEBMWS

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22. M,unimuls

Mammals are known only from the Santa Lu ch Formation Bolivia.Peradectes airstriniimwas based ona specimen collectedfrom tlie Umayo Formation a t Laguna Uinayo inPeru(Sig&, 1971,1972; Crochet, 1980) anda second specimenfrom Uie Santa Luca

Formationat Tiupampain Bolivia was later referred to this species(Marshall Muizon, 1988). The earliest knownPerudecfesis fromthe baseof the early Paleocene inNorth America (Van Valen, 1988:29) and is common thereafter in early Tertiary age rocks inNorthAmerica and Europe (Crochet, 1980).

Robert/ioflsferterias most closely related toProcuroloanreglriniafrom the middle Paleocene of Brazil (Marshallet al . , 1983a),a l though other more pr imi t ive members of the subfamilCaroloaineghiniinae are knownfrom the Maastrichtian in NorihAmerica(Marshal l et a l , ,1989) . The genera an d specieso fMi'crobiotlieriidae, Didelphidae, Hathliacynidae and K ollpaniidaeknown only Erom theSnnta Luca Formationa t Tiupampa,and theyall representthe earliest known m embersof these faniiliesin SouthAinericri(MarslinIl Muiimii, 1988; MiuslinIlet d., 989).

Cf. Cintolestessp. is the only known member of the orderLeptictida inSouth America.Cintolestesis recordedfroin theMnnskichtinnand cnrly Paleocene in North America (MarshallMuizon, 1988).

The pantodontAlcidedorbignyais the only known member of thisorder in South America. Pantodonts are fist known in the middPaleoceneo f North America wherePanfolambdabatlrniodonrepresentsa potential descendant ofAlcidedorbignya(MarshallMuizon, 1988). Among the New World pantodonts,Alcidedorbignyais the earliest and most primitive member y et known.

The mioclaenine Hyopsodontidae are the only representatiyesofthis familyin South America. In North America, members of thisgroup are 6rs t recordedin the early Paleocene (Marshall Muizon,1988; Van Valen, 1988).

The earliest known notoungulatesare represented byPeruUieriimtaltiplanense(Perutheriidae) fromthe Umayo Formation at LagunaUmayo in Peru (Marshal let a l . , 1983b) ,Sintpsonolus(Henricosborniidae) from the Mealla Formation in northweArgentina (Pascua1ef al., 1979), and cf. HenricosborniidaeorOldlicldtlioinnsiidncfrom lhe Siintri Luchi Forinntionnt TiupniiipninBolivia(Miusha11 Muizori, 1988).

1I

B, THE CRETACEOUS/PALEOCENE (KIT) BOUNDARY INTHE ANDEAN BASIN

Based011 the above discussion of chronostratigraphic ranges, thtaxa canbe grouped into five categories (see Table 3):

1. Those which presentlyare known onlyfrom the EI MolinoFormation and its stratigraphic equivalents in tlie Andean basThese are currently regarde$ as "endemics" which are potentiausefulfor intra-basin correlation' although their usefulnessi ncalibrating these rocks wittithe geologic time scalehas yet to befirmly established. Included arePitcapristis(Sclerorhynchidae),Prtcabafis(Rhombodontidae), the three speciesof Dasyaf is(Dasyat idae) ,Coelodrrs toncoensis (Pyc nod on t idae). andDoliclroclIanipsa (Dolichochampsidae).


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2. Taxa fromthe EI Molino and Santa Lu ch formations (andtheirstratigraphic cquivalcnls) which are "endemics" in the Andean basin.Included areGasteroclicpea bronisai(Clupcidae).Andinichtliysbol ivianensis , (Andinichthyoidea, Andinichthyidae) ,Hoffsterterichtliyspucai, Incaichthys suarezi(familiesincertae sedis)and?Roxochelys vilavilensis(Podocnemididae).

3 .Taxa known only from the lower and basal middle EI MolinoFonnation which are currently regardedas late Cretaceous "guidefossils" basedo n their records elsewhere in the world. Included are

SchizorhizaandIschyrhiza(Sclerorhyncliidae), apparentlyLepidot es(Semiono t idae ) which has on ly one dub ious record in theCretaceous/Paleo cene of India ,Enclzodiis(Enchodontidae),?Apaleodris (family indet.), Ichthyotringoidei (gen. andsp. indet.),Sleplranodiis(Eotiigonodontidae) and dinosaurs.

4. Taxa which occur in the EI Molino and/or Sant 'a Lucaformations and, which based on records from elsewhere in the world.lransgresscdthe K/Tboundary arid arc of dubiousor no valueindefiningthis boundary. These includeLepisosteus(Lepisostcidae),Os eog os s d ae, Cypriniform es, Ch aracior mes,S lu forni es,Cyprii iodoiit iforrnes. Perc iform es, Polypteriformes, D ipnoi.Leptodactylidne. Podocncniididac. Laccrtilia, Boidae, Madtsoiidae,Dyrosauridne. and Lcptictida.

5. Taxa recorded only in theSanta Luca Formation which are theearliest or only records for thesegroups in South America. Theydonot occur in association with diagnostic Cretaceoustaxa andtherefore are regakdedas Paleocene. Included are Ceratodontidae n.gen. andsp., Gymnophiona, Aniliidae, Tropidopheidae, Sebecidae,Peradectidae, Caroloameghiniidae, Microbiotheriidae, Didelphidae,Hathliacynidae, K ollpaniidae, Pantodonta, Hyopsodontidae andNotoungulata.

As demonstrated by group3 above), apparent late Pretaceoustaxa occur only in tlie lower and basal middle members of the EIMolino Formation. I t therefore appears that theK/rboundary occursabovethe basal middle member, whilethe upper El Molino membermay b e regardedas early Paleocene based on palynological study ofa stratigraphic equivalent in norlhwestern Argentina (Quattrocchioelal. ,1986) andod the absenc e of diagnostic Cretaceous taxa. W etherefore suggest &hathe K/T boundary lies somewhere in the upperpartof the middle member of theEIMolino Formation. In addition,tlie Sanla Lucla Formation, which also lacks diagnostic Cretaceoustaxa and conformably overlies the early Paleocene upper memberofthe EI M olino Formation, may be regarded as early, but no earliest,Paleocene in age.

C. PALEOENVIHONMENT

PishesFishes have been divided into four main categories (Myers, 1938)

based on their toleranceto fresh or salt water:1) primary freshwaterfishes found only in fresh water;2) secondary freshwater fishes,found normally in fresh water, but capable of entering and survivingin marine water;3) peripheral freshwater fishes, diadromous, which

migrate bctwccn fresh water and marine;nnd 4) marinc fishes.

The definition of these four categories is basedo n living fisliesand is not always evident with fossil forms (see below). For treason, the identificationof paleoenvironment based only o n oneor afew taxa is generally of dubious value.

1. Prim ary fre shwater fishes

a. Semionotidae

Th e fossil Semionotidae are generally considered as primfreshwate r fishes. A new genus is reported in Bolivia in the land basal middle members of theEIMolino Formation. Except forUie questionable Cretaceous/Paleocene record from India (Gayetal., 19841, Semionotidae are not known to survive beyond Cretaceous/Paleocene boundary. Their absence at Tiupampprobably relatedto the Paleocene age ofthe Santa Luca Forination.

The new genus fromAgut i CI:m und Hokl Cordillern, known onlyby scales and referredto the Semionotidae,seems endemictoBolivia and northern Argentina where it is recorded in both mand fresh w akr environments.

b. Osleoglossidue

Living O steoglossidae are restrictedto freshwater environments intropical South America, Africa and Asia. Some fossil remainsreported from fresh water (Green River basin, Wyoming, Gr1984; Sumatra, Sanders, 1934; Australia, Taveme, 1979), somebrackish water (mid dle Cretaceous of Zaire; Ta verne, 1976some from marine environments (Paleocene of Zaire; DarteveCasier, 1959; Turkmenistan, D anil'chenko, 1968; Niger, Cap1972, Gayet Meunie r, 1983; Eocene of Italy, Taverne , 1England, Woodward, 1901; Morocco, Arambourg, 1952; Denmark, Bonde, 1966). In well known marine Cretaceous l(Lebanon, Israel, Morocco, Italy, Portugal and Yugoslavia) wthe ichthyofauna is abundant,no fossils of this order a re represented.

Consequently, Osteoglossiformes a re consi dere das primaryfreshwater fishes following Nelson (1969) and Gayet (1987bInBolivia, Ostcoglossinac wcrcreportcdi n rill mcmbcrs of the EIMolino andin the San ta Lucia foniiatons, wlicrcns' Phareodontiwere reportedi n the Santa Luca Formation only (Tiupampa).n theEl Molino Formation, they occur in association with marine fis

c. Cyprinodontiformes

Nothing can be said about the paleoenvironinent of theCyprinidontiforines in Bolivia, because this group is too poknowI.

d. Polypteridae

Living Polypteridae are Ahi can freshwater fishes as are all of this group known from Senoniaito Pleistocene. In Bolivia heywere reported at Tiupampa (Santa Luca Formation) and occassociation with marine fishes at Vila Vila (lo we r EI MoFormation).

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e. Lungfishes

Ceratodonts are genera l ly associa ted wi th f resh waterenvironments by extrapolation based o n occurrence of extant forms.NeverUieless, ceratodon ts occur in undoubtedly marine deposits inZaire (Casier, 1961), Niger dnd Algeria (Martin, 1984), Mali(Tabaste, 1963). Egypt (S cha d, 1984) and Kansas (Schultze, 1981).Accordingto Uiese diffe rent authors, thesefaunasmaybe interpretedas d e ah assemblages of m arine, brackish and hesh water forms orthat they were tolerant ofa wide range of salinity.Lepidosirenids are restrictedto fkesh water env ironmen ts.

2. Secondary freshwaterrilies

fresh water species. This family includeslhb gcnus Ccrr~roponrrrsfrom thecoastal waters of the New World,a species wliicli entersBrazilian rivers (Berra,1981). Centropomidae a re presenta tTiupampa and Criadero d e Loro in Bolivia.

d. Cypriniformes

Cypriniformes are living freshwater fishesandfossil remainsattest,as f a r as i s known,to a s imi la r env i ronment . On ly

Ranrallicfitliysfrom Ein Jabrud (Israel) which may be considered aan ancestral Cypriniform es (but is nota true Cypriniformes)ismarine. Cypriniformesare reported in the lower and basal middlemembers of the EI Molino Formation in association with marifishes.

a. Lepisosteidaee. Characiformes

Most ofthe living Lepisosteidae are foundin fresh water, but onespecies may tolerate brackish or even marine conditions and is foundalongUie coast of the Gulfof Mexico (Sutlkus,1963). Bccouse oftheir Uiick scales coveredwith ganoine which are more resistan1llinnany oflier parts o fth fish, the fossil Lepisosteidae are often reportedon scaks only, which may be transported far from their biotopewithout damage. In Bolivia. Lepisosteidne are always found inassociation with marinefislies including selachians, pycnodonts andeotrigonodonts in the lower and basal middle membersof the ElMolino Formation, but theyare always disassociated, indicatingpostmortem movement. In the only locality where no true marinefishes were found( .e.Tiupampa), Lepisosteidae are absent butUlisabsence maye the consequence of age (Paleocene) and not ofenvironment.t pisosteidae ha da nearly world-wide distributionduring Cretaceous and early Tertiary times. Subsequently, theydisappeared from all the continents except North America wherethey are now restricted to the southeastern part of the USA andCentral America.

b. Clupeidae

Gasteroclicpea branisuiis a Clupeiformes endemicto the Andeanbasin. Clupeiform,es are generally marine fishes with the exceptionof some South American li.ving genera(RnnmoRnsrer, frisrignsrcr;G6ry.1969; Cioncp 'ercirii, 1985). IIIUic middle Eocene oft l icGreen River Format ion (Wyoming, USA),KniglrtiaandDiploniystuswhich are knownto be marine, are reported in strictlyfresh water environments (Grande, 19820. b).Gaslerocfupeabranisuiis reported at Tiupam pa (Santa Lucia Formation) where nomarine fishes are found, and in some levelsof the EL MolinoFormation (at Agua Clara) in association w i h marine fishes. In somegreen m ar l levels a t Agua Clara ,Gasferoclupeais found inassociation only with brackish-water o stracods. Consequently,Gasteroclupeamaybe considered, with some confidence, as a

secondary Jeshwater clupeid.e. Centropomidae

The family Centropomidae, as defined by Greenwood (1976). iscomposed of marine and fresh water genera.Lares, to which theBolivian genus maybe compared, is knownto include marine and

Living Characi formes are f reshwater f i shes (only a fewcxccpt ions enter brnckisl i waters) w hi lesonic fossils wercapparcnlly adaptedto brack

gayet+marshall+sempere 1991 mz-pc vertebrates of bolivia

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