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Home , 33rd parallel south, Carcharoides, Coquimbo Formation, Navidad Formation, Temblor Formation

Cainozoic Research, 3-18, 2006 4(1-2), pp. February

An Early elasmobranch fauna

from the , Central , America

³* Mario+E. Suarez Alfonso Encinas² & David Ward ¹,

'Museo Paleontoldgico de Caldera, Cousino, 695, Caldera, Atacama, Chile, [email protected]

2 Departamento de Geologia. Universidad de Chile, Plaza Ercilla 803, Santiago Chile. [email protected]

J David J. Kent ME4 4AW UK. Ward, School of Sciences, University of Greenwich, Chatham Maritime, Address for correspon-

dence: Crofton Court, 81 Crofton Lane, Orpington, Kent BR5 1HB, UK. E-mail: david@.ws. [* corresponding author]

Received 5 March 2003; revised version accepted 12 January 2005

A rich elasmobranch assemblage is reported from the Early Neogenemarine ofthe lower member of the NavidadFormation,

Central Chile. The fauna comprise Squalus sp., Pristiophorus sp., Heterodontus sp., Megascyliorhinus trelewensis. cuspi-

data, Isurus hastalis, Carcharoides and Cal- Odontaspisferox, oxyirinchus,Isurus hastalis, totuserratus, sp.

for the the Miocene The of lorhinchus sp., all ofwhich are reported first time in Early ofChile. presence Carcharoides totuserratus sup-

the Miocene for the lower ofthe basal Navidad Formation. The Chilean fossil elasmobranch fauna is ports Early age part representedby

deep water and shallow water taxa, which probably were mixed in a submarine fan. Certain taxa suggest warm-temperate waters. The

Early Miocene fauna from the Navidad Formation show affinities with other faunas previously reported from the Late and

Neogene of and New Zealand.

Se describe rica asociacion de fosiles veniente de los sedimentos marines del Inferior de la Formación una elasmobranquios pro Neogeno

central. La fauna Navidad, Chile comprende Squalus sp., Pristiophorus sp., Heterodontus spp., Megascyliorhinus trelewensis, Carcha- rias cuspidata, Odontaspis ferox, hums oxirinchus, Cosmopolitodus hastalis, Carcharoides totuserratus, Myliobatis sp. and Cal- lorhinchus todos los cuales el Mioceno de Chile. La de Carcharoides sp., son reportados por primera vez para Temprano presencia to-

edad miocena la basal del miembro inferior de la Formatión Navidad. Los tuserratus permite proponer una tempranapara parte compo-

de la fauna de fósiles formas de habrían sido mezcladas al nentes elasmobranquios incluyen agua profunda y somera que depositarse en

abanico submarino. la de La fauna de la Formatión Navidad un Algunos taxones sugieren presencia aguas temperadas-cálidas. presenta

afmidades faunísticas faunas de el tardfo de con otras elasmobranquios previamente reportadas para Paleógeno y Neógeno Argentina y

Nueva Zelandia.

Key words:: Miocene, Elasmobranch, Teeth, Chile Navidad Formation.

from Introduction remains this formation, commenting briefly on a

large tooth, which had been recovered from the Matanzas

First described by Darwin (1900), the Navidad Formation locality. That toothwas described as Carchariasgiganteus

also similar from is one of the largest onshore exposures of Tertiary sedi- Philippi and compared with a one La Her-

These fos- radura Both teeth mentary successions in western . locality (). were

“ siliferous have been known since the last illustrated in classic work Los outcrops century Phillipi’s (Phillipi, 1887)

Covacevic & through the work of a number of authors; Phillipi (1887), fosiles cuaternarios y terciarios de Chile".

(1942, 1968),Osorio (1976), Tav- Frassinetti cited the of in their Darwin (1900), Tavera (1980) presence Lanina sp.

Valenzuela Covacevic & invertebrate fauna recovered from the Punta Alta era (1979), Martinez & (1979), locality

Frassinetti (1980), Martinez-Pardo (1990), Troncoso and recently Suarez and Encinas (2002) yielded a brief

fossil (1991), Troncoso & Romero (1993), who described the report about the vertebrate assemblage from the

lower member of the Navidad Formation. presence ofbivalves, gastropods, scaphopods, brachiopods, echinoids, Bryozoa, barnacles, cephalopods, corals, fo- The present contribution is the result of several field trips raminifera, discoasters, , , tree trunks and carried out in the Navidad Formation during the years plant . Fossil vertebrates such as fish are, however, 1998-2002.These trips allowed us to gather a significant

collectionoffossil vertebrates, fish teeth, scarce or absent judgingfrom the major part of the litera- mainly including ture cited. Philippi (1887)provided the first description of an unexpectedly rich elasmobranch assemblage. -4-

Figure 1. Location of study area. A,B- Location of study area; C- Distribution ofthe Navidad Formation along the coast of Central

Chile; D- The Early Miocene Punta Perro and La Boca sections (indicated by T’ and ‘2’ respectively).

The material reported in this paper comprises ’ teeth, Historia Natural (MNHN) in Santiago, Chile. dental of and All plates rays, holocephalians. are new re- cords for the Navidad Formation.

Geological setting

Materialand methods The Navidad Formation was first described by Darwin

(1900). He assigned the term “ formation at

The material described in this paper was obtained by sur- Navidad” to all the sedimentary rocks exposed between face collecting from two sites located along the sea cliffs Estero Yali to the north and Punta Topocalma to the south. between Punta Perro and the town ofLa Boca (Figure Id). It consists of approximately 500 m of mudstones, silt-

isolated dental of and and which form It comprises shark teeth, plates rays, stones, conglomerates a succes- holocephalians. Additionalassociated vertebrate remains, sion of relatively undeformed strata. The Formation crops comprising both osteichthyan and cetacean teeth,were also out between 33° 45' S and 34° 15’ S latitude and 71° 30’ W

lh outside the All in the 6 collectedbut fall the scope of present work. longitude, 130kilometres southwest ofSantiago,

in for km north and the specimens described this paper have been deposited region ofChile, and extends 70 to south in the Seccion Paleontologla, of the Museo Nacional de 30 km east to west (Figure 1c). The Navidad Formation -5 -

crops out extensively along the coastal cliffs ofthe Navidad town of Rapel (M. Suarez, personal observation). During

region, the eroded banks of the Rapel River and the hill- the field trips we carried out in this area we found fossil

sides inland It (Chambers, 1985). rests unconformably vertebrates at several sea cliff sites between Estero

upon the Permo- crystalline , the Maitenlahue and the town of Pupuya.

Cretaceous Punta Topocalma Formation, or the Eocene Rfo In the northern extreme ofPunta Perro, fossil sharks’ teeth

Formation foundin the sandstone of intertidal Topocalma (Cecioni, 1978). are grey-green the zone,

Etchart (1973), Tavera (1979) and Ceccioni (1978) studied immediately above the basal conglomerate. At the southern

the ofthe Navidad Formation in detail.Etchart of Punta fossil teeth become stratigraphy part Perro, more common

divided the Miocene succession into the Middle (1973) along the sea cliffs, within thin, discontinuous lenses of

Miocene La Boca Formation and the Late Miocene Navi- varying lithology such as conglomerates and coquina beds

dadFormation. Tavera (1979) divided the Navidad Forma- with gastropods, bivalves and scaphopods. SouthofPunta

tion into three from oldest in the Punta Alta and the of members, to youngest being: Perro, locality near town Ma-

Lincancheo and Ceccioni Navidad, Rapel. (1970, 1978) tanzas (Figure Id), fossil teeth are scarcer and are associ-

defined the Navidad Group as formed by different units of ated with lithologies similar to those of Punta Perro (Tav-

Cretaceous- Within this he defined age. group era, 1979; Covacevic and Frassinetti, 1980; this paper).

the Miocene- Punta Perro-La Era Two mainfossiliferous sites Formation, were recognised which yielded

which is the Navidad Formation of Tavera. the elasmobranch equivalent to assemblages (Figure 1c). These are lo- Ceccioni divided this formationin four members: I, II, III cated along the sea cliffs between Punta Perro and La Boca

and IV. town and belong to the base of this formation. The first and

richest assemblage was recovered from a locality named

“La Boca,” to the northofLa Boca town. The second local-

of the Navidad Formation is referred Age ity to as “Punta Perro”, which is at the south of

Punta Perro. Two sections, where we have found the most

There is the of the Navidad some disagreement over age representative fossil assemblages, are describedin detail in

Formation. It has been described as Early Miocene based this section.

on macrofossils (Tavera, 1979) who suggested that the ob-

served palaeontological differences among members could

be attributedto facies variations.Osorio (1979) suggested a Stratigraphic sections

maximum of 11.5 Ma age based on ostracods. Martinez-

Pardo (1990) suggested a 19-10 Ma age based on plank- La Boca Section

tonic . Frassinetti & Covacevic (1981) sug-

Miocene for the 1 gested an Early-Middle age Navidad This site is located about km north ofLa Boca town and

member and based and is on stratigraphical correlation, as- shown as point 1 (Figure 2a). The section was measured

Late-Middle Late Miocene for the Lin- from the intertidal signed a to age zone to the top ofthe cliff.

cacheo member. These authors proposed the same Late- BASE: Base ofsection lies below sea level.

Middle Miocene for to Late age the lower part ofthe Rapel UNIT 1: (3.0 m). Conglomerate with granite, tonalite and

member. maficrock clasts that show variationin size from a huge up Recent in the lower of the investigations part Navidad to boulders exceeding one metre in diameter.The clasts are

Formationhave been carried out teams from the Colum- by well rounded in general with the exception of the bigger bia University, Universidad de Chile and the Museo Na- ones, which are angular. The rock is clast-supported, some-

tional de Historia Natural. Using Sr chemostratigraphy times a with sandy matrix rich in quartz, are scarce, 24 of shell material from lower they analysed samples the usually broken and consist mainly ofbivalves, gastropods

member of Punta Perro and from the Matanzas area and and solitary corals.

obtained of 20.2 ± 0.6 Ma for Punta Perro and an age an UNIT 2: (4.0 m). Medium to coarse-grained sandstone of 23.0 ± 0.6 Ma for Matanzas age (K. Gregory, pers. with some interbeddedmudstoneand siltstone. Sandstones

These last in with those deter- show comm.). ages are agreement planar cross bedding, current ripples, high-energy

mined the elasmobranch in this laminationand by assemblage reported parallel massive texture. Rip-up clasts, car-

(see conclusions below) and with those determined bonized wooden paper fragments and pumice clasts are common

by most ofthe formerauthors. in the sandstone and there are some water escape struc-

tures, as well as breccia formed by mudstone and pumice

clasts. Tuffaceous mudstone and siltstone are interbedded

Localities with the sandstone forming beds several metres long which

are broken and show syndepositional thrusts and are occa- Over the collectors looked for fossil inver- years, amateur sionally brecciated forming intraformational conglomer- tebrates in the Navidad Formation and sharks’ reported ates. Fossils are abundantand are concentratedin lenses of

teeth and teleost fish and cetacean remains. Fossil verte- dm scale within the sandstone. include bivalves, They gas- brates have also been farther inland from La small reported tropods, solitary corals, brachiopods, scarce cetacean Cueva and from locality (D. Frassinetti, pers. comm.) sev- remains, abundant elasmobranch teeth, leaves. Teredo

eral other still unstudied fossiliferous localities close to the bored logs and small branches. -6-

2. sections with the of the fossil A- Punta Perro B- La Boca section Figure Stratigraphical provenance elasmobranchs; section; -7-

UNIT 3: (3.0 m). Coarse to medium-grained sandstone subangular clasts of granite and volcanic rocks that passes

with some thin interbedded mudstone and siltstone layers. laterally into coarse-grained sandstone. In the rest of the

The unit has an erosive base forming a channel, several unit there are some thin horizons offine conglomerates that

metres wide cutting the lower unit. There are some thin show neithererosional contacts nor imbrication.At the top

conglomerate beds without erosive bases, composed of of the unit there are abundant ovoid concretions that are

angular quartz and lithic clasts interbedded with sand- usually joined forming levels parallel to the main bedding

stones. The sandstone shows upper flow regime parallel ofthe unit. There are some vertical burrows and very few

lamination, current ripples, mudstone and pumice clasts, fossils, mainly bivalves and gastropods.

concretions, and logs and branches of that are often TOP: Recent erosional surface.

drilled by Teredo. Siltstone and mudstonecontainabundant

leaves.

UNIT 4: (2.0 m). Orange, light brown, medium-grained Sedimentology

massive sandstones, occasionally with small pumice clasts,

The Miocene succession with marinefossiliferous which sometimes form spherical aggregates. begins a

out UNIT 5: (1.40 m). Medium-grained sandstones alternating conglomerate that crops only in the La Boca section

with siltstones in beds on a cm to dm scale. The sandstone (Figure la). The clasts are well rounded, in general mud-

shows flow lamination. stone siltstone condi- upper regime parallel or are lacking, indicating high-energy

UNIT 6: (8.0 m). Dark grey, medium-grained sandstone. tions. There is a huge range in clast size, from coarse sand-

meter They are massive in general but there are some levels that stone to boulders that exceed one in diameter. This

show high-energy parallel lamination.There are abundant conglomerate marks the beginning of the Early Miocene

marine that covered ofthe of concretions, cm to dm in diameter, that usually link up transgression most territory

forming concretionary beds with irregular forms parallel to Argentina and Brasil (Uliana & Biddle, 1988). It is inter-

beach where reworked alluvialfan sedi- the bedding of the unit. The fossil content is scarce and preted as a waves

concentrated in small lenses formedby abundantfragments ments. The presence of huge boulders alternating with of bivalves and gastropods. small clasts and sandstone indicates intense tectonic activ-

TOP: (2.0 m). Dark, fine-grained homometric sandstones ity. The succession continueswith mudstone, siltstone and

sandstone beds. The fossiliferous content is of with a modem soil developed upon. composed marine vertebrates and invertebrates and plant remains that

Punta Perro Section indicate both marine and continental influence. There are

This section is approximately 600 metres south of the abundant rounded pumice clasts in some layers that indi-

volcanic with the sedimenta- northern end of the Punta Perro promontory as point 2 cate activity contemporaneous

(Figure 2b). The section is measured from the intertidal tion. The presence of load structures and water escape

marks indicates .Brecciated zone to the top of the cliff. rapid horizons,

lamina- BASE: Base of section lies below sea level. large slumps, floating clasts, high energy parallel

tion and massive sandstone indicate UNIT 1: (2.50 m). Medium to coarse-grained quartz sand- gravity-driven trans-

port in whichslides, slumps, fallof clasts, stone with irregular wavy bedding. The unit yields some processes granu-

hard bands, cemented with calcite. The lar flows and turbidity currents are the main sedimentary very concretionary The interbedded siltstones and mudstones have fossil content is low and comprising logs, many bored by processes. foraminifera and ostracods of Teredo bivalves and gastropods. planktonic characteristic , sedimentation(Matmez and Valenzuela, 1979; UNIT 2: (2.15 m). Yellow-brown, medium-grained mas- deep-water

Osorio, We this of the succession to sive sandstone,rich in quartz. In the lowermostpart there is 1979). interpret part formed fossiliferous horizon with have been in a submarine fan. The sedimentation an 8 cm thick bivalves, gastro- would have been pods, solitary corals, cetaceans and elasmobranchs. This controlledmainly by gravity-driven proc-

esses alternating with the settling offine grains during quiet level gradually passes to the top to a 65 cm intermediate

This of the succession a concretion horizon. The top of this unit is formedby a 70 periods. part represents major,

rapid sea-level rise relative to the basal beach con- cm thick coarse-grained sandstone, with some granite and very glomerate. mafic clasts up to one metre in diameter. Some of these

clasts are well rounded,the bigger ones being sub-angular.

This part of the unit has abundant vertical burrows and a

few burnt wood fragments. Systematic Palaeontology

UNIT 3: (7.0 m). Light grey to white siltstone and mud-

of follows stone. They show rare small spherical concretions some (The terminology used here that ofCappetta, 1987)

cm to dm in diameter that usually contain plant fragments.

There are abundantostracods, gastropods, and planktonic

foraminifera. Class Chondrichtyes Huxley, 1880

UNIT 4: (15.0 m). Light yellow-brown, medium-grained Subclass Elasmobranchii Bonaparte, 1838

sandstone with load structures. Massive in general but also Order Squaliformes Compagno, 1973

showing high-energy parallel lamination.At the base there Family Squalidae Blainville, 1816

is a fine conglomerate, 1 to 2 metres thick, formed by Genus Squalus Linnaeus, 1758 -8-

3. b- (SGO-PV-877). Scale bar 1 cm. dorsal view, Figure a, Squalussp. (a) lingual view, (b) c-e- Pristiophoms sp, (SGO-PV-878; 879;

Scale bar 1 lateral view, mesial lateral view, f-h- 880). cm. (c) (d) view, (e) Squatina sp. (SGO-PV-881). Scale bar 1 cm. (f) lingual

view, occlusal basal view. I-I-Heterodontus Scale bar 5 (g) view, (h) sp. (SGO-PV-882). mm. (i) lingual view, (j) lateral view, (k)

occlusal basal view, -Heterodontus I view, (I) m-n sp.( SGO-PV-883; 884). Scale bar cm. (m) occlusal view, (n) occlusal view. -9-

Squalus sp. Family Pristiophoridae Berg, 1959

Figures 3a, b Genus Pristiophorus Muller and Henle, 1937

Material — One tooth

Pristiophorus sp.

Locality — La Boca Figures 3c-e

— The of this — Description crown tooth is large and both Material Three rostral spines lingual and labialfaces are very smooth. The cusp is trian-

gular in outline. The mesial cutting edge presents delicate Localities— La Boca, Punta Alta serrations, while those on the distal cutting edge and distal cusplet are coarser. The crown looks rather high in labial Discussion — One broken rostral spine from Punta Alta view, having more than two times the height of the root. was included in a faunal list as Lamna sp, by Frassinetti&

The apron, which reaches the floor, is well developed and Covacevic (1981). Recent examination of this material is backward-directed. In the showed that it is slightly lingual view, crown a pristiophorid rostral spine. Other well- presents a depression on the mesial edge of the uvula. The preserved pristiophorid spines fromLa Boca locality share latteris small but prominent and the infundibulumis rather the characteristic elongated andlaterally compressed crown large. The root is high having a row of foramina that are with anteriorand posterior cutting edges. In these teeth the sub-parallel to the limitof the crown-root. Belowthe limit peduncle is rather low and its base extended forward. The of the crown, under the mesial and distal regions, the sur- available material from the NavidadFormation, which in- face presents two well-marked depressions and dispersed cludes rostral spines of both juveniles and adult individu- foramina. In basal the is is here This view, root slightly concave expos- als, assigned to Pristiophorus sp. genus was

small foramina with distribution. ing some an irregular recently reported from the Miocene of the Bahia Inglesa

Formation(Suarez & Marquardt, 2001 j but, because ofits

Discussion — The and mesiodistally narrow apicobasally larger size, the northernspecimens are markedly larger than

of this that it is those described here. deep crown specimen suggests an upper Pristiophorus sp. have been reported tooth. The weak serrations on the mesial cutting edge and from the , (De Vries & De Muizon, the rather distal coarser cutting edges on the cutting edge 1985) and from the of Argentina (Cione & Ex- and the distal cusplet on the single Chilean Squalus tooth posito, 1980;Cione, 1988; Arratia & Cione, 1996). This is

it resemble cause to two serrate . the first occurrence of pristiophorids from the Early Mio-

1. S. weltoni Long 1992, from the Eocene La Meseta For- cene of Chile and the southernmost occurrence of this mation, of Island, Seymour (Long 1992a). This group from the western rim of South America.

listed species was originally as Squalus sp. by Welton

- (1978 unpublished PhD thesis) from the Late Eocene -

Oligocene of the Pacific coast of the USA. Order Squatiniformes Buen, 1926

2. Squalus occidentalus (Agassiz 1856) (= S. serriculus Family Squatinidae Bonaparte, 1838

Jordon & Hannibal, 1923) from the Middle Miocene of Genus Squatina Dumeril, 1906

California, USA. The in similarity age and morphology strongly suggest that the Chilean specimen belongs to this

However, more materialis needed before it can be species. Squatina sp. confidently assigned. Figures 3f-h

The Late Palaeocene species, Megasqualus orpiensis

(Winkler 1874) differs from the Chilean specimen in hav- Material — One tooth ing far weaker serrae on the distal cutting edge and distal

cusplet. Locality — La Boca

The earliestChilean squaloids are represented by teethand

fine to the and — spines belonging genus Centrophoroides Description This single tooth possesses a low cusp, comes from the Late Cretaceous (Maastrichtian) of the slightly recurved lingually and distally. The specimen is

Formation Suarez and Quiriquina (Suarez, 2001; Cappetta, elongated mesio-distally with convex lingual and labial

The record ofthe in South faces. The is almost flat 2004). Neogene genus Squalus root in basal view and exhibits a

America is and is limited scarce to material from the Early central foramen.

Miocene-LateMioceneofUruguay (Perea & Ubilla, 1989).

Suarez etal. the of — Recently reported presence Squalus sp. Discussion Teethof Squatina exhibit a very low degree fromMiddle-LateMiocene sedimentsofthe FormationLa of intraspecific variation, which makes it difficult to estab-

northernChile. This is the first of Portada, record Squalus lish species from isolated teeth. After comparing this tooth from the Miocene of Chileand South America. with those from ofextant of Early jaws specimens Squatina sp.

(probably Squatina armata, Phillipi) from Chilean waters

(M.S. collection), it was tentatively identifiedas a lateral

Order 1958 Pristiophoriformes Berg, tooth (third to six position). Both the mesial and distal cus- - 10-

massive in plets are more extant specimens and the latter lateralteeth ofa more common Francisci-group species (H.

has a more extended angle of lateral fallin the fossil form. vincenti (Leriche)). He described the anterior teeth ofthe

Such features, insufficient however, are to support a spe- Francisci-group species as a separate species; H. war-

cific determination and it be denensis 1966. can only designated as Casier, This was because, although from a Ward & Squatina sp. Bonavia (2001: 142) comment simi- rarer species, the large Portusjacksoni-group anterior teeth

tooth from larly regarding a single the Miocene of Malta. are much more readily surface-collected than the smaller

Prior to this report, Squatina sp. has been recorded from anterior teeth of Francisci-group species.

the Late Cretaceous (Maastrichtian) of the Quinquina For- The Chilean Francisci-group teeth closely resemble those

mation, centralChile (Suarez, 2001) and from the Middle (unnamed) from Round MountainSilt Memberofthe Mid-

Late Miocene of the Bahia Inglesa Formation, northern dle Miocene TemblorFormation, of Bakersfield, Califor-

Chile (Suarez y Marquardt, 2003; Suarez etal, 2004). This nia, USA ( Hill Bone Bed).

is the first record of Squatina from the Early Miocene of Considering the limited numberof available specimens for

Chile. comparison, we can only designate the Chileanmaterial as

Heterodontus sp. This genus has been previously reported

from the Neogene ofArgentina (Cione & Pandolfi, 1984),

Order Heterodontiformes Berg, 1937 from the Pisco Basin (De Muizon & DeVries, 1985), and Heterodontidae 1851 Family Gray, more recently from the Bahia Inglesa Formation, northern Genus Heterodontus Blainville, 1816 Chile (Suarez et al, 2004). This is the first record of Het-

erodontus in the Early Miocene of Chile.

Heterodontus spp.

Figures 3i-n Superorder Galeomorphii Compagno, 1973

Order Carcharhiniformes Compagno, 1973

Material — One anteriortooth and two lateral teeth. Family Megascyliorhinidae Pfeil, 1984

Genus Megascyliorhinus Cappetta & Ward, 1977

Localities — Punta Perro, La Boca.

Discussion — Reif (1976) divided the extant species of Megascyliorhinus trelewensisCione, 1986 Heterodontus into two groups, the Francisci-group, andthe Figures 5g-k

Portusjacksoni-group. The Portusjacksoni-group have ro-

bust clumpy anterior teeth that lack lateral cusplets in the Material— One tooth

adult, and have large ovoid molariformlateralteeth which

lack or have a much reduced longitudinal ridge. Teeth of Locality — La Boca

the Francisci-group of species have more gracile anterior

at teeth least of lateral and — bearing one pair cusplets, nar- Description This tooth is large having a crown strongly

lateral teeth with raised lon- rower, occlusally sigmoidal a recurved lingually and flanked by one pair of cusplets. The

anterior tooth from the has gitudinal ridge (Reif, 1976).The crown two cutting edges extending about one third of

Punta Perro locality (Figures 3i-l) is characterised a the crown’s below the by height apex. Both lingual and labial

smooth rounded surface and absence of the lateral faces ofthe exhibit vertical apical crown strong folds,being more

This is characteristic of the delicate cusplets. Portusjacksoni-group and sinuous on the former. The root is high and of Heterodontus Two lateral teeth wide and also species (Reif, 1976). shows a well-developed lingual protuberance

3m-n) rather and in occlusal and lobes (Figures are narrow sigmoidal two divergent separated by an incomplete medial

view. show a fromwhich In view the exhibits They prominent longitudinal crest groove. lingual root one pair of large parallel ridges radiate labially and lingually. The root is lateral foramina. low and basally flat. These are typical of the Francisci-

Discussion — Based its unusual group. on morphology, this tooth Two of this material interpretations are possible. can be immediately assigned to the genus Megascylio-

1. There is a species present that shares characters of both rhinus (Cappetta & Ward, 1977). Its size, the characteristic

groups. ornamentationof the crown, and the root (with an incom-

2. There are two species present. plete medial groove), indicates that the Chilean material

We are inclined towards the second possibility. Francisci- should be referred to the species M. trelewensis (Cione,

teeth are in the Cretaceous of 1986), The taxonomic of group present Early Europe. relationships this genus have been

Portusjacksoni-group teeth, the more derived condition, are subject to a number of differing opinions. It was first con-

in the Late Cretaceous of Kazakhstan and Uzbeki- present sidered to be a scylliorhinid by Cappetta & Ward, (1977)

stan Both but (David Ward, pers. observation). groups occur in later, Cappetta (1987: 113) proposed that the histology, the ofN.W. Paleogene Europe. Interestingly, Casier (1966: the possession ofan orthodont core, that it not suggest may in his 53), monograph on London Clay Fish, erroneously be a scylliorhinid. Specimens examined morerecently have associated the anterior teeth of an uncommon Portusjack- an osteodont histology characteristic of carchariniforms

(H. woodwardi, Casier) orectolobids soni-group species with the more and some (David Ward, pers. observation). - 11 -

4.a-i- Carcharias 886; 887; 888; Figure cuspidata (SGO-PV-885; 889; 890; 891). Scale bar 1 cm. (a) lingual view, (b) lingual view, (c)

labial view, (d) labial view, (e) lateral view, (f) lingual view, (g) lingual view, (h) labial view, (i) lingual view, j, k- Odontaspis ferox

(SGO-PV-892; 893). Scale bar 1 cm. labial view, (k) (j) labial view. 1- Carcharias sp. (SGO-PV-897). Scale bar 1 cm. lingual view,

m- Isurus Scale bar 1 cm. view, o- oxyrinchus (SGO-PV-894). lingual n, Cosmopolitodus hastalis(SGO-PV-895). Scale bar 1 cm.

(n) lingual view, (o) lingual view. - 12-

Cione (1986) described il- these the of Megascyliorhinus trelewensis, authors, teeth C. cuspidata can be separated

lustrating its osteodont with He from the modem C. due histology a radiograph. taurus to a major development of referred it to galeomorphii incertae sedis. Nishimoto & the root in the anterior. Carcharias cuspidata is known in Karasawa referred (1991) it to the orectolobiformes, as did the fossil record from the to Pliocene(Cappetta,

Purdy etal. (2001). Nishimoto & Karasawa that the Teeth of the argued 1987). genus Carcharias were recently re- flattened base and lack of labial root marginal foraminae ported from the Late Cretaceous (Maastrichtian) of the

are oforectolobiforms. However, these charac- Chile The suggestive Quinquina Formation, (Suarez, 2001). present is

ters also are present in many fossil carcharhiniform genera, the first record ofC. cuspidata from the Neogene ofChile.

including Premontrea, Palaeogaleus, Stenoscyllium, and

Scyliorhinus and are not unique to orectolobiforms. We

feel that the balance ofevidence supports the placement of Carcharias sp.

Megascyliorhinus within the carchariniformes, rather than Figure 41

the orectolobiformes.

This is from species easily distinguished the Eocene spe- Material — One tooth

cies M. cooperi and the Plio-MioceneM. miocenicus (see

Cione, 1986) by its coarser crown ornamentationand root Locality — La Boca

morphology (partially covered basal root groove).

The teeth of also known from the — Megascyliorhinus are Discussion Although this single large tooth lacks most

Neogene of New Zealand (Keyes, 1984; Pfeil, 1984). of its root and denticles, it preserves the typical fine stria-

The Navidad occurrence is the first fossil record of this tions on the lingual face of the crown that have been tradi-

in the Pacific of South America. genus margin tionally used to characterise the species O. acutissima

Such striations not in the (Agassiz, 1843). are present ge-

nus Odontaspis (Kemp, 1991; Ward, 2001) implying that

Order Berg, 1958 the Navidad material represents the genus Carcharias. Familia Muller & 1839 Odontaspididae Henle, Longbottom (1979: 59) assigned it to O. acutissima, two

Genus Carcharias 1810 teeth from the Miocene of Ecuador. These Rafinesque, specimens are

virtually identical with that discussed here from the Navi-

dad Formation, and we believe they should be referred to Carcharias 1883 cuspidata Agassiz, as Carcharias sp. Figures 4a-i

Material— Several teeth Genus Odontaspis Agassiz, 1838

Localities — Punta Perro, La Boca

Odontaspis ferox Risso, 1826

Description — All the specimens ofC. cuspidata from the Figures 4j-k Navidad Formation characterised are by a crown with complete or nearly complete cutting edges and a single pair Material— Two teeth of greatly reduced lateral cusplets. A large upper-anterior from tooth La Boca locality (Figure 4i) has a massive main Locality — La Boca that is flanked cusp by one pair of very small and recurved denticles. In lateral view the crown is to be — straight, tending Description The two specimens share a smooth cusp flexuous at the Its enameloid apex. lingual is smooth and its with incomplete cutting edges and exhibiting two pairs of lateral reach the foot of the well- cutting edges nearly lateral cusplets. One anterior tooth (Figure 4j) shows a

developed crown. One small tooth identifiedas a symphy- strongly lingually directed cusp with one first pair of high seal 4e-f) has a lateral and (Figures strongly sigmoidal cusp, one pair cusplets one greatly reduced second pair. A of small and massive root. A “crusher” very cusplets a pos- lower tooth (Figure 4k) has a cusp which is sigmoidal in terior tooth is root and (Figures 4g-h) small, having a heavy lateral view. In this specimen the first pair of lateral cus- lateral well-developed cusplets. Upper and lower teeth as- plets are twice the height ofthe second pair. signed to the same species (Figures 4a-d) have more deli- cate main characteristic of individuals. — cusps juvenile Discussion The teeth of Odontaspis can be separated

from those of Carcharias in that the former have more

Discussion — The teeth from the Navidad member are prominent and higher lateral cusplets. In addition, as is the

similar those ofC. illustrated very to cuspidata by Purdy et case with the Chilean material, the cutting edges of the al. (2001) from the River North Caro- Pungo Formation, crown are incomplete and the root lobes are deeply arched lina. Althoughconventionallyreferred to, and and narrow. Oliver Schneider superficially (1936) reported the presence similar the Recent to genus Carcharias the broad of teeth from the ofla , cusplets Odontaspis elegans Boca, seen in lateral teeth would that a southern suggest separate genus Lebu, Chile. Giventhat this material was not illus- would be more As out appropriate. has been pointed by trated and has been lost, it should be considered an unreli- - 13-

able record. in Odontaspis ferox occurs the Neogene ofthe Localities — La Boca, Punta Perro

Bahia Inglesa Formation, northern Chile (Suarez et al,

2003). This is the first record ofthis from the — The teeth have species early Description upper an unomamented

Neogene of Chile. crown with serrated The cutting edges. principal cusp is flanked by a single pair of lateral cusplets that are more

Genus Isurus Rafinesque, 1810 massive than its lower counterparts. One upper anterior

tooth (Figure 5 a) is asymmetrical and shows an erect crown

with an extreme apex slightly lingually directed. The labial

Isurus 1810 face oxyrinchus Rafinesque, of the crown is rather flat and the lingual face moder-

Figure 4m ately convex. In this tooth the lateral cusplets are very low

and The which worn. root, is rather low, has a poorly de-

Material— One tooth upper veloped lingual protuberance and short lobes. A tooth,

identified tentatively as an anterior (Figure 5b), lacks the

Locality — La Boca mesial and shows cusplet (due to breakage) a distally slight

curvature. The mesial edge is convex, while the distal one

Discussion — This tooth is almost identical with the is less The lateral sec- so. cusplets are more developed than in ond anterior tooth of dried of Isurus the tooth. upper a jaw oxy- previous A more lateral tooth (Figure 5c) is rinchus (MNHN collection). records ofthis Neogene spe- strongly distally directed, having a pair ofwell-developed

cies in Chile were previously reported by Long, (1993) and serrated triangular cusplets. The height ofthe cusplets from the ofEl Rincon and is about Neogene (Coquimbo Formation) a quarter the height ofthe main cusp. The lingual from the Bahia This also and labial face Inglesa locality. species seems to of the crown is smoothly convex. Three be abundant in the Late Mioceneof Pisco Peru Muizon lower teeth (De (Figures 5d-f) are symmetrical, having erect

& De Vries, 1985). serrated with slender conical cusps high, cusplets. The cus-

plets are very fine in more anterior rows, resembling the anterior teeth of Carcharias cuspidata or of Odontaspis.

Genus Cosmopolitodus Glickman, 1964 As in teeth from the upper jaw, the lateral cusplets of the

teethof the lower jaw become broader at more distal posi-

tions. The serrated ofthe edges lower teeth maybe incom-

hastalis 1843) in the anteriorteeth Cosmopolitodus (Agassiz, plete most (Figure 5d) or complete in 4n-o Figures more distal teeth (Figure 5f).

Material— — Two upper teeth Material Several teeth

— Locality La Boca Discussion — etal. Purdy (2001) referred the fossil genus

Carcharoides to the Recent carcharhinid genus Triaeno-

“ ”

Discussion — The is genus Cosmopolitodus used in rec- don. Ward & Bonavia (2001) rejected this assignment on that ognition the species hastalis is ancestoral to the great the basis of its dental formula, general morphology and white shark and not related the Makos Isurus with closely to ( ) histology, an opinion which we fully concur.

(Glyckman, 1964; Siverson, 1999 and Ward & Bonavia, The Chileanmaterial comprises serrated anterior teeth of

2001). the odontaspid-type and serrated upper lateral teethof the

et al. the Isurus for lamnoid Purdy (2001) use name xiphodon type which allows us to assign them to Carchar- broad-crowned specimens of hastalis. We agree with Ward oides totuserratus, Ameghino, 1901a (Cappetta, 1987). A & Bonavia in (2001), considering Isurus xiphodon as a second species, Carcharoides catticus (Philippi, 1846) has dubium.Whether there nomen was one or two coeval spe- non-serrated edges. The latter appears to be restricted to

cies of in the Pliocene is outside of the Cosmopolitodus Early Northern Flemisphere (Cione & Exposito, 1980; Cap- the of this however, it does petta, 1987) whereas Carcharoides is scope paper, appear very totuserratus, appar- hastalis unlikely. Cosmopolitodus (as I. hastalis) was pre- ently restricted to the (Cione & Ex-

viously reported from the Pisco Formation, southern Peru 1980).Its is the first posito, presence recorded for this spe- (De Muizon & De Vries, and from the El Rincon cies in Chile 1985) and its second occurrence in the Neogene of locality, Coquimbo Formation, Chile(Long, 1993). This is South America. the most abundant shark species in the Middle Late Mio-

cene of the Bahia Inglesa Formation, northern Chile

(Suarez etal, 2004) Superorder Batomorphii Cappetta, 1980 b

Order Myliobatiformes Compagno, 1973

Family Myliobatidae Bonaparte, 1838

Genus Carcharoides Ameghino, 1901

Carcharoides totuserratus 1901 Ameghino, Myliobatis sp.

Figures 5a-f Figures 5m-n - 14-

Figure 5. a-f- Carcharoides totuserratus (SGO-PV-898; 899; 900; 901; 902; 903). Scale bar 1 cm. (a-f) lingual view, g-k- Megascylio-

bar rhinus trelewensis (SGO-PV-904). Scale 1 cm. (g) oclusal view, (h) lingual view, (i) basal view, (j) lateral view, (k) labial view

Scale bar 1 occlusal view. Callorhinchus m,n- Myliobatissp. (SGO-PV-905). cm. (m) view, (n) posterior o-. sp. (SGO-PV-906).

Scale bar 1 cm. Occlusal view. - 15 -

Material— One medial tooth of Callorhinchuscf. callorhynchus from the Pisco Forma-

tion, Peru. However that material was not formally de-

Locality — La Boca scribed or illustrated. In Chile the oldest record of holo-

This isolated medial tooth characteristic fishes worn, presents a cephalian comes from Late Cretaceous rocks ofthe

which myliobatoid morphology, includes a hexagonal con- Quiriquina Formationin central and southern Chile (Stahl,

smoothocclusal surface and tour, a polyaulacorhizous root. 2001; Suarez, 2001b).

This material is the second and oldest record of holo-

Discussion — Taxonomical identificationsbased on iso- fishes cephalian in the Miocene ofChile and the first con- latedteethof difficultconsider- firmed of myliobatoids are extremely occurrence the group along thewestern margin of the wide dental variation found ing within the group (Wel- South America. ton & Zinsmeister, 1980; Nishida, 1990). Due to the frag- mentary nature of this material, we believe that it is not reasonable to assign this specimen to any given species Palaeoecology within Myliobatis. Long (1993) reported the myliobatoid

Aetobatus from the of El Rincon, (Co- Neogene locality Although precise depths have not been estimated yet, pre- and quimbo Formation) more recently (Suarez & Mar- vious authors who collected and studied microfossils from

2003) from the Middle- the lower member of the Navidad quardt, reported Myliobatis sp. Formation suggested a

Late Miocene of the Bahia Inglesa Formation. This is the deep environment of deposition for the sediments of this first occurrence of this genus in the Early Miocene of unit (Martinez & Valenzuela, 1979;Osorio, 1979). Osorio

Chile. (1979) recovered ostracods from the lower memberofthe

Navidad Formation and based on thepresence ofBradyela

normani, an species restricted to great depth, pro- 1832-41 Superorder Holocephali Bonaparte, posed a bathial environment for the deposition ofthe Navi-

Garman, 1901 Family Callorhynchidae dad Formation. He also suggested that the occasional oc-

1999 of Subfamily Callorhynchinae Stahl, currence some shallow water species ofostracods could

Genus Callorhinchus 1798 be Lacepede, interpreted as probable redeposition in a deeper facies.

Sharks from the genus Squalus, for example, may inhabit

boreal from the intertidal to temperatewaters ranging zone

Callorhinchus 900 in sp. up to m depth (Compagno, 1984; Purdy et al.,

5o Figure 2001). Compagno, 1984 (p. 112) also commented on the

distributionofthis shark, which seems to be correlated with

Material One fragmentary left mandibulardentalplate. a water temperature ranging between 7° to 8° C and 12° to

15° C. Among the extant sawsharks of the genus Pristio-

La Boca Locality phorus we can find some temperate forms that prefer shal-

low and estuaries bays and other forms such as the tropical

- The outlineofthe dental is Description incomplete plate species Pristiophorus schroederi that may occur on or near rather in The and the bottom from 640 square shape. symphyseal lingual margins at depths to 915 m (Compagno, little are a convex and the symphyseal margin is straight. 1984). The Squatina shows a broad distribution

There is median tritor. in both a single large, centrally placed, The cool temperateto tropical waters ranging in depth characteristic of the allows between morphology plate us to recog- the intertidal zone to over 1300 m (Compagno, nise it as one left dentary (See Suarez et al, 2004) 1984: 138).

Extant ofHeterodontus from species may occur the inter- Discussion After with dental tidal comparisons plates from zone to 275 m depth, but they are more common in

extantspecimens ofchimaeroid fishes from Chileanwaters waters shallower than 100m(Compagno, 1984; 154). The

cf. of (eg: Hydrolagus macrophthalmus, Hydrolagus affinis presence this shark in the lower memberofthe Navidad and Callorhinchus the Formation is remarkable because callorhynchus) (M.S. collection) presently it is an inhabi-

Navidad’s material can be assigned to the genus Cal- tant of warm-temperate and tropical waters that prefers lorhinchus with doubt. because of the any Also, overall temperatures above 21° C (Compagno 1984; 154). shape of its tritor, the fossil is very similar with Miocene The extant sand tiger shark Carcharias inhabits shallow, dental of described and and from plates Callorhynchus sp. recently temperate tropical waters the surface zone to a figured by Suarez et al. (2004) from the Bahia Inglesa maximum of 191 m depth (Purdy et al, 2001). On the other

northern Chile. the Formation, Regarding extant species hand, Odontaspis ferox occurs in warm temperate and these authors out the of the tritor is and it is believed pointed shape strongly tropical seas to prefer depths up to 420 m determinated the of the individualand is by ontogenetic stage (Compagno, 1984). Isurus oxyrhinchus a coastal and the variation. intraspecific It suggests that tritor shape can't oceanic shark occurring in both temperateand tropical wa- be used sufficient criterion as a to distinguish species ters down to at least 152 m (Compagno, 1984: 243). Like among the genus Callorhynchus. Consequently we as- other large lamniforms sharks, Cosmopolitodus hastalis

the new material Previ- signed only as Callorhynchus sp. was probably a cosmopolitan pelagic species during the

in De the ously (Cappetta Muizon, 1981) reported presence Miocene (Cappetta, 1987;Ward, 2001). Myliobatis shows - 16-

a wide rangeofdistributionbetweentemperate and tropical Pacific. The early Miocene elasmobranch faunas from the waters & Callorhinchus (Cione Exposito, 1980). is com- Navidad Formation are very similar with those reported

monly found in deep waters but it can move to relatively from the Neogene ofNew Zealand(1984) and Late Paleo-

shallow coastal waters of 16-38 m (Didier, 1993). gene-Early Miocene of Argentina (Cione & Exposito

Palaeoecological informationabout the two remaining fos- 1980).

sil sharks found in the lower member of the Navidad For- mation, Megascyliorhinus and Carcharoides, is difficultto obtain since their possible relationships with extant elas- General remarks and conclusions mobranch forms have not been satisfactorily clarifiedyet.

Based uponthe associated fauna and the nature of the de- The elasmobranch assemblage from the lower memberof

where posits Megascyliorhinus teeth have been found, the Navidad Formation is currently represented by seven

& Ward Cappetta (1977) suggested that this genus inhab- families: Squalidae, Pristiophoridae, Squatinidae, Hetero- ited of depths around 150-200m. A pelagic habitatfor this dontidae, , Myliobatidae and Callorhynchidae all

shark wouldbe possible considering that Cione (1986: 111) which are reported for the first time from the Early Mio- noted that the teeth of Megascyliorhinus are similar to cene of Chile. The presence ofSqualus and Callorhinchus those reported in the three recent filter-feeding sharks: the represents the first confirmed record of these two taxa in

megamouth shark pelagios (Taylor et al, the Early Miocene of South America. The occurrence of

Yabumoto the shark 1983; et al, 1997), whale Rhincodon Carcharoides totuserratus, an uncommon shark species

and the shark Cethorhinus. basking with a chronostratigraphical range restricted between the

The abundance ofteethofCarcharoides totuserratus in the Late Oligocene and Early Miocene (Suarez & Marquardt, lower member of the NavidadFormation, seems to be pro- 2003) supports a minimalEarly Miocene age for the basal observed portional to the abundance of Carcharias and it part of the lower member of the Navidad Formation. probably indicates that the former fossil species frequented Observations about habitat, distribution and biology of

shallow temperate-warmwaters. extant chondrychthians are certainly a useful tool for the

Most of the elasmobranch genera recorded in the lower understanding ofthe palaeoeological conditions ofits fossil member ofthe Navidad Formation the influenceof relatives from the Navidad Formation.Based suggest on a compari- warm-temperate waters. son with their living relatives, theelasmobranch faunafrom

the lower memberof the Navidad Formationrepresent dif-

ferent ecological habitats. Squalus, Odontaspis,

Palaeobiogeography Megascyliorhinus and Callorhynchus suggest relatively

deep waters while Heterodontus, Carcharias and probably

the fossil elasmobranch fauna Among of the Navidad For- Carcharoides totuserratus suggest shallow waters. The mation five we recognised genera that currently inhabit remains of sharks are likely deposited together in a subma-

Chilean waters. These are Squalus, Squatina, Isurus oxy- rine fan or turbidite. The presence of Heterodontus and

and Callorhinchus. Fromthe rinchus, Myliobatis remaining Odontaspis suggests warm-temperate waters. The latter is

Heterodontus and Carcharias known from in with evidence taxa only are agreement previously supported by mi- South American waters. Heterodontus occurs in waters of cropalaeontological information(Martmez-Pardo & Valen-

Peru and Ecuador while Carcharias is known from the zuela, 1979; Martmez-Pardo, 1990; Martmez-Pardo &

Atlantic margin of southern Brasil and Argentina (Com- Martmez-Guzman, 1997)and also by macropalaeontologi- pagno, 1984). The genus Pristiophorus, which at present cal data (Covacevic & Frassinetti, 1980). has not been recorded in South America, is common in In comparison with other southern Neogene elasmobranch waters ofNew Zealandand SouthernAustralia (Compagno, assemblages, the Navidad faunas indicate certainaffinities

1984). The fossil occurrences of Pristiophorus and Het- with those previously reported from the “Patagoniano” of erodontus, documented by De Muizon& De Vries (1985), Argentina (Cione & Exposito, 1980; Cione, 1986) and

Cione & Exposito (1980), Clone & Pandolfi (1984), Suarez those from New Zealand (Keyes, 1984; Pfeil, 1984).

& et al., Marquardt (2003), Suarez 2004) and by the pre-

sent work, indicate that during the Neogene and particu- Acknowledgments

larly the Miocene, sawsharks and bullhead sharks were

widespread along the Pacific andAtlantic margins of South We wouldlike to thank to Jose Valenzuela and Jose Valen-

America. The Early Miocene faunafrom the Navidad For- zuela Jr., Alexander Vargas, David Rubilar and Pedro

mation similar the faunas of fossil are to elasmobranchs Parra who collected part ofthe fossil specimens described

from the Late Oligocene and early Neogene fromArgentina here. Ruben Martinez and Daniel Frassinetti from the

reported by Cione and Exposito (1980) and Cione (1986), Museo Nacional de Historia Natural and Jacobus le Roux

the of Heterodontus, from sharing presence Pristiophorus, Isu- the Departamento de Geologfa de la Universidad de

rus Isurus Carcharoides totuserratus Chile read this A. Encinas hastalis, oxyrhinchus, critically manuscript. was sup-

and Megascyliorhinus trelewensis. The occurrence of the ported by Proyecto Fondecyt 1010691,Programa MECE

latter two species in the Navidad Formation is remarkable Educacion Superior UCH0010, Beca PG/50/02 ofthe De-

the limited record in South de de Chile. given fossil America,extending partamento Postgrado yPostitulo-Universidad

the Neogene geographical rangeof these taxa to the eastern - 17-

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