New multituberculate mammals from the Hauterivian/Barremian transition of Europe (Iberian Peninsula)

NEW MULTITUBERCULATE MAMMALS FROM THE

HAUTERIVIAN ⁄ BARREMIAN TRANSITION OF

EUROPE (IBERIAN PENINSULA)

by AINARA BADIOLA , JOSE I. CANUDO and GLORIA CUENCA-BESCOSGrupo Aragosaurus, Paleontologıa, Facultad de Ciencias, Universidad de Zaragoza, Pedro Cerbuna 12, E-50009 Zaragoza, Spain; e-mails: [email protected],

[email protected], [email protected]

Typescript received 18 March 2008; accepted in revised form 5 August 2008

Abstract: New multituberculate mammals from the Hauter-

ivian ⁄ Barremian transition of Europe are described. They

were found in the late Hauterivian-early Barremian fossilifer-

ous locality of La Cantalera (Josa, Teruel, Spain), one of the

Early Cretaceous sites in the Aragonese branch of the Iberian

Ranges, in northeastern Iberia. The fossils have been assigned

to at least three taxa on the basis of nine isolated teeth: a

new pinheirodontid taxon, Cantalera abadi gen. et sp. nov.;

a representative of the eobaatarid Eobaatar; a taxon described

as Plagiaulacidae or Eobaataridae gen. et sp. indet.; and other

as Plagiaulacida indet. These fossils have increased the reso-

lution of European Early Cretaceous multituberculate mam-

malian biostratigraphy and palaeobiogeography: the oldest

representative of Eobaatar is described here; a taxon is

assigned to ?Plagiaulacidae, in which case it would be the

first of this family in the Iberian Peninsula; and the discovery

of a new late Hauterivian pinheirodontid taxon demonstrates

greater biodiversity and a wider distribution for these multi-

tuberculates than was previously known. The mutituberculate

fauna of La Cantalera consists of endemic taxa (Pin-

heirodontidae), which were restricted to what is now Wes-

tern Europe, and others (Eobaataridae) which have also been

described in Asia. Consistent with the Iberian record of late

Barremian gobiconodontid mammals, the presence of Eobaa-

tar in Iberia with representatives from the late Hauterivian

to late Barremian, as well as in the Aptian or Albian of Mon-

golia, indicates that faunal exchanges between Europe and

Asia could have existed for most of the Early Cretaceous,

either sporadically or constantly.

Key words: Pinheirodontidae, Eobaataridae, ?Plagiaulacidae,

Early Cretaceous, Europe, systematics, palaeobiogeograhy.

When Pangaea broke up at the end of the Jurassic, a

significant separation between Laurasia and Gondwana

took place, causing increased differentiation of their

respective faunas. In the Early Cretaceous Laurasia was

divided into three palaeobiogeographic units: North

America, Europe and Central Asia (Russell 1993). These

landmasses, rather than being continuous, formed an

archipelago. The combined effect of plate movements and

eustatic sea level changes during the Early Cretaceous

(Skelton et al. 2003) allowed the coastal or shallow sea

areas to be used during sea level lows by the terrestrial

tetrapods to move between these three major continental

areas. The position of Europe, with its connections to

Asia, North America and Gondwana, makes it a challenge

to unravel the palaeobiogeography of terrestrial tetrapods

at this time. The expression ‘biogeographic ambiguity of

Europe’ defines this situation very well (Russell 1993;

Canudo 2006). The Early Cretaceous, therefore, is a key

period in investigating the geographical distribution, ori-

gin and evolution of many continental vertebrate faunas,

including mammals. To date, the most complete and con-

tinuous Early Cretaceous mammalian fossil record in

Europe has been found in the Iberian Peninsula, where

immigrant (Gobiconodontidae Chow and Rich, 1984)

and endemic faunas (the Dryolestidae Crusafontia cuen-

cana Henkel and Krebs, 1969) were present (Cuenca-

Bescos and Canudo 2003, 2004). Multituberculate fossils

dominate the mammal assemblages of the Iberian Penin-

sula (Canudo and Cuenca-Bescos 1996; Crusafont and

Gibert 1976; Cuenca-Bescos et al. 1996; Hahn and Hahn

1992, 1999, 2000, 2001, 2002), which also consist both of

endemic taxa (Pinheirodontidae Hahn and Hahn, 1999

and Paulchoffatiidae Hahn, 1969), with representatives

from the Late Jurassic, and others which, to date, are only

known in western Europe and Asia (Eobaataridae Kielan-

Jaworowska et al. 1987). New multituberculate finds from

the Iberian Peninsula (Canudo and Cuenca-Bescos 1996;

Cuenca-Bescos and Canudo 2004; Hahn and Hahn 2006;

Badiola et al. 2007a, b), together with those from Britain

(Sigogneau-Russell and Kielan-Jaworowska 2002; Butler

[Palaeontology, Vol. 51, Part 6, 2008, pp. 1455–1469]

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2008.00822.x 1455

and Hooker 2005; Sweetman 2006), have increased the

resolution of the Middle and Late Jurassic and Early

Cretaceous multituberculate biostratigraphy and biogeo-

graphy.

The most abundant and diverse Early Cretaceous multi-

tuberculate fossils are recorded in Western Europe, from

the Berriasian of Portugal and England and from the

Barremian of Spain. Fossils from the Valanginian are

scarce, and no Hauterivian specimens have previously

been described (Kielan-Jaworowska et al. 2004; Hahn

and Hahn 2006). In this paper, the first record of late

Hauterivian multituberculates in Europe is reported.

These have been recovered in the late Hauterivian-early

Barremian site of La Cantalera (Josa, province of Teruel),

one of the sites in the Aragonese branch of the Iberian

Ranges. The latter is becoming one of the most impor-

tant areas in the study of Early Cretaceous vertebrate

faunas, mainly multituberculates, because of the abun-

dant and stratigraphically well-correlated fossiliferous

beds. To date, late Valanginian, late Hauterivian and

early and late Barremian multituberculate fossils have

been recovered (Badiola et al. 2007a, b), of which those

of the Barremian are the most abundant and diverse

throughout the world (Canudo and Cuenca-Bescos 1996;

Cuenca-Bescos and Canudo 2004; Cuenca-Bescos et al.

1996; Ruiz-Omenaca et al. 2004). Further sampling in

the Iberian Range, which is currently being undertaken,

is expected to yield crucial information that will increase

what is known about the phylogeny, systematics, palaeo-

biology and distribution of the Early Cretaceous multi-

tuberculate faunas.

Geologically, La Cantalera site is an outcrop of Early

Cretaceous clays (‘Wealden facies’), specifically clays from

the limestones and marls of the Blesa Formation, located

in the Oliete Sub-Basin, one of the sub-basins of the Early

Cretaceous Maestrazgo Basin (Text-Fig. 1). This unit has

had a complex history, including a lower part with allu-

vial to lacustrine sedimentation (the site of La Cantalera).

The fossiliferous beds comprise grey clays with abundant

vegetal remains, ostracodes (Cypridea [Ulwellia] soriana

Kneuper-Haack, 1966) and gastropods (Viviparidae

indet., Basamotophora indet.) that are reported to be

TEXT -F IG . 1 . Geographical location and geological setting of the late Hauterivian–early Barremian site of La Cantalera (Oliete Sub-

Basin, Maestrazgo Basin). Modified from Aurell et al. 2004.

1456 P A L A E O N T O L O G Y , V O L U M E 5 1

resistant to periods of drought (Brookes Knight et al.

1960; Mayoral and Sequeiros 1983), and abundant car-

bonate nodules that were formed in palaeosols (Aurell

et al. 2004). There are also charophytes (Atopochara tri-

volvis triqueta Grambast, 1968 nov. com. Martın-Closas,

1996, Hemiclavator adnatus Martın-Closas and Grambast-

Fessard, 1986 nov. com. Schudack, 1989) and other mac-

roplant fossil remains. The presence of the charophyte

Atopochara trivolvis triqueta, subzone triqueta, indicates

that the fossiliferous beds are late Hauterivian–early Bar-

remian in age (Canudo et al. 2002). The site was probably

formed in a lucustrine environment with periodic

droughts. It is likely to have been a rich and diverse ter-

restrial ecosystem with a complex and varied faunal

assemblage and abundant vegetation in the surrounding

area (Ruiz-Omenaca et al. 1997). The highly diverse ver-

tebrate fossils are composed mainly of isolated teeth, dis-

articulated, complete or fragmented bones, eggshell

fragments, and coprolites. The vertebrate assemblage con-

sists of fishes (scarce fossils), frogs, lizards, crocodiles,

turtles, pterosaurs, dinosaurs and mammals (Ruiz-

Omenaca et al. 1997, 2005; Canudo et al. 2002). The

study of the vertebrate fossils from La Cantalera is still in

progress. To date, nine isolated teeth of multituberculate

mammals have been recovered, which are here assigned

to at least three different taxa.

Abbreviations

Institutional. FCPT, Fundacion Conjunto Paleontologico de Ter-

uel-Dinopolis, Teruel, Spain.

Localities. CAN, La Cantalera site.

Dentition. We use capital letters (I, C, P, M) for upper dentition

and lower-case letters (i, c, p, m) for lower dentition. Cusps are

labelled following Kielan-Jaworowska et al. (2004, text-fig. 8.28,

modified from Hahn and Hahn 1998a, text-fig. 1a): cusps of the

labial (b, B) and lingual (l, L) rows are numbered mesio-distally,

indicating their corresponding number and letter; a capital letter

(L, B) in the upper cheek tooth row and a lower-case letter (l,

b) in the lower cheek tooth row (e.g. B3: third labial cusp of an

upper tooth; l2: second lingual cusp of a lower tooth). The cusp

formula is that proposed by Kielan-Jaworowska et al. (2004,

p. 279), which is indicated by the number of cusps in consecu-

tive rows given from labial to lingual, separated by a colon. We

have here added the corresponding letter for each side for an

easier reading (e.g. 3B:4L).

Measurements. L, Length; W, width, which are given in mm. L

corresponds to the mesio-distal length measurement of the

crown, taken as oriented parallel to the mesio-distal axis,

whereas W is the maximum width measurement between the

labial and lingual margins of the crown.

Systematic use. The heterogeneous knowledge of multi-

tuberculates results in important unresolved problems

regarding their systematics and the interrelationships

among the different groups. In some of them, the whole

dentition is known in situ, whereas in others only a few

isolated teeth are present. In contrast, some taxa are rep-

resented by complete skeletons (Kielan-Jaworowska and

Gambaryan 1994). Consequently, cladistic analysis does

not produce reliable results, and different systematic

propositions have been addressed (Kielan-Jaworowska

and Hurum 2001 and references therein). Within the sub-

order Plagiaulacida, all authors consider that the families

are the real evolutionary and systematic units, definable

in each case by their special apomorphies. Here, therefore,

we shall not follow any particular systematic proposition

for plagiaulacids to explain the relationships between the

different families (e.g. Kielan-Jaworowska and Hurum

2001; Hahn and Hahn 2006) until new finds allow us to

make a more extensive and reliable phylogenetic study.

SYSTEMATIC PALAEONTOLOGY

Order MULTITUBERCULATA Cope, 1884

Suborder PLAGIAULACIDA McKenna, 1971

[pro Plagiaulacoidea Ameghino, 1889]

Family PINHEIRODONTIDAE Hahn and Hahn, 1999

Genus CANTALERA gen. nov.

Type species. Cantalera abadi sp. nov.

Derivation of name. After the name of the river nearby.

Diagnosis. A pinheirodontid described on the basis of

P4 ⁄ 5 and M1, which are characterized by having abun-

dant, long, and strongly-marked radiating enamel ridges

from the tip to the base of the cusps, which intermesh at

their distal ends with the neighbouring ridges; these form

a labyrinth-like structure on the floor of the central valley

on M1. P4 ⁄ 5 rectangular in outline and considerably

lengthened, a tooth longer than it is broad, with both

rows of cusps, 2B:4L, arranged clearly parallel to the

mesio-distal axis. M1 with a 2B:4L cusp formula, without

B3, with central valley open disto-labially, and with

prominent and small cuspules on the disto-lingual and

mesio-labial corners of the crown; the first one located

above a cingulum-like structure, which is strongly later-

ally protruding, and the second one placed above the

thick cingulum that encircles the mesial margin.

Differential diagnosis. In addition to the differences with

respect to other pinheirodontids apparent in the cusp

B A D I O L A E T A L . : N E W M U L T I T U B E R C U L A T E M A M M A L S F R O M T H E H A U T E R I V I A N ⁄ B A R R E M I A N T R A N S I T I O N 1457

formula and the presence of abundant, long and strongly-

marked radiating enamel ridges throughout the height of

the cusps on P4 ⁄ 5 and M1, which intermesh basally with

the neighbouring ridges, and form a labyrinth-like struc-

ture on the floor of the central valley on M1, Cantalera

gen. nov. can be distinguished from ?P5 of Sunnyodon

Kielan-Jaworowska and Ensom, 1992 by its larger size,

and by the absence of two labial cusps arranged symmet-

rically in the middle of the length of the tooth, with two

small mesial and distal cuspules; from P4 of the Pin-

heirodontidae gen. et sp. indet. from Portugal (Hahn and

Hahn 1999) by its larger size, and by the absence of a

labial row of cusps covering the whole crown longitudi-

nally, with three cusps instead of two; and from M1 of

Iberodon Hahn and Hahn, 1999 by its larger size, and by

the absence of an enlarged L3 and an additional row of

small cuspules on the lingual wall of the crown. Lavocatia

Canudo and Cuenca-Bescos, 1996 is like Cantalera gen.

nov. in having cusps covered by prominent radiating

enamel ridges but they do not intermesh basally with the

neighbouring ridges. Moreover, Cantalera is larger in size

than Lavocatia.

Cantalera abadi sp. nov.

Plate 1

Derivation of name. Dedicated to Jose Marıa Abad, who found

the site of La Cantalera.

Holotype. Left M1, FCPT (CAN 1 ⁄ 1609).

Paratype. Left P4 ⁄ 5, FCPT (CAN 1 ⁄ 934).

Referred material. A fragment of right M1, FCPT (CAN

1 ⁄ 1610).

Diagnosis. As for genus, monotypic.

Description

M1. FCPT (CAN 1 ⁄ 1609) (holotype; L: 2.15; W: 1.52) exhibits a

well-preserved crown. It is nearly rectangular in outline, with a

2B:4L cusp formula. The lingual row of cusps is slightly broad-

ened distally. The labial row of cusps is shortened, with only

two cusps. B3 is missing; consequently, the central valley opens

disto-labially. The strongly-marked and long radiating enamel

ridges, which extend from the tip to the base of the cusps, inter-

mesh basally with the neighbouring ridges, forming a labyrinth-

like structure on the floor of the central valley. At the base of

the disto-lingual corner of the crown, there is a strongly laterally

protruding cingulum-like structure with a small cuspule. A small

cuspule is also present mesio-labially above the thick cingulum

which encircles the mesial margin. FCPT (CAN 1 ⁄ 1610) is a

fragment of a right M1. There is only the mesial portion of the

crown preserved, which is very similar in morphology to that of

the complete M1 described above. The first two lingual cusps

(L1 and L2) and the first labial one (B1) are present. These are

densely covered by thick and prominent radiating enamel ridges.

P4 ⁄ 5. FCPT (CAN 1 ⁄ 934) (paratype; L: 2.16; W: 1.33) is a P4

or a P5. Small portions of its mesio-labial, disto-labial and dis-

to-lingual corners are broken. Neither of the roots is preserved.

The tooth is rectangular in outline in occlusal view and consid-

erably lengthened mesio-distally. The cusp formula is 2B:4L.

Both rows of cusps are oriented clearly parallel to the mesio-dis-

tal axis. The apices in some cusps are also partially broken (B2

and L2). The lingual row of cusps covers the whole crown longi-

tudinally, whereas the labial one is considerably shortened, with

only two cusps confined to the mesial half of the crown. On the

distal margin of the crown, only the last lingual cusp (L4) is

present. This is confined to the disto-lingual corner, but due to

its large size it occupies almost the whole distal margin. The

cusps exhibit different sizes; their heights increases considerably

distally; L4 and B2 are the widest and highest cusps. Cuspules

are absent. The radiating enamel ridges are strongly developed.

They are long and prominent and cover the cusps densely from

the tip to the floor, intermeshing basally with the neighbouring

ridges. The tooth is wider in the middle than at the mesial and

distal margins. Both mesial and distal margins are gently

rounded. The mesial margin bears a thick but faint cingulum.

Comparison. The absence of the disto-lingual ridge on M1 pre-

vents it from being assigned to Plagiaulacidae Gill, 1872, Eobaa-

taridae, Allodontidae Marsh, 1889, Zofiabaataridae Bakker, 1992,

Arginbaataridae Hahn and Hahn, 1983, or to genus Glirodon

Engelmann and Callison 1999 (family incertae sedis). The disto-

lingual ridge is lacking, by contrast, in Paulchoffatiidae and

Pinheirodontidae, as in the M1 from La Cantalera. The latter

exhibits a 2B:4L cusp formula, with the absence of B3, and the

central valley opens disto-labially. These dental features are also

present on M1 of the paulchoffatiids Kuehneodon simpsoni Hahn,

1969 and Pseudobolodon oreas Hahn, 1977, and in M1 of the pin-

heirodontid Iberodon quadrituberculatus Hahn and Hahn, 1999.

These multituberculate fossils were found in the Late Jurassic and

Early Cretaceous (Berriasian) deposits from Portugal, respec-

tively. The morphology of the M1 from La Cantalera, however, is

closer to the pinheirodontid taxon than the paulchoffatiid taxa

EXPLANATION OF PLATE 1

Scanning electron micrographs of gold-palladium-coated teeth of Cantalera abadi gen. et sp. nov. (Pinheirodontidae,

Multituberculata) from La Cantalera (Josa, Teruel), Spain. A–E, left M1, holotype FCPT (CAN 1 ⁄ 1609). F–J, left P4 ⁄ 5, paratype FCPT

(CAN 1 ⁄ 934). A, F, occlusal; B, G, mesial; C, H, distal; D, I, lingual; and E, J, labial views.


PLATE 1

BADIOLA et al., Cantalera abadi, gen. et sp. nov.

A B C

D

F G H

I J

E

1 mm

mentioned above. One of the diagnostic characters of pinheir-

odontids is the presence of the last cusp in each row of M1 either

enlarged or reduced. Both are reduced in I. quadrituberculatus,

whereas in the M1 under study, L4 is enlarged and B2 reduced.

There are other differences between the M1 of I. quadritubercula-

tus and the specimen from La Cantalera (see differential diagno-

sis). The low cusp which is present above the cingulum-like

structure in the M1 under study, located in the disto-lingual cor-

ner of the crown, could be an incipient radiating ridge like that

of the Berriasian pinheirodontid Sunnyodon notleti Kielan-Jaw-

orowska and Ensom, 1992 of the Purbeck Group of Dorset, Eng-

land, or an incipient row of small cuspules like that of I.

quadrituberculatus. It seems evident that it is not the typical fully

developed disto-lingual ridge of the eobaatarids. Moreover, eoba-

atarids have an M1 that is 3B:4L instead of 2B:4L.

Elongate and molariform upper premolars usually belong to

P4 or P5. The basic cusp formula of P4 in the suborder Plagiau-

lacida is 2-3B:4L, although most of them exhibit 3B:4L on P4

and P5, with the exception of Paulchoffatiidae, which have the

third row of cusps (row bb sensu Hahn and Hahn 2004: 120) in

P4 and P5, and of Pinheirodontidae, Albionbaataridae Kielan-

Jaworowska and Ensom, 1994, and the Allodontidae Morrison-

odon brentbaatar (Bakker, 1998), which exhibit a P5 with the

additional cusp row bb. Therefore, our specimen without row

bb and cuspules, and with a cusp formula of 2B:4L, could

belong to a P4 from a pinheirodontid taxon, or to a P4 or P5

from one of the following allodontid and eobaatarid taxa: the

Late Jurassic (Tithonian) allodontid Psalodon potens (Marsh,

1887) of North America, the Early Cretaceous eobaatarid Eobaa-

tar Kielan-Jaworowska et al. 1987 of the Iberian Peninsula (early

and late Barremian) and Asia (Aptian or Albian), or the late

Early Cretaceous (Albian ⁄ Cenomanian) eobaatarid Janumys ere-

bos Eaton and Cifelli, 2001 of North America. The presence of a

rectangular outline and cusps with considerably different heights,

however, allows us to assign the upper premolar of La Cantalera

to Pinheirodontidae instead of the other families mentioned

above. There are other differences too. In the P5 of P. potens the

inner flank of the lingual row is worn, forming a shearing edge;

the P5 of Eobaatar is characterized by having its main row of

cusps (the lingual one) obliquely oriented to the mesio-distal

axis, forming a cutting edge: consequently, the distal margin of

the crown is formed only by the lingual row of cusps; and the

P4 of J. erebos is clearly shorter and narrower than the tooth

from La Cantalera, and its cusps are less distinctly separated

from one another. The upper premolars of pinheirodontids

which have been interpreted as P4 do not have the third addi-

tional cusp row bb. They exhibit two rows of cusps, of which

the labial one is shorter than the lingual one, as in the premolar

from La Cantalera. We therefore assign it to the family

Pinheirodontidae. However, our specimen could belong either to

a P4 or a P5, because there is an upper premolar tentatively

assigned to the fifth premolar in the pinheirodontid Sunnyodon

notleti that does not have the row bb (Kielan-Jaworowska and

Ensom 1992, p. 107). We thus assign the premolar under study

to P4 ⁄ 5. The occlusal morphology of the P4 ⁄ 5 from La Cantal-

era is close to the Berriasian taxa S. notleti and Pinheirodontidae

gen. et sp. indet. from Portugal (Hahn and Hahn 1999) in hav-

ing cusp-rows oriented parallel to the mesio-distal axis, though

this character is more accentuated in the premolar from La Can-

talera. Other differences between them have also been described

(see differential diagnosis).

The great density and prominence of the radiating enamel

ridges observed in the teeth of Cantalera abadi gen. et sp. nov.

also exist in M1 of the albionbaatarid Proalbionbaatar plagiocyr-

tus Hahn and Hahn, 1998b and in P5 of the pinheirodontid Lav-

ocatia alfambrensis Canudo and Cuenca-Bescos, 1996, which

come from the Late Jurassic of Portugal and Early Cretaceous

(early Barremian) of Spain, respectively. However, the upper

premolar from La Cantalera differs from that of L. alfambrensis

(see differential diagnosis), and the M1 of La Cantalera can also

be distinguished from that of P. plagiocyrtus by its clearly differ-

ent number and arrangement of cusps. P. plagiocyrtus has a

greater number of cusps, and their apices are displaced some-

what to the labial side. The specimens under study belong to a

new pinheirodontid genus, which is characterized by a number

of autopomorphic characters (see diagnosis).

Family EOBAATARIDAE Kielan-Jaworowska et al. 1987

Genus EOBAATAR Kielan-Jaworowska et al. 1987

Eobaatar sp.

Plate 2

Referred specimens. Left P5, FCPT (CAN 1 ⁄ 935), and other left

P5 partially worn FCPT (CAN 1 ⁄ 1611).

Description

P5. FCPT (CAN 1 ⁄ 935) (L: 1.63; W: 1.35) exhibits a well-pre-

served crown, except that the apices of some cusps (L2-L4 and

B2) are slightly broken. Neither of the roots is preserved. The

cusp formula is 2B:4L. The main row of cusps, the lingual one,

is obliquely oriented to the mesio-distal axis. It extends from the

mesio-lingual corner towards the disto-labial corner of the

crown. The labial row of cusps is shortened, with only two cusps

(B3 is missing), and the disto-labial portion of the crown is con-

siderably narrowed, comprising only the last lingual cusp (L4).

Consequently, the occlusal surface is distinctly narrow, with a


Scanning electron micrographs of gold-palladium-coated teeth of Plagiaulacidae or Eobaataridae gen. et sp. indet. (Multituberculata)

from La Cantalera. A–E, left P1 ⁄ 3, FCPT (CAN 1 ⁄ 936). F–J, fragment of a right p4, FCPT (CAN 1 ⁄ 937). A, F, occlusal; B, mesial; C,

H, distal; G, mesio-labial; D, I, lingual; and E, J, labial views.


PLATE 2

BADIOLA et al. Eobaatar sp.

A

B

C

D

F G

H I

E

1 mm

triangular outline. In the lingual row of cusps, L2 and L3 are the

widest and highest cusps, whereas L1 and L4 are similar to one

another in size. In the labial row, B2 is considerably bigger than

B1. The cusps are conical, well separated from one another, and

covered by short and thin radiating enamel ridges. The latter are

only present on the cusps. There are no cuspules. The lingual

wall of the tooth extends below the cusps to form a smooth and

wide surface or wing (sensu Kielan-Jaworowska et al. 1987,

p. 12), which is strongly laterally protruding. Lateral to the two

labial cusps, the labial wall likewise forms a smooth wing,

although it is narrower and only mesio-labially protruding. The

mesial margin of the crown is somewhat concave in the middle,

whereas the distal margin is convex. Both the labial and lingual

walls are roughly concave in the middle. FCPT (CAN 1 ⁄ 1611) is

the other left P5, with severe wear on the lingual and distal mar-

gins of the crown. The last lingual cusp (L4) is worn away. Con-

sequently, the crown length value could be smaller than the

actual measurement (L: less than 1.96; W: 1.33). The morpho-

logical features of this specimen, however, are similar to those of

the P5 described above.

Comparison. The crown outline, and the cusp formula and cusp

arrangement of the P5 teeth under study are typical of the eo-

baatarid Eobaatar: the cusp formula is 2B:4L; the lingual row of

cusps is obliquely oriented to mesio-distal axis; the labial row of

cusps is shortened, and the distal portion of the crown is consid-

erably narrowed, comprising only the lingual row of cusps; and

lateral to the cusps of the labial row there is a small wing. The

size of the better-preserved P5 from La Cantalera (L: 1.63; W:

1.35) is similar to that of Eobaatar magnus Kielan-Jaworowska

et al. 1987 (L: 1.7; W: 1.2 in each of the four specimens) from

the Aptian or Albian beds at Khovboor (Gobi Desert, Mongolia),

but the eobaatarid of La Cantalera differs from E. magnus by the

absence of lingual cusps distinctly increasing in height distally,

and by having lateral to the cusps of the lingual row a smooth

and wide wing that covers the whole lingual margin longitudi-

nally and is strongly laterally protruding. The lingual wing in

E. magnus is not present or is incipiently developed disto-

lingually, though it is clearly less laterally protruding than the

one from La Cantalera. The P5 from La Cantalera seems smaller

than the holotype specimen of Eobaatar hispanicus Hahn and

Hahn, 1992 (Hahn and Hahn 1992: text-fig. 5; L: 1.92, W: 1.2)

of Galve (early Barremian), and differs from it in the absence of

lingual cusps distinctly increasing in size distally, and by having

a narrower occlusal surface, which exhibits a clearly developed

triangular outline, and strongly laterally protruding lingual and

labial wings. However, because the differences between the P5

specimens from La Cantalera and the P5 of E. magnus and

E. hispanicus are only slight or the features above mentioned

could be variable intraspecifically, other lower and upper teeth

must be studied in detail in order to be able to conclude that

the morphological differences observed between the premolars

from La Cantalera and those of E. hispanicus and E. magnus are

sufficiently great to regard the studied specimens as belonging to

a different species.

Plagiaulacidae or Eobaataridae gen. et sp. indet.

Plate 3

Referred specimens. Left P1 ⁄ 3, FCPT (CAN 1 ⁄ 936), and frag-

ment of a right p4, FCPT (CAN 1 ⁄ 937).

Description

P1 ⁄ 3. FCPT (CAN 1 ⁄ 936) is an anterior upper premolar, proba-

bly P2 or P3 (L: 1.06; W: 0.68). A small portion of both roots is

preserved. The crown is well preserved. It is roughly rectangular

in shape in occlusal view, with labial and lingual distal bulges,

and somewhat elongate mesio-distally. There are two lingual and

two labial cusps, which do not form clear longitudinal lingual

and labial rows. The cusps are similar in size, although B1 is

slightly smaller than the others and is displaced towards the

mesio-labial corner of the crown. B2 is confined to the midpoint

of the labial margin and placed transversally between the two

lingual cusps. The cusps are covered by thin and radiating

enamel ridges. These are short and present only on the cusps.

The mesial margin is roughly rounded, and two very small cus-

pules are present. The distal margin is convex due to the pres-

ence of a talonid-like ‘‘heel’’. The lingual margin is somewhat

concave in the middle, whereas the labial one is concave between

B1 and B2 and between the B2 and the talonid-like ‘‘heel’’.

p4. FCPT (CAN 1 ⁄ 937) is a fragment of a right p4 (L: bigger

than 2.26; W: 1.24). It cannot be known exactly how many cusps

and serrations are present on the cutting edge, because the tooth

is broken mesially. The projection of the first serration forwards

with a reasonable downward curvature suggests that at least a

quarter part of the mesial margin may be missing. The preserved

serrations are roughly of the same length, have the same distance

between them, and are bent similarly downwards. The first two

and last two serrations are worn or less marked than the others.

A prominent and dorsally worn shelf-like concave structure is

located close to the disto-labial corner. This structure has been

called the posterior wear shelf by Kielan-Jaworowska et al.

(1987: 6), and it seems to belong to several worn labial cusps

instead of only one. These could have been small and fewer in

number than the cusps on the cutting edge. The absence of


Scanning electron micrographs of gold-palladium-coated teeth of Eobaatar sp. (Eobaataridae, Multituberculata) from La Cantalera.

A–D, left P5, FCPT (CAN 1 ⁄ 935). E–I, left P5 with severe wear on the lingual and distal margins of the crown, FCPT (CAN 1 ⁄ 1611).

A, E, occlusal; B, F, mesial; G, distal; C, H, lingual; and D, I, labial views.


PLATE 3

BADIOLA et al. Plagiaulacidae or Eobaataridae gen. et sp. indet.

A

B C

D

F

GH

I J

E

1 mm

1 mm

horizontal erosion on the cutting edge indicates that the premo-

lar could have been used for cutting, instead of for shearing.

Comparison. The morphology, outline and number of cusps of

the anterior upper premolars are quite similar in the Plagiaulaci-

dae, Eobaataridae, Allodontidae and Arginbaataridae, where they

are short, rounded and tricuspid, whereas in the Paulchoffatiidae

and Pinheirodontidae they are tri- to tetracuspid, with or with-

out additional very small cuspules. Anterior upper premolars are

unknown in Hahnodontidae Sigogneau-Russell, 1991 (Hahn and

Hahn 2004, tables 1, 7). Nevertheless, some exceptions with

respect to the number of cusps and the arrangement of cusp-

rows have been described. Pentacuspid or hexacuspid P1 ⁄ 3 are

present in ?Plagiaulacidae gen. et sp. indet. (Kielan-Jaworowska

and Ensom 1992, pl. 5, figs. 5–6) and in the paulchoffatiid Kie-

lanodon hopsoni (Hahn, 1987), whereas in others there appears a

transitional stage towards P4, with two clear longitudinal rows of

cusps (usually in paulchoffatiids, e.g. Pseudobolodon krebsi Hahn

and Hahn, 1994) or even with three longitudinal cusp-rows (in

albionbaatarids). However, only the plagiaulacid Parabolodon

elongatus (Simpson, 1928), the eobaatarid Eobaatar? pajaronensis

Hahn and Hahn, 2001, and the specimen described here exhibit

a P1 ⁄ 3 mesio-distally lengthened with cusps not arranged in

clear longitudinal rows. The morphology of the P1 ⁄ 3 from La

Cantalera differs from P. elongatus and Eobaatar? pajaronensis for

the reasons set out below, and seems to belong to a different

taxon. The P1 ⁄ 3 under study can be assigned either to Eobaa-

taridae or to Plagiaulacidae. The size of the P1 ⁄ 3 from La Can-

talera is similar to that of E.? pajaronensis, but it differs from it

in having four cusps instead of three, a distal margin that forms

a fully developed talonid-like ‘heel’, which is missing in E.? paja-

ronensis, and the last lingual cusp confined to near the disto-lin-

gual corner. In E.? pajaronensis, by contrast, this has been

displaced towards the middle, alone forming the distal margin of

the crown. P. elongatus is like the P1 ⁄ 3 from La Cantalera in

having a similar talonid-like ‘heel’ behind the cusps, and in the

last lingual cusp being confined to near the disto-lingual corner.

However, the premolar from La Cantalera is smaller than the

smallest premolar (P3) of P. elongatus, and there are four cusps

instead of three. The presence of more than four cusps, but less

than 15, on the cutting edge in the p4 fragment from La Cantal-

era and the fact that the second serrations that is not distinctly

longer than the others allow us to assign it to Eobaataridae or

Plagiaulacidae, as opposed to Paulchoffatiidae, Pinheirodontidae,

Allodontidae, Zofiabaataridae, Glirodon, or Arginbaataridae. The

morphology of the p4 fragment, however, is closer to a p4 of the

Plagiaulacidae than of the Eobaataridae. In the p4 specimens

known within the family Eobaataridae (Eobaatar and Sinobaatar

Hu and Wang, 2002), only one labial cusp is located close to the

distal border of the crown. In contrast, the size of the worn

shelf-like concave structure of the p4 from La Cantalera suggests

that possibly more than one labial cusp existed in the original

unworn tooth, as is the case in plagiaulacids. Therefore, the p4

and the P1 ⁄ 3 from La Cantalera could belong to the family Pla-

giaulacidae rather than to the Eobaataridae. However, because of

the fragmentary state of the p4 under study and the absence of

other tooth specimens, we have assigned the P1 ⁄ 3 and p4 in

question neither to a specific family nor to a specific taxon.

Plagiaulacida indet.

Text-fig. 2

Referred specimens. Fragment of an I2, FCPT (CAN 1 ⁄ 939), and

fragment of a ?ix, FCPT (CAN 1 ⁄ 938).

A B

1 mmC D

TEXT -F IG . 2 . Scanning electron micrographs of gold–palladium-coated teeth of Plagiaulacida indet. (Multituberculata) from La

Cantalera. A–B, fragment of an I2, FCPT (CAN 1 ⁄ 939). C–D, fragment of a ?iX, FCPT (CAN 1 ⁄ 938).


Description

I2. FCPT (CAN 1 ⁄ 939) is a fragment of an I2 (L: c. 0.94; W:

0.53). Its distal margin is broken, and we cannot see how many

cusps there were. The cusp of the mesial margin is the highest

and widest cusp, and this seems the only functional one. The

labial flank of the crown is convex, whereas the lingual one is

broken.

ix. FCPT (CAN 1 ⁄ 938) is a fragment of an ix. There is a very

small fragment of the crown and another of the root preserved.

Neither the enamel development nor the crown morphology can

be described in this specimen.

Comparison. The I2 and ix seem to correspond in size to the

same multituberculate taxon described above for P1 ⁄ 3 and p4 or

for P5. However, they are too fragmentary for it to be possible

to observe the diagnostic characters of these incisors (the num-

ber of distal cusps on the I2 and the enamel development in the

?ix) in order to assign them to a specific family within the sub-

order Plagiaulacida. Therefore, they are provisionally described

as Plagiaulacida indet.

PALAEOBIOGEOGRAPHICALIMPLICATIONS

The late Hauterivian multituberculate fossil assemblage

from La Cantalera consists of some taxa (Pinheirodonti-

dae and probably ?Plagiaulacidae) that were restricted to

what is now Western Europe, whereas others (Eobaatari-

dae) have also been described from Asia. The two premo-

lars described as Plagiaulacidae or Eobaataridae gen. et

sp. indet. could belong to a new plagiaulacid taxon, in

which case it would be the first in the Iberian Peninsula.

To date, these multituberculates have only been described

from the Berriasian deposits of England (Owen 1871;

Simpson 1928; Kielan-Jaworowska and Ensom 1992,

1994; Sigogneau-Russell and Kielan-Jaworowska 2002;

Hahn and Hahn 2004). Pinheirodontids are described

from the Berriasian deposits of England and Portugal

(Kielan-Jaworowska and Ensom 1992; Hahn and Hahn

1999), and in the late Hauterivian (this paper) and early

and late Barremian deposits of Spain (Canudo and

Cuenca-Bescos, 1996; Cuenca-Bescos et al. 1996; Badiola

et al. 2007a). Eobaatarids are present in Europe (England

and Spain) and Asia (China, Japan and Mongolia). Those

from England are located in the Early Cretaceous (Weal-

den) deposits of Cliff End (Wadhurst Fm., south Eng-

land) and of the Isle of Wight (Wessex Fm.), which are

dated as Valanginian and Barremian, respectively: at the

Cliff End site, Loxaulax valdensis (Woodward 1911) has

been described from a few specimens (Clemens 1963,

1971; Clemens and Lees 1971), and two Barremian speci-

mens from the Wessex Formation of the Isle of Wight

have also been assigned to the genus Loxaulax Simpson,

1928, without specific determination, by Butler and Ford

(1975). New multituberculate fossils have been recovered

in this area, the study of which by Steven C. Sweetman is

still in progress. Two species of the eobaatarid Eobaatar

are described from the early Barremian of Spain: Eobaatar

hispanicus and Eobaatar? pajaronensis (Hahn and Hahn

1992, 2001, 2002). New specimens which belong to Eoba-

atar have recently been identified in the late Barremian

bed at Vallipon (province of Teruel), in the Aragonese

branch of the Iberian Ranges, in northeastern Iberia

(Badiola et al. 2007a). Some of these were previously

described as Eobaatarinae indet. (Cuenca-Bescos et al.

1996). In this paper, the oldest representative of Eobaatar

is described. It comes from the late Hauterivian–early

Barremian site of La Cantalera (Josa, Teruel), another site

in the Aragonese branch of the Iberian Ranges. The age

of the Asiatic localities that have yielded multituberculate

fossils is still subject to debate. Such fossils come from

the Jehol and Tetori Groups of China and Japan, and

from the Gobi Desert in Mongolia. The only eobaatarid

taxon described from the Jehol Group, in the Yixian For-

mation, is Sinobaatar lingyuanensis Hu and Wang, 2002.

The Jehol Biota is currently thought to date from about

130–120 Myr (Meng et al. 2006 and references therein),

and so is Barremian to Aptian in age (Gradstein et al.

2004). There are also two other formations (the Shahai

and Fuxin formations; e.g. Wang et al. 1995) that are

stratigraphically younger than the Yixian Formation,

which have yielded some multituberculate fossils that are

still unpublished (Kusuhashi, pers. comm. 2007). Two

other species of Eobaatar are described from the Aptian

or Albian beds at Khovboor (Gobi Desert, Mongolia; Kie-

lan-Jaworowska et al. 1987): Eobaatar magnus and Eobaa-

tar minor Kielan-Jaworowska et al. 1987. The age of the

fossiliferous beds located in the Japanese Tetori Group is

more obscure, because there are no good radiometric

dates available and reliable index fossils are rare. The age

of the Kuwajima Formation ranges from Valanginian to

Aptian (Matsukawa et al. 2006; Rougier et al. 2007),

although the Zircon U-Pb ages collected from tuff beds

indicate a shorter period, Barremian or early Aptian

(Kusuhashi et al. 2006). The multituberculate remains of

the Kuwajima Formation were previously described as Eo-

baataridae indet. (Takada et al. 2001), but recently have

been assigned to new eobaatarid taxa (Kusuhashi 2008).

The occurrence of Eobaatar in the Iberian Peninsula,

from the late Hauterivian to late Barremian, and in Asia

during the Aptian or Albian, indicates that some faunal

exchange could have taken place between these areas dur-

ing the Neocomian. The palaeogeographic models indi-

cate that it was possible (Smith et al. 1994; Smith and

Rush 1997). Other palaeontological data support this

hypothesis, as well as the vertebrate faunal exchange


between other parts of Laurasia and Gondwana. The

mammalian family Gobiconodontidae is recorded in the

?Berriasian of Africa (Morocco), in the early and late

Barremian of Europe (England and Spain), in the Hauter-

ivian to Aptian of eastern and central Asia, and in the

Aptian–Albian of North America (Cuenca-Bescos and

Canudo 2003; Sweetman 2006; and references therein),

and some Barremian dinosaur taxa (at generic level) are

observed to be common to the Iberian Peninsula, Asia

and North America, indicating intermittent faunal

exchange between these areas of Laurasia during the Bar-

remian (Milner et al. 2000; Cuenca-Bescos and Canudo

2003; Canudo 2006). However, the presence of Eobaatar,

together with the presence of an euhelopodid dinosaur, in

the late Hauterivian deposits of La Cantalera, in the Ibe-

rian Peninsula (Canudo et al. 2002; Ruiz-Omenaca et al.

2005), suggests that an Early Cretaceous geographical

connection between Europe and Asia could have existed

from the Hauterivian to the Barremian (for c. 11 myr),

either sporadically or constantly for most of the Early

Cretaceous, thus making mammalian and dinosaur faunal

exchange possible.

The area of origin of the family Eobaataridae is still

unknown. The paucity of the fossil record for these multi-

tuberculates, and the absence of well-dated deposits in

Asia, currently inhibits the use of cladistic analysis for

investigating the phylogenetic and palaeobiogeogaphic

relationships between the different taxa. However, the

oldest representative of Eobaatar at La Cantalera, together

with those of the early and late Barremian found in other

Iberian localities, as well as the species from Asia, will be

essential in studying the area of origin and the dispersals

of this eobaatarid.

CONCLUSIONS

The multituberculate fossil assemblage of La Cantalera

(Teruel, Aragon), one of the sites in the Aragonese

branch of the Iberian Ranges in northeastern Iberia,

indicates that the Early Cretaceous (late Hauterivian)

multituberculate fauna of the Iberian Peninsula was

fairly diverse. The fossils consist of endemic western

European taxa (Pinheirodontidae) that are known from

the Late Jurassic and other taxa (the eobaatarid Eobaa-

tar) that were originally described from the Aptian-Al-

bian deposits of Mongolia, Asia. On the basis of the

palaeobiogeographic interpretation of certain mammals

(Gobiconodontidae) and dinosaurs (some Ornithopoda)

a European-Asian connection during the late Barremian

has been previously suggested. New multituberculate

finds from the Hauterivian ⁄ Barremian transition of the

Iberian Peninsula, at La Cantalera site, suggest that the

Early Cretaceous geographical connection between these

areas could have existed from the late Hauterivian,

either sporadically or constantly for most of the Early

Cretaceous, thus making mammalian and dinosaur fau-

nal exchange possible.

Acknowledgements. We thank Drs A. Currant and J. J. Hooker

for their help and giving access to specimens in the Natural His-

tory Museum of London. The journal referees (one of them was

Dr Zofia Kielan-Jaworowska) of this paper provided helpful

comments on an earlier version of the manuscript. Dr Nao

Kusuhashi (Institute of Vertebrate Palaeontology and Palaeoan-

thropology, Chinese Academy of Sciences, Beijing, China) pro-

vided information on the Asian multituberculate faunas.

Financial support was provided by the Ministerio de Educacion

y Ciencia (MEC) (research projects CGL2004 ⁄ 03393 ⁄ BTE and

CGL2007 ⁄ 62469 ⁄ BTE), the Universidad de Zaragoza (‘Financ-

iacion de Grupos Consolidados’ 2008), and the Gobierno de Ar-

agon (Direccion General de Patrimonio Cultural de Gobierno de

Aragon and Programa Europa XXI de Estancias de Investigacion

CAI-Gobierno de Aragon) of Spain. A. Badiola also acknow-

ledges support from the Programa Juan de la Cierva of the

Ministerio de Educacion y Ciencia of Spain.

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New multituberculate mammals from the Hauterivian/Barremian transition of Europe (Iberian Peninsula)

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