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Bivalves du Trias tariquide de Los Pastores

(Algésiras), Espagne. Leur signification

Suzanne FRENEIX

Laboratoire de Paléontologie, UMR 8569 du CNRS,

Muséum national d’Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

Freneix S. 1999. — Bivalves du Trias tariquide de Los Pastores (Algésiras), Espagne. Leur

signification. Geodiversitas (21) 2 : 137-146.

MOTS CLÉS

Bivalves (mollusques).

Trias,

lariquides,

Algésiras,

Espagne,

taxinomie,

paléobiogéügraphie,

biostratigraphic,

paléoécologie.

RÉSUMÉ

La dalle calcaro-dolomitique Fossilifère, découverte dans le roc de Los

Pastores, à Pouest d’Algésiras contient deux lits, plus carbonatés, à faunule de

bivalves. Ceux-ci se présentent en valves dissociées de moules internes a la

surface supérieure de la dalle et de moules externes incomplets, non détermi¬

nables, à la surface inférieure. Sept espèces, en concordance avec les especes

trouvées dans d’autres localités d'Espagne, ont été identifiées : ? Geri>illia

joleandiy Leptochondria albertis ? Posidonia obliqua-, Costatoria {Costatorta)

goldfussiij Lyriornyophoria suhlaevis, Neoschizodus (N.) iaevigatus,

Psetidocorbula gregaria. Après une révision taxinomique, la signification de

cet aSvSemblage est considérée : (1) selon la distribution stratigraphique de ces

espèces, parmi les domaines trîasiques germanique, alpin-téthysien et

d’Espagne, Page suggéré est I.adicn supéricur-Carnien inférieur ; (2) quelques

données de corrélations ressortent du biofaciès précédent souvent cité dans le

Trias des Catainnides et des cordillères bétiques du domaine <• sépharade

Cependant, le biofaciès à Lyriomyophoria stiblaevis prédominante paraît carac¬

téristique de ce gisement méridional d’Andalousie ; (3) cette thanatocénose

reflète un paléoenvironnement marin, peu profond, de haute énergie.

GEODIVERSITAS • 1999 • 21 (2)

137

Freneix S.

ABSTRACT

Bivalves from tbe Taricjuide Trias of Los Pastores (Algesiras), Spahi.

A fossiliteroiis liinestone slab from the 1 riassic (ocalit)' of the Roch of Los

Pastores (Algcsiras W), yieldcd two beds of a bivalve fauniile. The bivalves are

preserved as dissociated valves of internai nioulds on ihc upper sidc of the

slab and as iinidenrifîabic exrcrn.il moiilds on its lower side. Seven spccies,

similar with taxa knowatrom urher localitlcs in Spain bave been Identified:

}GervilHa joleaueit, Leptoihoiulria alberti, ^Posicloma obliqua. Castaioria

{Costatoria) goldjùssii, Lyriomyophoria sublaevist Neoschizodus {N.) laevigatus,

Pseudocarbulugt'egaria. 7 hc signifîcance of ihis assemblage îs considered after

a systemutic siudy: (1) uccording to the stracigrapliiail distiibutioti of these

seven spccies. particularly .imong rhe Germanie, the Alpine-tethyan and the

Spanish Triassic realms. the âge stiggcsted is upper badinian-lower Caniian;

(2) somc corrélations appc.ir with the association: "^.Gervilba joleaudi,

Leptochûudriii aiberti. Neoschizodus laevigaïus. Myophoria goldfussii,

Lyriomyophoria sublaevisx this last speeies is known from sevcral fossüerous

localiries in the Catalanid and Beric Ranges into the ‘^Sephardic” rcalm.

Howcvcr, ihc siudied bivalve assemblage» with the dominant speeies

Lyriomyophoria sublaevis, shows a characicristic biofacics. endémie of this

Southern province of Andalusia; (3) rhe Lyriomyophoria biofacies» with its

condition of préservation, signifies a high energy, shallow, marine palco-

envjronincnt.

INTRODUCTION STRATIGRAPHIQUE ET

STRUCTURALE

La dalle carbonatée ayant livré les bivalves» objet

de ce travail, provient d’un niveau du Trias

appartenant à une succession mésozoïque origi¬

nale qui apparaît dans les rochers de Los

Pastores, à la sortie ouest de la ville d’Algésiras.

Cet affleurement d’un-demi kilomètre carré est

traversé par la route N-34Ü vers Tarifa. Il s'inter¬

cale tectoniquement dans les flyschs crétacés-

paléogènes du Campo de Gibraltar.

Los Pastores, le roc de Gibraltar et le Jebel

Moussa (Rif marocain) sont situés de part et

d’autre du Détroit de Gibraltar. Ils ont été ras¬

semblés (Durand-Delga 1972) dans un même

ensemble tectonique et paléogéographique, les

« Tariquides D’appartenance strucntrale discu¬

tée, cct ensemble se situe près du contact majeur

qui sépare les Aines externes (flyschs allochrones)

des zones internes (Dorsale calcaire), dans Parc

qui unit cordillères bétiques et Rif.

Le Trias de Los Pastores (Baudclot et al. 1993,

fîgs 1, 2) comprend de bas en haut :

1. marnes et gypses clairs (plus de 80 m) ;

2. pélites colorées (20 m) ;

3. grès clairs (20 m) à stratifications obliques ;

4. péliics colorées (30 m) avec quelques interca¬

lations de bancs de dolomies plus ou moins cal-

careuses ;

5. dolomies grises à passées pélitiques sombres

(30 m), que surmontent de puissantes dolomies

(Trias supérieur) et calcaires (Lias inférieur) dans

le rocher de Gibraltar.

Des associations palynologique.s ont été dégagées

des termes 2 et 4 (Carnien inférieur) et du

terme 5 (Carnien moyen). L’horizon grès carbo¬

nate à bivalves couronne une barre (2 m) formée

de bancs calcaro-dolornitiques amalgamés. Elle se

place aux deux tiers supérieurs du terme 4, dans

des pélites lie de vin et verdâtres. Le point fossili¬

fère étudié est à environ 200 m au sud du carre¬

four entre la route N-340 et la voie du Poligono

KEYWORDS

Bivalves,

Triassic,

“Tariauides”,

Algesiras,

Spain.

taxonomy,

pal CO b i ogeogra P h)’,

biosrraiigraphy.

paleoecology.

138

GEODIVERSITAS • 1999 • 21 (2)

Bivalves triasiques d’Andalousie

Industrial. En 1988, avant que la zone ne soit

recouverte de déblais des grandes carrières, au

sud de la route, la barre à bivalves se suivait,

plongeant à l’est en série normale, en conservant

ses caractères.

L’étude paléontologique entreprise est la suite

logique de la publication sur le Trias des

« Tariquides » (Baudelot et al. 1993).

TAPHONOMIE DES BIVALVES

Une dalle calcaro-dolomitique lossilifère, mesu¬

rant 35 cm X 23 cm et 7 cm d’épaisseur, a été

extraite de la l-ormation triasiquc du gisement

précédemment cité par M. Durand-Dclga et

M. Esteras au cours d'une mission géologique

récente. Celte dalle présente, sur chacune de scs

faces horizontales, un lit coquillicr à mollusques

bivalves. A la face supérieure, les coquilles

incluses dans cette assise gisent à ptac la face

convexe orientée vers le haut, en valves dissociées

et en fragments au nombre de quatre-vingts envi¬

ron. Cet assemblage, suppose-t-on, a été déposé

par des courants à la surface du sédiment, les

valves désunies trouvant une position d'équilibre

sur leur face concave. A la surtace inferieure de la

dalle, les bivalves, plus rares, ne sont observables

que par leur face interne dont il ne subsiste que

le contour ou quelques fragments d’empreintes

externes. Ces derniers ne sont pas déteiminables.

Par contre, sept espèces onr été identifiées (dont

deux avec doute) dans l’horizon supérieur, en

concordance morphologique exrerne avec celles

publiées pur les auteurs espagnols, car leur état de

conservation, très incomplet, ne donne pas accès

aux caractères internes.

SYSTÉMATIQUE PALÉONTOLOGIQUE

Pour le matériel examiné, Pétude systématique

comprend une synonymie restreinte, la distribu¬

tion srratigraphique et paléobiogéographique des

espèces, plus particulièrement, leur distribution

connue en Espagne.

Ce matériel est déposé au Laboratoire de

Paléontologie du Muséum national d’Histoire

naturelle de Paris (LPM, MNHN).

Classe BIVALVIA Linné, 1758

Sous-classe PTERIOMORPHIA Beurlen, 1944

Ordre PTERIOIDA Newell, 1965

Sous-ordre PTERJINA. Newell, 1965

Supcrfamille PTl:IU.^CEA Gray, 1847

Famille Bakevui liiDAK KJng, 1850

Genre Genùllia Delrance. 1820

? Gervillùi joleaudi Schm'idu 1935

(Fig. 1 A. B)

Gennilia sp. - Joleaud 1912 : 77, pl. 1.

Gennlleui joleaudi Schmidt, 1935 : 53, pl. 4, figs 17,

18, 20.

Gerviliia joleaudi Schmidt - Lerman 1960 : 34, pl. 3,

figs 14, 15 - Mâiquez-Aliaga, Hirsch & l.opez-

Garfido 1986 : 210, figs 4E, F. - Màrquez-Aliaga &

Momoya 1991 : pl. 2, fig. 3 (3).

MATt-RlF-l.. — Six spécimens de taille réduite

(n'* L]*IVl-R. 62091J, de 15 à 25 mm de diamètre

antéro-posiéneur, de 5 â 7 mm de diamètre timbono-

vcnrral. cerninenienr juvéniles.

DiSTRIBUTlON. — Trias moyen d’Algérie : Anisien-

Ladinicn intérieur d’Isracl. En Espagne : cordillère

Ibérique, Ladinien moyen (Baléares), Ladinien supé¬

rieur d’Hcnarejos (faune de Cuenca), Chelve (faune

de Valence) ; Ladinien supérieur-Carnien de la cor¬

dillère Bérique, Prébérique de Murcie (Cehegin),

Subbérique de Jaén (Hornos Siles)...

Remarques

Leur assignation hypothétique est fondée sur la

forme étroirc légèrement incurvée ventralement,

sur l’nngic de 25” environ du bord dorsal et de

Taxe d'allongement de la coquille et sur les stxies

corn marginales distales en relief de la région pos¬

térieure et sur l'aile po.stérieure obtuse. G. joleau-

di « forma juvenis » Schmidt (1935 : 56, pl. 4,

fig. 19) du Langobardien d’Espejeras est peut-être

proche de nos exemplaires, scs dimensions sont

de 14 mm de longueur et de 4 mm de largeur.

Superfamille PECTINACEA Rafinesque, 1815

Famille AviCULOPECTlNlDAE

Mcek & Hyaden, 1865

Sous-famillc Aviculopectininae

Meek Hayden, 1865

Çimee. Leptochondria^ixxxvex-, 1891

GEODIVERSITAS • 1999 • 21 (2)

139

Freneix S.

Leptochondria alberti (Goldftiss, 1836)

(rig. ic)

Monotis alberti Çio\àï\xi^s, L836 : 138, pi. 120, fig. 6a,

Pecten inaeqiihmatiis Goldfuss - Wurm 1911 : 102,

pl. 6, fîgs 8-10.

Velopecten albenit Goldfuss — Schmidt 1935 : 61,

pl. 4, fïgs 30, 31.

Chlaruys { Velatd) alberti Goldfuss — Virgili 1958 : 464,

pl. 8, Ag. 2.

Peeten alhertii Goldfuss — Lerman I960 : 40> pl. 4,

% 14.

Leptochotulria alberti (Guldfuss) — Marquez-Aliaga,

Hirsch & Lopez Garrido 1986 ; 212, fig. 4G (avec

synonymie). — VUrquez-Aliaga & Montova 1991 :

118, pl. 2, fig. 3 (2b, 2c, 2d).

MatERIRI.. — Un fragmeru d’une valve gauche

(n'’ LPM-R. 62092) de 20 mm environ de hauteur,

présentant des costules radiales de deux ordres ; celles

du second ordre apparaissent par intercalation. Ce

fragment se rapporic sans aucun doute ;l L. alberti^

espèce très commune au \ rias.

Dlstributk'im. — Trias germanique (Buntsandstein à

Lettenhohle) et IVias alpin (Scythien à Carnien).

Anisien-Ladinien inférieur de I ransjordanie, d'I.sraël.

En Espagne : cordillère Ibérique, Ladinien d’Esporlas

(Baléares), terme carbonate supérieur du Muschelkaik,

faune de Teriicl (Royuelâ), de Cucncu (Henurejos) ;

cordillère Bétique, Subbétique de Jaén.

Famille POSIDONIIDAE Frech, 1909

(jcnre Posidonia Bronn, 1828

? Posidonia obliqua Haucr, 1857

(Fig. ID)

Posidonomya obliqua Haucr, 1857 : 152, pl, 2, fig. 9.

- Philipp 1904 ; 94, pL 6, figs 23, 24.-Wurm 1913 :

574, dI. 19. fig. 6.

Posidonia oblictm Hauer - Virgili 1958 : 451, pl. 6,

fig-1‘

Posidonia obliqua Hauer in Philipp — Ichikawa 1958 :

187, pl. 23. fig. 6.

MatëRIEI. — Une valve gauche et trois valves incom¬

plètes (iT LPM-R. 62093).

Distribution. — Trias alpin (Ladinien des Alpes du

Sud). En Espagne : Muschelkaik supérieur des

Catalanides (Composines).

Re.M ARQUES

La valve gauche ovalaire, de 20 mm de longueur

et 13 mm de hauteur, d'indice des diamètres

L/H = 1,5, se rapproche des dimensions données

par V^irgili avec L/H variant de i,4 à 1,5, tonre-

fois de caille plus petite. L'ornementation est

composée de huit ou neuf cordons (ou lamelles)

concentriques assez espacés, épais, irréguliers ; le

bord postérieur est oblique, mais le spécimen a

été déformé par compaccion.

Sous-classe RA1.AEOHETERODONTA

Newell. 1965

Ordre TRIGONK.’)IDA Dali, 1889

Sous-ordre TRIGONIINA Dali, 1889

Superfamille MYOIMIORIACEA Bronn, 1849

Famille MyüFHORIIÜAH Bronn, 1849

(ou Famille CtiSTATORlIDAE

Newell ik Boyd, 1975

= MlNETRicîONitDAF Kobayashi, 1954)

Genre Costatoria Wangen, 1906

Sous-genre Cosxaxoria s.s.

Costatoria (Costatoria) goldfussii

(Alberti /// Zieten, 1830)

(Fig. lE)

Trigonia goldfiessii Aberti in Zieten, 1830 : 94, pl. 71,

fig. 1.

Myoplymit ktliani Schmidt 1935 : 79, pl. 5, figs 31,

32.

Myophoria goldfitssi (Zieten) — Virgili 1958 : 480,

fig. 58 n* 9.

Costarorta (CostatoriaJ goldfitssi (Alberti in

Zieten) — Allasina/ 1966 : 69b, pl. 50, figs 7-10.

Costatoria gûldfusd (Alberti in Zieten) — Cox 1969 :

473, fig. D 6.3, 3a. - Tamura 1972 : 69, pl. 1. figs 17-

20. - Marquez-Aliaga, Hirsch ik I.épcz Garrido

1986 : 216, fig. 4B. C. - Marquez-AHaga ôc Montoya

1991 ; 120, pl. 1, figs 2, 7, pl. 2, fig. 4. - Pérez Lôpez,

Fernandez, Solé de Porta & Aldrquez-Aliaga 1991 :

144, pl. 1, figs 2, 5,7.

MaîTriiU. — Une valve droite, une valve gauche et

un fragment d’empreinte externe (n“ LPM-R, 62094).

Dl.STKItlL 1 It'i.V. — Espèce curasiatique (depuis

l’Europe jusqu*en .Asie). Miischelkalk-Kcuper inlériciir

d’Allemagne ; l.adiuien-Carnien des Alpes du Sud. En

Espagne, l.adinien supérieur-Carnicn inférieur de la

cordillère Ibérique : provinces de Teniel, G.uenca et

Valence, branche Castellane (diverses formations dtiiit

celle de Royuela), cordillère côtière catalane.

140

GEODIVERSITAS • 1999 • 21 (2)

Bivalves triasiques d’Andalousie

Fig. 1. — A. B, ? GervUlia joleaudi Schmidl. deux valves gauches (LPM-Ft 62091) ; C. Lapiochondria alberti (Goldfuss). valve

gauche (LPM-R. 62092) : D. ? Posidonia obliqua Hauer. valve gauche iLPM-R. 62093) : E. Costatoria {Cosîatona) goldfussii (Alberli

in Zieten), valve droite (photo de gauche) et Lynomyophona sublaevis (Schmidt), valve droite (LPM-R. 62094) :

F, G, Lyriomyophoria sublaevis (Schmidt) , F, valve gauche , G deux valves droites (LPM-R. 62095 LPM-R. 62096) ,

H. Neoschizoaus {Neoscmzoous) laevigatus (Goidfussj. valve gauene (LPM-R. 62097) ; I. J, Pseudocorouia gregana (Munster in

Gotdfuss) ; L valve gauche, photographie du haut (LPM-R. 62098) : J. autre valve gauche (photo du bas) avec Lyriomyophoria

sub/aev/s (LPM-R. 620099). Clichés Denis Serrette (LPM, MNHN). Échelles : 1 cm.

GEODIVERSITAS • 1999

Freneix S.

Remarques

La valve droite la mieux conservée, de dimen¬

sions : H = 12 mm, L = 12 mm, est de forme tri-

gono-ovalc avec un crochet prosogyre, de

position antérieure ; le flanc a dix à dou7;e côtes

radiales légèreinent granuleuses que séparent des

intervalles lisses plus larges que les côtes.

Quelques côtes fines apparaissent pre.s de Tumbo

par intercalation. Des costules radiaires peu

apparentes s’observent au bord de t’aréa niai

conservée.

Genre Lyriomyophoria Kobayashi, 1954

Lyriomyophoria snblaevis (Schmidt» 1935)

(Fig. lE, F, K)

Myophorid sublaedis Schmidt, 1935 ï "^8, pi- 5, figs 27-

30. - Virgili 1958 : 478, pl. 11, % 3.

Lyriomyophoria sublaiw (_Schinidt) — Marquez-Aliaga

& Montoya 1991 : 122. pl. 1, fig. 3. - Marquez-

Aliaga & Martine-/ 1996 : 108.

MATÊRiEL. — Onze valves gauches, trois valves droites

assez différentes des formes typiques décrites par

Schmidc d’Espejeras (Alicante) et de Celicgin

(Murcie) du Ladinien supérieur. Elles représentent un

morphotype nouveau de rang infrasubspécifique.

Distribu I ION. — Çette c.spèce est prédominante

parmi la faunule étudiée de Los Pastores ; elle est fré¬

quence dans les cordillères Ibérique ci Bétique, parti-

culièrcnieni abondante dan^ le Ladinien

supérieur-Carnien du Prébétique de Murcie, selon

Marquez-Aliaga & Martine? (1996 : 108, 109).

Descrifuon

La forme rrigone est d’une hauteur moyenne de

15 mm, d’une longuear moyenne de 16,5 pour

un nombre de cinq spécimens, ayant les dimen¬

sions suivantes :H-12;l4;15;17et

L=+10:16;17; 20, La longueur peut être

légèrement inléricure, égale ou un peu supérieure

à la hauteur ; la movetinc de l'indice des para¬

mètres L/H esr proche de 114,8 %. Le galbe esc

peu convexe, les' crochets sont situés aux deux

cinquièmes environ antérieurs de la longueur ,

Fangle umbonal oscille entre 85" et 100“. l.^exrré-

mité anréricurc est o\'alaire en continuité avec le

bord ventral, tandis que le bord postérieur est

légèrement convexe. La carène marginale est

assez saillante (Fig. IF) précédée d'ulie dépres¬

sion antécarénale ; Paréa est étroite, subdivisée

par un sillon médian ; l’écus.son est inobservable.

Le flanc porte trente a irenie-cinq côtes commar-

ginales, régulières, denses dont les intervalles

s'élargissent avec la croissance. Sur des hauteurs

successives par intervalles de 3 mm, i partir du

stade observable le plus juvénile, le nombre de

costules concentriques descend de douze à neuf,

à six, puis i cinq tout en s'épaississant et s'incur¬

vant au niveau du sillon ancccarénal et sur la

carène marginale, avant de sc poursuivre sur

l’aréa. Cette co.stulation est plus dense, plus fine

que celle connue par les illustrations des synty'pes

de Schmidt ou de celles d'autres auteurs (Virgili

1958 ; Marquez-Aliaga bi Montoya 1991), soit

dans le Muschelkalk supérieur de la cordillère

côtière catalane, soit dans le Ladinien supérieur-

Carnien des cordillères béfiques (zone.s externes :

Prébétique et Subbérique).

Les différences entre les spécimens de Los

Pastores et ceux d’autres localités, relatives à la

taille plus petite, Findice de.s dimcn.sions L/H

inférieur, les lignes de costules de croissance plus

fines et régulières, nautori.sciiL pas leur attribu¬

tion à un nouveau .statut subspéciliquc pour ces

variants. Ce sont des morphoty'pes ou morphes

(Simpson 1961 : 178), soir des formes parmi un

extrait de population fossile de faible effectif,

donc de rang infrasubspécifique.

Genre Neoschizodus Giebel, 1855

Sous-genre Neoschizodus s.s.

Neoschizodus (Neoschizodus) laevigatus

(Coldfass, 1837)

(Fig. IH)

l.yrodon laevigaium Goldfuss, 1837 : 197, pi. 135,

fig. 12 a, b.

Myophorid faevi^ata Alberti - Defretin, Durand-Delga

&Lambert i943 : 191, pl. 1, figs 10-12.

Myophoria laevigata Ziethen - V^irgili 1958 : 476,

pl. 1L fig. 9.

Neoschizodus Lievigintis (Goldfu.ss) - tmx 1969 ; 475,

figs D 62, ? . 1 . b. — Newell 6c Boyd 1975 : 74, pl. 12,

figs C. D 141, figs 82 A'D. - Tamura 1981 : 12,

figs 12-18. - Marquez-Aliaga & Montoya 1991 : 121,

pl 1, fig. 1.

142

GEODIVERSITAS • 1999 • 21 (2)

Bivalves triasiques d’Andalousie

Neoschizodus luevigalus (Albcrti) - Farsan 1972 : 178,

pl. 45, fig. 7-3.

Neoschizodus {Neoschizodus) laevigatus (Goldfuss)

- Fleming 1987 ; 18, fig. 7 a, b.

MaiëRIEL. — Deux vaiveü droireü, une valve gauche

(n** LPM-R. 62097)) mctiiles internes de lormc civale-

arrondie de diamètres voisins de 22 mm, à carene pos¬

térieure peu m.uquée et d'aspect lisse, n’ayant

consersT que quelques lignes de croissance commargi-

nalcs.

Distribution. Furasiarique : Puntsandstein supé¬

rieur a Leirenkohle d'Allemagne : VC'frfénien a

Ladinien des Alpes du Sud ; Anisien-Ladinien de

Transjordanie et d Israël : Keuper d’Algérie ; Anisicn-

Norien d’Asie, lin Fspagne, de nombreux gisements

de l’Anisien, Ladinien int»ycn ei supérieur des c<-»r-

dillèrts in Marque/.-AÜaga & Mariinci; 1996 : 105,

Ibérique, 107, Pyrénées catalanes ; 109, Bétique orien¬

tale (Prébérique et Subbérique)...

Sous-classe HETERODONTA Ncuniayr, 1884

Ordre VENEROIL9A

FI. Adams & A. Atlams, 1856

Superfamille CkASSATLI-IAc.I.A l'érussac, 1822

Famille MVOrHOKIFAKDIlOAT

Chavan bi Cox \in Moore cd.], 1969

Genre Pseudocorbida Philippi, 1898

Psettdocorbiila gnegaria

(Münsrer in Goldfuss, 1837)

(Fig. Il,J)

Nucula gregariit Munster in Goldfuss, 1837 : 152,

pl. ! 24, Rg. 12 a, b.

Myophoriopsis gjr^tuia (Münster) - Schmidt 1935 :

84. pl. 5, (ig. 36. '

Psetidocorbulagregaria (Münster) - Farsan 1975 : 132,

pl. 2. figs 9-12.

Pseudocorhiilo gregario (Mün.sier in Goldfuss) — Cox

& Chavan 1969 : N 582, fig. 81, fig. 3a-

d. - Mdrquc/.-Aliaga, Hirsch & L6pc/-Garrido 1986 :

18, fig. 4C (avec synonymie).

MAI TKUa.. — I )eux valves gauches de hauleiir dc 7,5

et 9 mm pour une longueur de 12 er 10 mm (n“ LPM-

R. 62098).

Distribution. — Pseudocorbuln gregtiria est à large

répariiuon eurasiarique : du Mirschelkalk moyen et

supérieur à la Lettenkohlc d’Allemagne, Anisien des

Alpes du Sud, l.adinicn d’Afghanistan central. En

Espagne : Ladinien-Carnien des cordillères Ibériques

(terme carbonate supérieur du Muschelkalk (provinces

de Tcruel, Cucnca, Valence) ; I.adinien .supérieur de

la branche Aragonèse ; Anisien-Ladinien des

Catalanidc.s ; I.adinien de la cordillère Bétique

(Subbétique dc Jaén, de C^adi/.-Algésiras-Boyar) ;

Ladinien-Carnien des Baléares.

Ri:marqui-;vS

Leur forme est suberigone, à carène postérieure

peu élevée, délimitant une aire postérieure étroite

limitée par un bord postérieur rectiligne obliqtie,

puis droit à son extrémité distale. C'es individus

se .superposent, d une pan a la figure-texte don¬

née par Virgili (1958, fig. 59-5) pour l’espèce

gregaria et d autre part à celle de Myophoriopsh

(Pscndocorbula) keuperina (Quenstede 1851 î Vir¬

gili [958, fig. 59-2), espece mise en synonymie

avec la précédente par Marquez-Aliaga et ai

(1986).

SIGNIFICATION DE LA FAUNE DE

BIVALVES ET CONCLUSIONS

A partir de l’assemblage de bi\'al\e.s étudié, pro¬

venant du gisement meme de Los Pastores dc la

série Fariquidc, il est possible d'envisager sa

signification d’ordres biosrrarigraphit]ue, paléo-

biogeographique, palcoécologique.

1. La disiribulion de ces espèces citées dans le.s

divers domaines germanique, alpin-tcthy.sien et,

plus particulièrement, dans celui dc l’Espagne est

portée sur la Tableau 1. Leur association suggère

d’assigner un age Ladinien supéricur-Carnien

inférieur à cette fiitme du rocher de Los Pastores,

sans exclure toutefois le Carnien .seul d’après les

corrélation.^ biogéographit|ues avec le Trias

d’Espagne.

2. Les analyses, entre autres, de Hirsch (1977)

dans le domaine sépharade permettenr de relever,

en eflei, certaine.^ affiniiés avec : (a) le Trias cata¬

lan, en patuculicr le Carnien inférieur à

Costaioria goldfussiP le Langobardien avec

Lyriomyphoria suhlaevis > (b) la faune de Royuela,

région de Tcruel des chaînc.s ibériques, conipte

tenu des especes signalées, Ctervillia joleaadi^

Leptochondria albenP Costatoria goldfttssii,

lyrunnyphoria snhlaevis. Neoschizodus laevigatus^

dont Fage sc .siiLierait a la limite du Ladinien

supérieur et du Carnien inférieur ; (c) la meme

association spécifique précédente des cordillères

GEODIVERSITAS • 1999 • 21 (2)

143

Freneix S.

Tableau 1. — Distribution straligraphique des espèces de bivalves citées du Trias germanique, du Trias alpin-téthysien. du Trias

d'Espagne. Dessin Françoise Pilard (LPM. MNHN).

ESPÈCES

Muschelkalk

Keuper inf.

Anisien

Ladinien inf.

Ladinien sup.

Carnien

Gervillia joleaudi

— - -

Leptochondria alberti

m m

- - -

—

— — _

Posidonia obliqua

Cosîatoria goldfussii

- - -

—

Lr_:z

Lyriomyophoria sublaevis

Neoschizodus laevigatus

- - -

— - _

—

Pseudocorbula gregaria

—

- - -

Trias germanique : — - Trias Alpin-Téthys : — Trias d’Espagne :

bétiques, zone interne, prébetique de Jaén, récol¬

tée dans une assise carbonatée du Ladinien supé¬

rieur ; (d) le membre somniital de la tormation

de Fuentc Aledo (Murcie) des cordillères

béciques, zone imerne, livrant Costatorui goldftis-

sîi et Gervillia cb jnleuudiy qui serait datée du

Ladinien supérieur ; (e) toutefois, malgré les affi¬

nités relevées avec ces divers domaines paléobio-

géographiques, l’a-ssemblage de bivalves étudié

présente une particularité : la prédominance sur

les autres especes de Lyriomyophoria îubhxevh

définissant un biofacics à Lyriamyophorin du

groupe /-. degiwSy signalé dans plusieurs horizons

triasiques de différeiucs cordillères (Martin-

AJgarra et ai 1993 ; Marqucz-Aliaga craL 1996).

3. Les caractères taphom>mû]ue.s de l’assemblage

indiques préccdemmeiu dont, en particulier, la

désarticulation des valves stabilisées, leur surface

externe convexe tournée vers le haut de la

couche, laissent supposer un certain hydrodyna¬

misme du milieu. Comme aucune direction pré¬

férentielle d’orientation des valves n est obser-

v'able, on peut en déduire qu'il s'agit d’une sim-

migie allochrone. La morphologie fonctionnelle

des espèces, la sédimentologie originelle de leur

milieu de vie supposent des implications. Les

modes de vie tous filtreurs saspensivores se repar-

ti.ssent en deux groupes : épifaunique et infau¬

nique.

L'épifaunc comprend les especes : ? Gervillia

joleandiy !Aptoebondria nlhcrti^ ? Posidorna ohti~

cjun. Lc.s trois genres représentés [Gervillia,

Leptoebondria, Posidonia) sont cpibcnthiqiics,

pscLidoplanctoniqucs. oscillant.s ou pendants,

fixés par des fils de byssus h des algues ou autres

supports flottants ou fixés (Kauflmanii 1969).

Les juvéniles devaient être facilement déraches tic

leur support par l’agitation de.s eaux.

L'endütaune est composée des quatre espèces

libres, louisseuses dans des sédiments meubles :

Costdtorin goldfusiii, Lyriomyophoria sublaevis,

Neosebizodus laevigatus, Pseucorbula gregaria. Les

144

GEODIVERSITAS • 1999 • 21 (2)

Bivalves triasiques d’Andalousie

Myophoriidiie auxquelles s'ajoute le crassarellacê

Peudocorhula .souvetit opportuniste (Marquez-

Aliaga & Garcia-Gil 1991)» sont des fouisseurs

superficiels actifs, en position de vie à la limite

sédiments-eau que leurs zones exhalante cr inha¬

lante dépassent parfois.

En tant qu’ancêtres supposés des Trigoniidae, ils

avaient la capacité, comme Lyriornyophoria

(Stanley 1977 : 896). de s’adapter aux sédiments

instables et aux variations de salinité.

4. Conclusions : la launc de bivalves étudiée,

bien que de faible diversité, a permis de la situer,

au Trias, dans le Carnien du domaine sepharade.

Elle SC caractérise par un biofaciès a Lyrio-

myophoria sublaevis. Son gisement de Los

Pastorcs évoque un paléoenvironnement marin

d'ctagcmenc infra-littoral peu profond, en bordu¬

re d’un escran gréso-carbonatc, pem-ctre peri-

delraïque, car soumis à des fluctuations

d’hydrodynaml-sme (courants, \^gucs, marées),

de salinité, de température, sous climat tropical.

Remerciements

Je remercie vivement M. le Professeur

M. Durand-Delga pour m’avoir confié Tétude de

cet intéressant matériel, ]:iOLir avoir participé à la

rédaction du texte .stradgraphique introductif et

déposé la dalle lossililerc (n'’ ET. 674), enregis¬

trée SOU.S le rT 1993-1, dans les collections du

Laboratoire de Paléontologie du Muséum natio¬

nal d'Histoire naturelle de Paris (LPM,

MNHN). J'exprime ma gratitude, pour leurs

judicieuses critiques, aux rapporteurs ;

Mlle Annie Dhondt, M. Philippe Bouchet.

M. Manuel Esteras, Mme Anne-Marie Ohler.

Mes remerciements s'adressent aussi à M. Denis

Serrette pour les très bonnes photographies, à

Mme Françoise Pilard dessinatrice, pour les illus¬

trations, à Mme Danielle Quinrus pour la saisie

de texte, sans oublier M. Hervé Barrât pour

Taide apportée âu dégagement très difficile du

contour des fossiles.

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146

GEODIVERSITAS • 1999 • 21 (2)

Un paléoniscoïde (P/sces, Actinopterygii)

de Buxières-les-Mines, témoin des affinités

fauniques entre Massif central et Bohême

au passage Carbonifère-Permien

C. POPLIN

Laboratoire de Paléontologie, UMR 8569 du CNRS, Muséum national d’Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

cpoplin@mnhn.fr

Poplin C. 1999. — Un paléoniscoïde {Pisces, Actinopterygii) de Buxières-Ies-Mines, témoin

des affinités fauniques entre Massif central et Bohême au passage Carbonifère-Permien.

Geodiversitas 2^ (2) : 147-155.

RÉSUMÉ

Description d'un paléoniscoïde, Progyrolepis htyleri n. sp.» ti’uprcs neuf maxil¬

laires ec mandibules isoles sur des plaques de .schiste provenant du Permien

inférieur (Aucunien) du bassin de l'Aumance (Massif central). Os pièces, de

six à neuf centimètres de long, portent deux rangées de dents marginales

(petites externes, grandes internes) coniques à chapeau d’acrodine. La plaque

postorbitaire du maxillaire est assez longue, basse et trapézoïdale. L’attribu¬

tion générique de ce poisson se fonde sur la similitude de son ornementation,

très caractéristique, avec celle de P. spcàosus (Carbonifère supérieur de

Bohème). Ceci est une nouvelle indication de ce qu’il n'cxistair pas de barriè¬

re biogéographique entre les bassins Ümniques du Vfassif central et de

Bohême à la fin du Carbonifère et au début du Permien.

ABSTRACT

A paleoniscoid fPisces, Actinopterygiij from Buxières-Ies-Mines, evidence of

faunal relationships between Massif central and Bohemia at the turn over bet-

ween Carhonferous and Pennian.

Description of a paleoniscoid, Pro^rolepis heyleri n. sp., based on nine isolat-

ed niaxillaries and mandibles on shale slabs from the Lower Permian

(Autunian) of the Aiimance basin (Massif central). These specimens, six to

nine centirneters long, bave two row.s of marginal teeth (extcrnal small, inter¬

nai large)» conical with an acrodin cap. The postorbital plate of the maxillary

is rather long, low and trapézoïdal. The generic attribution of rhis fish is

based on the similitude ol its very charactcrisiic ornamentation with that of

P. speciosus (Upper Carboniferous, Bohemia). Fhi.s is a new indication that

rhere was no biogcographicaJ barrier between the basins of the limnic faunas

Massifeentrï Massif central and of Bohemia during the Upper Carboniferous and

Bohemia. Lower Permian.

KEYWORDS

Actinopterygii.

Paleoniscoid,

Autunian»

MOTS CLÉS

Actinopter)'eii»

paléoniscoïde,

Autunicn,

France,

Massif central,

Bohême.

GEODIVERSITAS • 1999 • 21 (2)

147

Poplin C.

INTRODUCTION

Les iictinopccrygicns du Permien inférieur du

bassin houiller de Buxières-les-Mines sont repré¬

sentés le plus souvent par des os isolés. C’esr

pourquoi aucune détermination précise n’a pu

erre donnée jusqu à présent dans les publications

consacrées a la paléoichthyologie de ce bassin

(Heyler 1984 ; Heyler ik Pt^plin 1990) Le maté¬

riel décrit ici est également très partiel car il

consiste en maxillaires et en mandibules isolés.

Mais leurs caractéristiques permettent de mieux

cerner leur appartenance taxinomique : ü s’agit

d’une espèce nouvelle attribuée au genre

Prog)>rolepis. Lincérct de cette étude ne se limite

pas à la systématique, die donne aussi quelques

indications sur les relations paléobiogéo¬

graphiques entre ce bassin et la Bohême aux

confins du Carbonilcre et du Permien.

Provenances GEtxutAPHiQüE et

STRATIGRAPIIIQUE

Les spécimens ont été récoltés dans la dernière

mine de Buxiéres (» Découverte n° III ») encore

exploitée par les Houillères des bassins du Centre

et du Midi dans le bassin de l’Aumance, qui fait

partie des bassins carbonifères et permiens de

l’AlIier (Steyer & Escuillié 1997). Us viennent de

la Formation de Buxières datée du Permien infé¬

rieur et attribuée h l’Ancunien en tant qu’unicé

lithostratigraphique (Steyer et al. en préparation).

Historique

l.es premières fouilles datent des années 1960-

1970 par D. Heyler et par des géologues dont

P. Debriettc qui a découvert le maxillaire choisi

comme holotypc du nouveau taxon décrit ici.

Heyler (1984 : 114) a mentionné cette pièce sans

la décrire Heyler ik Poplin (1990) ont signalé la

présence de pièces i.solées évoquant Progyrolepis

speciosus Fritsch, 187*^ de Bohème. Depuis 1996,

les activirés sur le terrain ont repris grâce à la

détermination des membres de Passociation

« Rhinopolis *> : ce sont eux, dont J.-M. Pouillon,

qui ont mis au jour les autres spécimens étudiés

ici.

Matériel ee fossilisation

Il s’agit de trois maxillaires et six mandibules iso¬

lés sur des plaques de schiste ci visible.s par leur

face latérale ou par leur face mésiale. L'absence de

traces de morsures écarte faction de prédateurs et

favorise pltitôr fidée d’une mort naturelle suivie

d’une décomposition sur place. N’étanr pas uses,

ces éléments ifiont du subir t|u'un transport

faible après leur désaruculadon. Cette h} 7 )Othcse

est soutenue par le fait qu’un ma.yillaii'e droit et

deux mandibules droite et gauche gisent à proxi¬

mité les uns des autres sur une même plaque de

schiste, suggérant qu’ils peuvent provenir du

même individu.

SYSTÉMATIQUE PALÉONTOLOGIQUE

Classe OSTEICH I HYES Huxley, 1880

Sous-Classe ACflNOPTERYGII

Woodward, 1891

Ordre PALAEONISGIFORMES Hay, 1929

Famille PYriOPTr-RinAi-; Aldinger, 1937

Genre Progyrolepis l'ntsch, 1895

Progyrolepis heyleri n. sp.

Holot\'PF. — n” BLIX 86, déposé dans les collections

du Laboratoire de Paléontologie du Muséum national

d Histoire naiiirdlc (Paris). Un maxillaire droit en vue

latérale sans contrepartie. Figuré in Heyler 1997,

% 9 -

LOCALÜ Ê-TVPE. —- Buxières-les-Mines (Allier,

France), Découverte III.

AC!K. — Permien inférieur,

ÉTi'MOl.OcaF. — Espèce dédiée à Daniel Heyler, pre¬

mier paléontologue à avoir étudié l’ichthyofaune de

Buxières-les-Mincs.

Auire MATlIniEE. C^ollecrlon personnelle de J.-M

Pouillon : n" J MP 177, empreinte de la face latérale

d'un maxillaire gauche et vue mcsialc du bord den¬

taire sans contrepartie (figuré in Poplin 195l'7b,

llg. 2} ; 11 “ jMP 194 A et B, extrémité anrérieurc de

mandibule gauche, partie et coturepartie. —

Collection de ! association « Rhinopolis » à Buxières-

les-Mines : n'^ BX 28089 R, grande pliuiue de schiste

(60 cm X 46 mi) sons œntrefxiriie, avec un maxillaire

droit en vue mésiale, une mandibule droite en vue

larérale (figurée in Poplin 1997b. fig. 1 ) et une mandi¬

bule gauche en vue mésiale ; BX O.V)895. une mandi¬

bule gauche en vue latcTde sans contrepartie ; BX M

093 1/2 et 2/2, mandibule partie et contrepartie ; BX

148

GEODIVERSITAS • 1999 • 21 (2)

Paléoniscoïde autunien de Buxières

F»g, 1. — Progyrolepis heyleri n. sp.. Autunien de

Buxières-les-Mines {Massif centrai, France) ; holo-

lype, colL MNHN, BUX 86 : maxillaire droit, face

iatérale. Échelle : 1 cm.

260996 (6), fragment de mandibule gauche en vue

latérale.

11 s'agit de collections privées.

DiAciNOSK. — Pro^‘n/lepis dont les maxillaires et les

mandibules ont 6 à 9 cm de long, ce qui suppose une

longueur totale de ranimai de 60 il 70 cm. Maxillaire :

laque postorbiiairc trapézoïdale, as.sez longue er

asse, plus haute posrérieuiemeni, avec une zone lisse

le long de son bord antérieur et un angle posréro*

inférieur marqué mai.s faible ; orneineiuation de fines

rides et vermiculaiiuns eu le long du bord dentaire, de

tubercules ; bord dentaire se relevaiu en courbe vers

l'avant et formani un bmtrreiet visible sur la face laré-

ralc sur rutile sa longueur. Mandibule : allongée, ma.s-

.sive, relevée vers l avant, .sans processus coronoïde,

ornée de rides irrégulières se résolvant en verrnicula-

tions le long des bords supérieur et inférieur. Dents

ntarginiilc.s cûntques pourvues d'un chapeau d'acro-

dine, à côtes vcrucale.s larges, dispo.sée>s en une rangée

latérale de dents nombreuses et petites et une rangée

médiale de dents moins nombreuses, haïUes (6 à

“ mm) à grande cavité pulpaire.

DF..SCRimON

Bien que les pièces soient isolées, la concordance

(le leurs caractères morphologiques, de leur raille

et de la nature de leur ornementation indique

sans conteste quelles appartiennent au meme

taxon.

Maxillaires {¥\gs 1-3)

I.a longueur des spécimens varie de 6,5 à 7,5 cm

environ (ce qui résulte d’une variabilité indivi¬

duelle et du fait que les extrémités des os ne sont

pas toujours intactes). Ils comportent» comme

chez les paléoniscoïdes» une partie sousorbiraire

basse et une plaque postorbitaire haute. Celle-ci

CSC trapézoïdale avec son bord supérieur plus

court que rintcricur et s'en écartant légèrement

vers Farrièrc de telle manière quelle atteint sa

hauteur maximale au niveau de son quart posté¬

rieur (18 à 22,8 mm) ; son bord postérieur des¬

cend obliquement jusqu’à l'angle postéro-

inférieur qui saille légèrement ver.s le bas. Le bord

demaire, Icgcrcmcnt sinusoïdal, se relève vers son

extrémité antérieure. Les dents sont portées par

l’habituel bourrelet longitudinal de la face interne,

mais qui apparaît aussi sur la face externe en for-

manr le long du bord inférieur un petit surplomb

masquant un peu la base des dents. Le rapport de

la longueur de la plaque postorbicaire sur sa hau¬

teur est 2,1 en moyenne. Le rapport des lon¬

gueurs de la plaque postorbitairc et du processus

sousorbiraire est 2 en moyenne.

L'oincmcntacion est complexe et originale. Sur la

plaque postorbiraircv elle est constituée de nom¬

breuses et fines rides irrégulières qui, grosso woekh

sont parallèles aux bords supérieur et postérieur

de l’os et forment parfois des boucles et de petits

tourbillons (Fig. 3A). Vers le quart antérieur de

la plaque, ces rides s’infléchissent vers le bas en

dessinant une ligne verticale nette (Fig. 3B). En

avant de celle-ci, quelques rides s’incurvent vers

le bas et l’avant et se résolvent en petits tour¬

billons ; le reste de cet espace est lisse jusqu’au

bord antérieur (Fig. 3B). Sur le bourrelet dentaire

rornementalion, longitudinale à l’arrière, se frag¬

mente en tourbillons puis en fines vcrmiciilations

à Favant (Fig. 3C). A proximité des dents, ce

sont de petits tubercules arroudi.s (Figs 3F. E)

ponant des stries rayonnantes à partir de l’apex,

comme chez R spedosus (Fritsch 1895, pl. 131,

fig, 15). À fort grossissement les rides révèlent un

relief « en duvet » comme chez Aloythomasia niti-

da Cross, 1953 (jessen 1968, fig. 3), avec une

fine crête médiane de laquelle partent de courues

crêtes latérales et régulières (Fig. 3E).

GEODIVERSITAS • 1999 • 21 (2)

149

Poplin C.

Fig. 2. — Progyrolepis heyleri n. sp.,

Autunien de Buxières-les-Mines (Massif cen-

tral^ France) ; coll. Rhinopolis BX 28089 ;

maxillaire droit, lace mèsiale. Échelle . 1 cm.

Cette ornementation varie d’un spécimen à

l’autre dans le détail à l’instar des dermato-

glyphes. La zone lisse le long du bord antérieur

de la plaque postorbitaire est rare chez les paléo-

niscoïdes : on peut penser quelle était recouverte

d'os sous- ou inlraorbituires du vivant de Tanimal.

Mandibules (Figs 4-6)

Les mandibules font 6 à 9 cm de long, donc un

peu plus que les maxillaires : cette difïerence est

courante cher les acrinoptér}^giens dont le maxil¬

laire est précédé par les os du museau. Files sont

massives, allongées et sans processus coronoïde.

Deux de CCS pièces ont le bord dentaire courbé

vers le haut et I avant comme sur les maxillaires.

Chez trois autres le bord dentaire semble recti¬

ligne : s’agit-il d une déformation post-mortem ?

La face latérale est consiituéc du dcntalosplcnial

suivi de rangulairc dont la suture est observée sur

Tune des pièces. Sur la face mésialc le dcntalo-

spléuial apparaît vcnrralcmcnt avec, au dessus,

Los de MeckcL Le long du bord dentaire, une

série de coronoïdes est suivie d^un large préatti-

culaire, mais les sutures ne sont pas nertes. Les

deux facettes articulaires, à l’extrémité postérieu¬

re de l’os de Meckel, .sont très apparentes sur les

faces latérale et mcsiale.

La fine ornementation consiste en rides longitu¬

dinales et en vermiculutions le long des bords

supérieur et inférieur.

Dents (Figs 1,2, 4-6)

l.es dents marginales .sont coniques, portent un

chapeau d'acrodinc ci sont lisses avec quelques

côtes verticales larges. Elles sont disposées en

deux rangées longitudinales : rangée externe de

nombreuses petites dents (hauteur moyenne

1,9 mm au maxillaire et 1,6 mm h la mandibule),

et rangée interne d’une dizaine de grandes dents

(hauteur rnoyenue 7 mm au maxillaire et 6 mm à

la mandibule) dont la base est un peu clargic par

une vaste cavité pulpaire, visible sur plusieurs

dents cassées, Sur le maxillaire n” Rhino

Bx.28089B (Fig. 2) les alvéoles des dents, internes

sont contigus, mais il ny a de dent en place que

dans une alvéole sur deux : cette disposition doit

erre liée au remplacement dentaire car les grandes

dents sont semblablement espacées sur les autres

pièces.

Langulaire et les coronoïdes sont couverts de

dents minuscules (Figs 5, 6).

DrscussioN

La présence du chapeau d’acrodinc est particu¬

lière aux actinopterygiens à l’exception de

Cheirolepis 1835 (Patterson 1982 ;

Arratia àc Clourier 1996). La plaque postorbitai-

re haute et longue du maxillaire ne s'observe que

chez les Actinopveri basaux (Gardlncr &

Schaeffer 1989) nommés de manière inhirmelle

« paléoniscoïdes » ou « actinopterygiens primitifs

fossiles ». En effet, ccitc plaque fait partie d’un

ensemble de traits morpho-fonctionnels liés au

suspensorium des mâchoires : bouche et joue

longues, hyomandibulaire et préopcrcule très

incline“s. Il en csrde même pour labscnccde pro¬

cessus coronoïde à la mandibule ; celui-ci, nés

rarement observé chez les palconisco'ides (Poplin

&: Véran 1996), est caracicristiquc des néoptéry-

giens (Gardiner 1984). Enfin la dUposicinn des

dents marginales, grandes médiales et petites

latérales, semble être primitive chez les actinopté-

150

GEODIVERSITAS • 1999 • 21 (2)

Paléoniscoïde autunien de Buxières

F»g. 3. — Progytolepis heylehr\. sp, Autunien de Buxières*les-Mines (Massif central. France) : A, B. colL Pouillon JMP 177. maxil¬

laire gauche, détails de l'empreinte de rornementation sur ta plaque postorbitaire ; A. centre de la plaque , B. ligne verticale anté¬

rieure ; C-F. holûtype, coll. MNHN, BUX 86, maxillaire droit, détails de l'ornementation ; C. partie postérieure du bourrelet dentaire ;

D. tubercules ; E. rides et vermiculations , F. partie antérieure du bourrelet dentaire. Échelles ; A-C, F. 2 mm , D, E, 1 mm

lygicns {Poplin &: IMcylcr P)93). Il ré.siilrc de ce

qui précède que cecce espèce de Buxières esr un

actinopeciygien paléoniscoïde.

Mais scs affinités au sein de ce groupe sont plus

difficiles à établir, force est de porter attention à

des caractères souvent traités comme mineurs :

détails de la süliouctte du maxillaire et de son

ornementation, hauteur comparée et état de sur¬

face des dents. Cette reciierclie n’est guère facile

car les de.scriptions des paléoniscoïdes ne sont pas

toujours poussées à ce point de détails. Mais par

chance fun de ceux-ci, rornementation, est suffi-

GEODIVERSITAS • 1999 • 21 (2)

151

Poplin C.

Fig. 4. — Progyrolepis heyleri n. sp.. Autunien de

Buxières-les-Mines (Massit central, France) ; coll.

Rhinopolis BX 28089 ; mandibule droite, face latéra¬

le Ang, angulaire ; Dsp, dentalosplènial ; f.a.l.. fos¬

sette articulaire latérale Échelle i cm,

samment caraccéristique et précis pour permettre

Tattriburion au genre Progyrolepis. l.a discussion

ci-après est limitée aux trois genres directement

concernés par ccue nouvelle espèce.

1. Heyler (1977i 1997) a étudié une mandibule

isolée du Permien de Lodève dont îl a fait Tbolo-

type du genre et de l'espèce Uscldsichtbys rtulvro-

dem Heyler, 1977. Ce taxtîu est remarquable par

la taille et là morphologie des dents marginales

internes. Ixur hauteur est plus grande que celle

de la mandibule à leur niveau . ceçi résulte du

fait que leur nioitié distale, qui a la forme

conique habituelle avec le chapeau d'acrodine,

esc portée par une base .aussi liante et considéra¬

blement élargie en forme de coupole auiour

d'une cavité pulpaire très vaste. En outre, le bord

dentaire porte de petites dents insérées en bou¬

quet sur des capsules hémisphériques.

Heyler (1997) a fait une première description du

maxillaire défini ici comme riiolotyjoe de

Progyrolepis heyleri. Remarquant sa caille, voisine

de celle de l’espèce de Lodève, ainsi que la hau¬

teur et la forme de ses grandes dents, il a émis

l’hypothèse que ce maxillaire pouvait appartenir

à une forme proche, voire a IJsclastchthys même.

Mais la base dentaire plus petite et l'absence de

petites dents disposées en bouquets lui ont Hit

évoquer également la possibilité qu'il s agisse

d’un « actinopréiy^gicn classique ». Cette dernière

proposition est confirmée par l’observation précise

des autrc.s pièces (Poplin 1997b) : bien qu'clargie,

la cavité pulpaire ne détermine pas de forme en

coupole comme chez ihclasichthys Heyler, 1977

si bien que les dents sont moins hautes que la

mandibule à leur niveau.

2. Nombre paléoniscoïdes ressemblent à l’espèce

Pragyroltpis heyleri par le.s grands traits de leurs

mâchoires : plaque posrorbitaire du maxillaire

trapézoïdale et plus haute posterieurement,

bouche incuivéc vers le haut et l'avant, grandes

dents marginales ayant une vaste cavité pulpaire,

ornementaiion de rides de ganoïnc. Parmi ces

paléoniscoïdes, deux genres se disiingueiu par

plusieurs caractères mineurs qu'ils partagent avec

l'espèce de Buxières : Nemutopiychius et

Progyrolepis.

L’espèce type de Nanahptyduus TTiie\\xm\ 1875,

N. d'raquair, 1S75, provient d’un milieu

estuarien du Carbonifère inférieur d'Ecosse. Ce

poisson, long d'une quarantaine de centimètres,

présente avec celui de Buxières les points com-

mun.s suivants : le rccouvicmenr de lu plaque

postérieure du maxillaire parles os postorbitaires

et la .surface des dents lisse à part quelques côtes

larges sur la face linguale. Mais ellcs'cn distingue

par l'angle posicro-inlcricur du maxillaire très

fort, la silhouette plus gracile de la mandibule et

rornementaiion du maxillaire faite de simples

rides régulièrc.s et de tubercules le long du bord

dentaire ( Jraquair 1877, pl. L figs 9, H ; pl. 26,

fig-s 1,5-7).

Progyrolepis speciasus (Eritsch, 1875), type du

genre Progyrolepis créé par Frirsch (1895 : 118),

vient du Carbonifère supérieur du ba.ssin intra-

montagneux de Kounov (Bohème). Ce poisson

est long de quelques 60 cm scion Scamberg

(1991), comme, probableineiit, l’espèce de

Buxières. il partage avec cette derntère, outre la

forme massive de sa mandibule^ l angle postéro-

inférieur peu marqué du maxillaire et, surtout,

rornemenration des deux mâchoires (rrit.sch

1895 : J19, pl. 131 fig. 12 : Stamberg 1991 : 53,

fig. l4, pl.s 5, 7, 9). En effet P. spedostù a exacte¬

ment la même ornementation, dans son aspect et

152

GEODIVERSITAS • 1999 • 21 (2)

Palconiscoïdc autunien de Buxièrcs

Fig. 5. — Progyrolepis heyleri n. sp., Autunien de

Buxières-les-Mines (Massif central. France) : coH-

Rhinopolis BX 28089 ; mandibule gauche face

mésiale. Cor. coronofdes : Dsp, dentalosplénial ;

t.a.m., fossette articulaire médiale . mk. os de

Meckel : Par, prearticulaire. Échelle : 1cm.

sa disposition, que les spccimens de Buxières. Il

est inhabituel de fonder une attribution géné¬

rique essentiellement sur Taspect des reliefs de

ganoïne des mâchoires ; mais cette ornementa¬

tion est à ce point originale que le lait de la

retrouver idcnritjuc clic/, la foi me de Boliéinc et

celle de Buxière.s me pousse a eu laire une syna-

pomorphie de Progyrolepis. I.e pan âge de ce

caractère, ajouté aux autres cités plus haut,, m’a

décidée à attribuer le poisson de Buxières au

genre Progyrolepis.

La création de l’espèce heyleri se justifie par les

différences qu’elle présente avec l’espèce P. specio-

sus :

- la surface des dents. Chez P spedosus (Fritsch

1895, pl. 1.32, figs 4-6) elle porte de nombrcusc.s

stries verticales entre lesquelles l’email esc couvert

de minuscules tubercules ovoïdes réguliers : ce

sont vraiscinblablemcnc des autapomorphies de

P. spedosus ;

- l’absence sur le maxillaire de P spedosus des

caractères suivants de P htyleri : ligne verticale

d’ornementation de la plaque postorbicaire, et,

en avant de cette ligne, surface lisse de recouvre¬

ment par les os sous- ou infraorbitaircs, enfin

bourrelet longitudinal du bord dentaire apparent

sur la face externe ;

- les proportions un peu différentes de la plaque

püstorbitaire, plus longue chez P. heyleri. Chez

P. spedosus le rapport de la longueur à la hauteur

de la plaque poscorbitairc est de 1,6 en moyenne ;

celui des longueurs de la plaque postorhirairc et

de la partie sousorbiiaire est de 0,9 en moyenne

(Sramherg 1991 ).

Ainsi le genre Progyrolepis comporterait actuelle¬

ment deux espèces ; P. spedosus (Fritsch, 1875) et

P. heyleri (présent travail). Je suis pleinement

d’accord avec Stamberg (1991) sur le fait que

l’espèce P. tricessimaloris Dunkie, 1946 n'appar¬

tient pas à ce genre en raison de leurs différences

ostéologiques, en particulier cclle.s du maxillaire

et du préopcrculc.

Avec un materiel limite aux maxillaires et mandi¬

bules, P he)>leri n’apporte pas d'arguments nou-

veau-K sur les affinités de Progyrolepis, comme sur

la suggestion de Fritsch (1895) selon lequel

Gyrolepis Agassi/., 1833 serait intermédiaire entre

Progyrolepis et Acrolepis Agassiz, 1833. Mais la

comparaison faite plus haut entre P. heyleri et

Nernasopiychins draquair, 1875 est significative :

elle vient h l’appui du groupeinem de ces deux

genres dans la famille des Pygopteridae

(Staniherg 1991).. Le cas de Watsouichehys peai-

nditts (draquaii, 1877) du Carbonifère inferieur

d’Écossc va probablemenr dans le meme sens. En

effet, malgré des différences (plaque postorbitaire

du maxillaire plus longue, mandibule moins

massive, bords dentaires droits), il présente avec

Progyrolepis une forte ressemblance concernant

l’ornementation (Stamberg 1998).

CONCLUSION

Définir aux niveaux du genre et de l’espèce trois

maxillaires et six mandibules isolés d’actinoptéry-

gien paléoniscoïde était une gageure car ces

pièces SC distinguent par des caractères mineurs.

Mais la chance a voulu que l'un de ceux-ci,

l’ornemenlation, ait etc déterminant en étant

retrouvé à Lidentique sur du matériel connu

depuis plus d'un siècle et provenant d’une autre

région et d'un autre étage straiigraphiquc. C est

ainsi que l’on peut résumer comment six pièces

GEODIVERSITAS • 1999 • 21 (2)

153

Poplin C.

Fig. 6. — Progyrolepts heylen n. sp.. Autunien de Buxières-les-

Mines (Massif cenîral, France) ; cûfl- Rhinopolis BX 260996(6) ;

détail de deux grandes dents marginales, vue latérale. Échelle ;

4 mm.

isolées de rAutunien de Buxières sont attribuées

à Progyrolepts heyleri ii. sp., dont le genre a été

décrit pour la première fois dans le Carbonifère

supérieur de Bohême par Fritsch en 1895. De

cette histoire sont tirées une leçon et une conclu¬

sion.

La leçon est que, dans l'étude d’un matériel fos¬

sile, il ne faut négliger aucun spécimen : une

pièce isolée peut donner des informarions utiles.

Il n’est pas mauvais de répéter cerie vérité pre¬

mière à ceux, rares heureuscmcnc, qui ne con.scn-

tent à travailler que sur les spécimens beaux et

complets. Il n’en re.ste pas moins que la mise au

jour à Buxières de tels spécimens en connexion

de Progyrolepts htylert est forcement souhaitée ;

ainsi pourront être précisée son anatomie et

confirmées ses relations phylogéniques avec

l’espèce de Bohême.

La conclusion est paléobiogéographique.

Progyrolepts est une forme d'accinoptétygicn pré¬

sente à Li fois à Buxières et en Bohême au tour¬

nant du Carbonifère et du Permien» comme

Paramhlypicrtts et des aeduclliformes (Sramberg

1985 ; Hcylcr éc Poplin 1990). C'est là un nou¬

veau témoin des fortes affinités fauniques

maintes fois constatées à cette époque entre le.s

bassins Hmniques du Nord du Massif central et

ceux de Bohême (làiplin 1994, 1997a) et une

démonstration de Pabsencc de barrière biogéo¬

graphique entre ces deux tcrriioircs à la limite

Carbonifere/Permien. Cette conclusion sera pro¬

bablement un leitmotiv des travaux en cours sur

les fossiles exhumés par les fouilleurs de l’associa¬

tion Rhinopoli.s à Buxières, en particulier les

acanthodiens, les xénacanthiformes, les autres

acrinopréiygiens et les tétrapodes, sans oublici les

invertébrés dont les insectes.

Remerciements

Je tiens à remercier l’association Rhinopolis et

J.-M. Pouillon pour le prêt des spécimens étudiés

ici, D. Heyler, H. Lelièvre et S. Stamberg pour

de fructueuses discussions, LL I.avina pour la réa¬

lisation de la planche et D. Serrette pour celle des

photographies.

RÉFÉRENCF.S

Anaiia Cî. & Clouticr R. 1996. — Rcassessment of

ihe morphology of Chcirolepis canadensis

(Aciitioprerygii): 165-199. in Schultze H. P. &

(’Uiiuiei R. (ecis), P)eoOHi*in jlsbei and plants of

Migoiiha (fichi'c» Canada- Veviug Dr., [Ticdrich

Pfeil., Müiicheri.

Fritsch A. 1875. — Ôher die Faun.i der Caskohle der

PiLsiier und Kakonii/en. ditzungsberichthe der K.

Piitnisehm (.^seiLvhafî der Wisseuschafien Fd. (.iregr,

l^raha, I I p.

— 189S. ^ Ftutna der Ga^kohle und der Kalbfeine

der Pevmformation Bnhmens. IfL 1. Kd. ('.rêgr,

Pr.ib.t. \Sl p.

(’rardincr R. 1984. — The iclaiiitiishlps «W tlu- palaeo-

niscid tlshes. a review based on new specinieiis of

AIftnta tind A'îüvthornasia froni the* Upper Devonian

of V(''esfern Australia. Bulletin nfthe British Muséum

(Nutural History). 3"^, '1: I7.V428.

Gardiiier B. ik Schaeffci B. (1989). Intcrielation-

ships of lowei actiiiopieiygian fishc.s. Zoological

Journal ajihe / inman Sonet)' 97; 135-187.

Heylci D. 1984. — Faune fossile dti IV-iniicn de

l'Allitt, Rei'Uc Scientifique du Bourbonnais :

103-122.

— 1977, Découvcric.s ichihyologiqiie.s dans le

Permien de l.odève : une nouvelle structure dentai¬

re. Géologie Méditerranéenne, IV. 3 : 189-204.

— 1997. Les veitchrc.s permiens du Bassin de

Lodève (Hérault) : Bilan. Bulletin de !a Soaété

d‘!Ustoirc NiîfttîcHe d'Autun 15"* : 5-28.

Moylcr D. & Poplin C. 1990. — Les Vertèbre^’ autu-

niens de* Buxière.s-lc.s-Mînes (Allier, France).

Bulletin du Muséunt national d'Histoire naturelle,

série 4, L 12 (2) : 225-239.

jevsen fL 1968. — Moythomasia nitida Cross und

M. cl. siriaia Cross, E3evonisclic Palaeoiii.sciden aus

dem oberen Plaitenkaik der Bergisch-Cladbach-

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Paléoniscoïde autunien de Buxières

Paffrarhcr Muide (Rheinisches Schiefergebirge).

Palaeontographica 128: 87-114.

Patterson C. 1982. — Morphology and interrelation-

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Poplin C. 1994. — Montceau-les-Mines, bassin intra-

montagneux carbonifère et permien de France :

reconstitution, comparaison avec d’autres bassins

d’Euramérique: 289-328, in Poplin C. Ôd Heyler

D. (eds), Quand le Aîassif central était sotts l'équa¬

teur : un écosystème carbonifère à Monîceau-les-

Mines. Editions du CTHS, Paris.

— 1997a. — Le premier Haplolépiforme (Pisces^

Actinopterygii) découvert en France (Carbonifère

supérieur au bassin de Blanzy-Montceau, Massif

Central). Comptes Rendus de l' Académie des

Sciences-, Paris 324 II a : 59-66.

— 1997b. — Le « Pscud’U.sclasichthys » de Buxières,

in Pouillon |.-M., Steyer J.-S., Debriette P. &

Rhinopolis (eds), Livret-guide excursion « Paléonto-

lo^e des Fades autuniens de Buxières-les-Mines, 9 nov.

1997», Association des géologues du Permien, ! p.

Poplin C. & Heyler D. 1993. — The marginal teerh

oF the primitive Fossil actinopterygians : systematics

and évolution, in Heidtke U. (Compiler), New

research on Permo-Carboniferous faunas, Pollichia-

Buch^ Bad Dürkheim 29: 113-124.

Poplin C. & Véran M. 1996. — A révision of the

actinopterygian fîsh Coccocephalus wildi from the

Upper Carboniferous of Lancashire. Paleontology,

Spécial l*apers, 52: 7-29.

Stamberg S. 1985. — Poissons permocarbonifères de

Tchccosiovaquic. Bulletin de la Société d'Histoire

Naturelle d'Autun 114 : 99-113.

— 1991. —Actinopterygians of the central bohemian

Carboniferous basins. Sbornik Narodniho Muzea v

Praze (Acta Musei Nationalis Prague') B XL VIL 1 -4:

25-103.

— 1998. — Preliminary results of the study of

Permian actinopterygian fishes from Buxières-les-

Mines (Allier, France). Acta Musei Reginae-

hradecensis 26; 173-178.

Steyer J.-S. & Escuillé F. 1997. — Le chantier de

fouilles paléontologiques dans le Permien inférieur

de Buxicres-Ies-Mines (Allier, France) en août

1996 : compte rendu préliminaire et perspectives.

Revue Scientifique du Bourbonnais 95 : 11-18.

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Carboniferous formations. Part L Palaeoniscidac.

Palaeontograpbical Societyj London: 1-183.

Soumis pour publication le 2 juin 1998 ;

accepté le 5 février 1999.

GEODIVERSITAS • 1999 • 21 (2)

155

Restes de Rhabdodon (dinosaure ornithopode)

de Transylvanie donnés par Nopcsa au

Muséum national d’Histoire naturelle de Paris

Xabier PEREDA SUBERBIOLA

Universidad del Pais Vasco/EuskaI Herriko Unibertsitatea,

Facultad de Ciencias, Laboratorio de Pateontologia,

Apartado 644, E-48080 Bilbao (Espana)

gpbmubej(5)ig. ehu.es

Laboratoire de Paléontologie, Muséum national d’Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

Philippe TAQUET

Laboratoire de Paléontologie, Muséum national d'Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

taquet@mnhn.fr

Pereda Suberbiola X. & Taquet P. 1999. — Restes de Rhabdodon (dinosaure ornithopode)

de Transylvanie donnés par Nopcsa au Muséum national d’Histoire naturelle de Paris.

Geodiversitas 2^ (2) : 157-166.

MOTS CLÉS

Dinos.îurc,

ornithopode,

Rhahaoelou,

baron Nopcsa,

Crctacc supcncur,

"Transylvanie.

RÉSUMÉ

Des restes de dinosaure provenant du Crétacé supérieur de T ransylvanie ont

été récemment retrouvés dan.s les collections du Muséum national d’Histoire

naturelle de Paris. 11 s’agit d’un dentaire, d’une scapula Fragmentaire et de

deux vertèbres caudales de rornirbopode Rhnhdodott (Igiianodontia). Certe

collection fossile a été donnée au Muséum par le baron Nopcsa en 1923. Elle

vient s'ajouter aux anciennes collections de reptiles fossiles provenant du bas¬

sin de Hapeg (Maastrichtien) qui sont déposées dans les musées de Londres et

de Budapest.

KEYWORDS

Dinosaur.

orniihopod,

Rhabihuioriy

Baron Nopcsa,

Late Cretaceou.s,

Transylvania.

ABSTRACT

Description oj some Rhabdodon remains (Dinosauria, Ornithopoda) frorn

Transylvania, given hy Nopcsa to the Muséum national d*Hisioire naturelle

(Paris).

Dinosaur remains from the Late Cretaceous of Transylvania bave reccnrly

been found in the collections of the Muséum national d’Histoire naturelle of

Paris. TTif matcrial consisrs of a dentary, a fragmentary scapula and c\vo cau¬

dal vertebrae frorn the orniihopod Rhabdodon (Iguanodoniia). This collec¬

tion is a présent of B.iron Nopcsa to ihc Mu.séum made in 1923. It adds to

the old collections of fossi! reptiles Ironi ihc Hatcg Basin (Maastrichtian)

kept at the muséums of London and Budapest.

GEODIVERSITAS • 1999 • 21 (2)

157

Pereda Subcrbiola X. & Taquet P.

INTRODUCTION

Les premiers restes de dinosaures du Crétacé ter¬

minal du bassin de Haçeg (actuel comté de

Hunedoara en Roumanie) ont été mis au jour en

1895. La découverte et l’étude des vertébrés fos¬

siles de Transylvanie sont étroitement liées à la

vie et a l’œuvre de Fercnc Nopesa (Tasnadi-

Kubac.ska 1945 ; Weishampel Ôc Reif 1984 ;

Pereda Subcrbiola 1996). Le baron Nopesa effec¬

tua de nombreu-ses prospections dans la région

de Hateg et entreprit des fouilles systématiques

dans plusieurs gisements maastrichtiens (notam¬

ment Sânpetrii et Valioara). Il réussir h récolter

une importante collection de reptiles fossiles,

principalement des dinosaures, qu’il décrivit dans

une série d’articles (Nopesa 1900, Î902a, 1902b,

1904, 1914, 1915, 1923a, 1925, 1929). La col¬

lection de dinosaures réunie par Nopesa, qui

inclut des restes d’ornitliopodcs {Tebruitosaurus

Nopesa, 1903 ; Rhtibclodon Matheron. 1869),

d’ankylosaures iStruthiosaurus Biin/el, 1870), de

sauropüdes titanosaures (Magyarûsaums Hiiene,

1932) et de theropodes. est l'une des plus impor¬

tantes du Créracé supérieur d'Europe

(WeishampeD^r rf/. 1991).

Le but de ce travail est de décrire quelques osse¬

ments de dinosaure ornitfiopode provenant des

sédiments finicrétacés de Transylvanie, récem¬

ment découverts dans les collections du Muséum

national d’TTstoire naturelle (MNHN) de Paris.

Ces os font partie de l'assemblage dinosaurien

récolté dans le bassin de Hateg et ont été donnés

par Nopesa au MNHN de Paris. Cette descrip¬

tion est l’occasion de faire un bilan du sort des

anciennes collections de reptiles fossiles de

Transylvanie.

PROVENANCE DU MATÉRIEL

Les os proviennent du Crétacé supérieur du bas¬

sin de Hateg, comme Pindiquc une étiquette

écrite en fran*;ais et conservée dans la meme

boîte que les fossiles, où on peut lire : « Crétacé

supérieur de Hongrie. Don de F. Nopesa '>. Le

cahier d'entrées du Muscutn national d'Histoire

naturelle de Paris permet de savoir que ces pièces

ont été données par Nopesa le 21 janvier 1923

pour échange éventuel avec d’autres spécimens.

Elles ont été numérotées par la suite, en 1944.

Nopesa éraii déjà venu en visite à Paris le 21 jan¬

vier 1904 pour étudier les reptiles fossiles de

France présents dans les collections du Muséum

et le 20 et le 21 mars 1905 pour étudier des osse-

menrs de dinosaures de Madagascar. Lorsqu'il fit

ce don et ces visites au Muséum, le protesseur

Marcelin Boule était titulaire de la chaire de

Paléontologie et directeur du laboratoire.

Il convient de rappeler que la Transylvanie

(Siebenbürgcn) Faisait partie de l'Empire austro-

hongrois jusqu'en 1920, date a laquelle elle fut

intégrée à la Routnanie par le traité de ’lrianon.

Nopc.sa étant d'origine hongroise, la faune de

dino-saure.s du bxs.sin de Hateg e.st décrite dans

ses rr.tvaux contmc provenant du Crétacé supé¬

rieur de Hongrie (Nopesa 1923a). Ce detail a

suscité des quiproquos parmi certains auteurs

modernes en ce qui concerne la géographie des

sites. De meme, les principaux gisements de la

région, à .savoir Sânpetru et Valioara (en langue

roumaine) sont décrits dans les articles de

Nopesa avec les noms hongrois de

Szenrpéterfiilva et Valiora respectivement,

l.es formations Sânpetru et Densus-Ciula, qui

ont livré les restes de vertébrés dans le bassin de

Hateg. sont d'âge maastrichtien (Grigorescu

1983 : Weishampel et ai 1991).

DESCRIPTION

Le matériel comprend quatre 05 , à savoir une

branche mandibulainc droite incomplète, deux

vertèbres caudales et une scapiila gauche frag¬

mentaire (Figs 1-3). Ces restes pourraient appar¬

tenir à un ou â plu.sieufs individus. Les fossiles

sont de couleur marron rougeâtre et conserv^ent

par endroits une gangue argileuse de couleur ver¬

dâtre â grisâtre. L'ensemble est inventorié sous le

numéro de collection MNHN 1944.2,

(MNHN 1944.2.1) (Fig. lA-C)

Il s’agit d'un dentaire droit de petite taille (lon¬

gueur conservée égale à 130 mm). 11 est assez gra¬

cile et élancé. Le spécimen est relativement bien

conservé mais ne porte plus aucune dent en

place. L’extrémité antérieure est peu effilée et

158

GEODIVERSITAS • 1999 • 21 (2)

Rhabdodon de Transylvanie du MNHN

Fig. 1. — Rhabdodon sp., collection Nopcsa : A*C, dentaire droit {MNHN 1944.2.1) ; A, vue médiale ; B, vue dorsale ; C, vue laté¬

rale. Échelle : 2 cm.

s'incurve venrromédialemcnt pour former une

courte symphyse mandibulaire. Les bords dorsal

et ventral du dentaire sont à peu près parallèles.

En vue latérale, le bord antérieur est rugueux et

orné de plusieurs foramens nourriciers. Le pré-

denrairc, absent, devait s'articuler avec le dentaire

de telle sorte que son extrémité postérieure soit

située à proximité du premier alvéole Les parois

latérale et médiale de la rangée alvéolaire, com¬

posée de dix alvéoles, ain.si que les parois inter-

alvéolaires ont été partiellement reconstituées

avec du plâtre peint en noir (une pratique jadis

courante). La rangée alvéolaire est légèrement

concave vers l'extérieur mais presque droite en

vue occlusale. Les premier et dixième alvéoles

sont les plus petits, tandis que les sixième et sep¬

tième sont ceux qui ont la plus grande dimen¬

sion. Le processus coronoïde est brisé et seule la

partie basale est conservée. Ce processus est situé

dans le prolongement de la rangée alvéolaire mais

latéralement par rapport à celle-ci. I.e splénial esc

absent, ce qui permet d'obseivcr venrralemcnt le

canal de Meckcl. C!e dernier forme un sillon

étnait en avant qui devient plus large et profond

vers rarnère. Cette partie de la mandibule esc

aussi partiellement reconstituée. La face latérale

du dentaire est très légèrement convexe vers

rcxtéricur et présente deux petits foramens nour¬

riciers â la hauteur des alvéoles sept et huit. Ces

foramens sont alignés en avant pat rapport à un

plateau latéral peu saillant situé au niveau du

processus coronoïde.

GEODIVERSITAS • 1999 • 21 (2)

159

Pereda Suberbiola X. & Taquet P.

Vertebres caudales (MNHN 1944.2.3 et 4)

(Fig. 2A-C)

Deux vertebres sont connues, dont une prove¬

nant de la partie antérieure et l’autre de la partie

médiane de la queue. La première comporte le

centrum et l'extrémiré proximale des processus

transverscs. Le centrum est platycoele à légère¬

ment amphicoelc, avec une surface articulaire

antérieure circulaire et une postérieure ovoïde. La

largeur du centrum (46 mm) esc plus importante

que la hauteur (42 mm) et que la longueur (envi¬

ron 40 mm). Des facettes articulaires en forme

de demi-lune destinées aux chevrons sont pré¬

sences sur la région ventrale. L'articulation avec

les chevrons est intervertébrale. Les facettes pos¬

térieures sont plus développées que les anté¬

rieures. Elles délimitent ventralemenr un sillon

profond de bible dimension. Les faces latérales

du centrum sont concaves. Les processus trans-

verses sont fragmentaires mais semblent disposés

horizontalentcni.

La vertèbre caudale moyenne est relativement

plus complète mais de plus petite taille. Elle reste

partiellement recouverte par du sédiment. Le

centrum cc une partie de l’arc neural, y compris

les postzygapophyscs, sont conservés. Lépine

neurale est brisée. La longueur du centrum

(38 mm) est plus importante que la largeur

(28 mm) ou que la hauteur (27 mm). Les sur¬

faces articulaires sont circulaires à Icgcrcmcnt

ovales. La surface ventrale présente un sillon lon¬

gitudinal peu marqué, délimité en avant et en

arrière par les facetieîf pour les arcs hémaux. Les

posrzygapophyses sont de petite taille et sîtuée.s

sur le bord postérolatéral de l’épine neurale. Elles

ne dépassent guère le niveau de la face postérieure

du centrum. Les processus transverses sont très

peu développés et situés sur la moitié postérieure

du centrum.

5c.z/>«^(MNHN 1944.2.2) (Fig. 3A-B)

Seul un fragment d’une scapiila gauche est

consers'é- l.e spécimen, long de 1SO mm, compor¬

te la région postéroventraie de l'extrémité proxi¬

male (y compris U cavité glcnoïde et une partie

de la surface pour le coracoïde) et le tiers proxi¬

mal de la lame scapulaire. Los présente une lé¬

gère courbure à convexité latérale adaptée à la

forme arrondie de la cage thoracique. La lame

Fte. 2- — Rhabdocton sp., collection Nopesa ; A. B, vertèbre

caudale antérieure (MNHN 1944 2-3) : A. vue postérieure :

B. vue latérale, C, vertèbre caudale moyenne, vue latérale

(MNHN 1944.2.4). Fchelle: 2 cm.

scapulaire, quoique fragmentaire, est mince

(42 mm de largeur mesurée au niveau de la cas¬

sure) et semble s’élargir distalement. Le bord

160

GEODIVERSITAS • 1999 • 21 (2)

Rhabdodon de Transylvanie du MNHN

Fig. 3. — Rhabdodon sp., collection Nopcsa ; A, B, scapula gauche (MNHN 1944.2.2) ; A. vue latérale : B, vue postérieure. Échelle:

2 cm.

antérodorsal de la lame est droit, tandis que le

bord postéroventral est concave. La surface laté-

nde de Textrémité proximale de la scapula pré¬

sente une forte proéminence sur le bord

postéroventral, en aplomb de la cavité glénoïde.

En vue ventrale, la cavité glénoïde est une

dépression plus longue que large, en lorme de

croissant. La surface pour le coracoïde est large et

rugueuse.

DISCUSSION

La matériel décrit est rapporté aux Ornichopoda

d’après sa morphologie générale. Les restes sont

de dimensions plutôt modestes et appartiennent

à un ornithopode de petite taille. L'arc neural et

le centrum d’une des vertèbres caudales sont

fusionnés : il ne s’agit donc pas d'un individu

juvénile.

GEODIVERSITAS • 1999 • 21 (2)

161

Percda Subcrbiola X. & Taquet P.

Le materiel de Hatcg consens à Paris peut être

rapproche des Iguanodontia de part la combinai¬

son de plusieurs caractères, .1 savoir : la présence

d’un dentaire à bords dorsal et ventral parallèles,

d’un processus coronoïde situe latéralement par

rapport a la rangée dentaire, d’un diastème entre

le prédentaire et le premier alvéole, d’alvéoles dis¬

posés sur le bord médial du dentaire, ainsi que la

présence d’un contrefort saillant sur le bord pos¬

térieur de la cavité glcnoïde sur la scapula

(Sereno 1986 ; Norman & Weisliampel 1990).

Le principal caractère diagnostique de Rhab-

dodon est la morphologie de ses dents et, plus

spécialement, rornementation de l'émail dentaire

(Brinkmann 1988 ; Weishampel et ul. 1991).

Léchantillon étudié ne comporte pas de dents

mais, à defaut. la morphologie du dentaire esi

comparable à celle du matériel de Rbabdodon

provenant de Transylvanie déposé a Londres

(Nopesa 1902a, 1925 ; Brinkmann 1988) et à

Budapest (Nopesa 1915). Le dentaire porte dix

alvéoles et possède un petit diastème entre la sur¬

face pour le predenraire et le premier alvéole, des

caractères que Ton retrouve chez Rbabdodon

(Weishampel et al. 1991). Il esc à noter que,

parmi les Iguanodontia, Rbabdodon présente le

plus petit nombre connu de dents mandibulaires.

De plus, la scapula présente une forte proémi¬

nence en forme de crochet sur le bord postérieur,

comme c’est souvent le cas chez Rbabdodon

(Brinkmann 1988). Le bord antérieur est droit

comme chez certains spécimens de Rhabdoçhm

provenant de Transylvanie (Brinkmann 1988) et

chez Tenontosaurus (Lorster 1990). D'autres sca-

pulae de Rbabdodon peuvent montrer un bord

antérieur concave, un caractère commun chez la

plupart des Iguanodontia (Brinkmann 1988 ;

Pincemaille 1997). Ces arguments nous amènent

à rapporter le matériel étudié à Rluihdodon.

Les ornithopodes du bassin de Hatcg incluent au

moins deux genres, riguanodontien primitif

Rbabdodon et l’hadrosauridé Telmatosaiirns

(Weishampel et al. 1991). Rbabdodon peut être

distingué de Telmatoumms par de nombreuses

caractéristiques (Nopesa 190(1, 1902a, 1904,

1915 , 1925 ; Brinkmann 1988 ; Weishampel et

ai 1991) Le matériel conservé au MNHN de

Paris peut être dilTérencic de l'elmatosaurus par le

fait que la mandibule ne montre qu’une ébauche

de batterie dentaire (bien développée chez tous

les hadrosaures) et que les facettes pour rurtitaila-

tion des chevrons sont fusionnées (diviséc.s chez

les hadrosaures).

Rbabdodon est par ailleurs le dinosaure le plus

abondant du Crétacé supérieur de Transylvanie

et d'Europe en général. Depuis Màtheron, qui

créa le genre en 1869 à partir de matériel trouvé

d,ms le Rogn.acien (Maastrichfien) de Pmvencc,

de nombreux restes provenant du Campanieii et

du Maastrichtien de plusieurs contrées euro¬

péennes lui ont été rapportés (voir Brinkmann

1988 pour un inventaire). Rbabdodon pré.senrc

un forte variation individuelle, dont la significa¬

tion n'est pas encore bien appréhendée (l*ereda

Subcrbiola Sanz l'999). Nopesa (1915) évoqua

un dimorphisme sexuel pour expliquer les diffé¬

rences mtjrphologique.s obscrvée.s sur du materiel

transylvain. Néanmoins, certaines d'entre elles

peuvent être imerprétees comme des différences

Lintogénctiqucfi (Brmkmann 1988) Ce dernier

auteur a proposé de rapporter tout le matériel de

Rbabdodon ï une seule espèce, R. priseiis

Marheron, 1869. Néanmoins, d'autres auteurs

estiment que la diversité .spécifique de

Rbabdodon est plus grande que celle envisagée

auparavant et plaident en faveur de la présence

de plu.sieurs espèces (Buffetaut & Le Loeuff

1991). Le matériel provenant de Transylvanie a

ainsi été rapporté a fcspècc R. robusttn Nopesa,

1900 (Pincemaille 1997 ; Weishampel et ai en

préparation). Léchantillon conservé à Paris

semble trop fragmentaire pour pouvoir te.ster

cette interprétation. Il est donc rapporté à

Rbabdodon sp. indci.

Enfin, la position phylogénétique de Rbabdodon

au sein des ornithopodes est discutée. Il a d’abord

été classé parmi les Camptosauridae (Nopesa

1902a, 1904), puis F.ipproché des

Iguanodontidae (Romer 1956). des Dryosauridae

(Milncr & Norman 1984) ou des HypsÜo-

phodonridae (Norman 1984 ; Brinkmann 1988),

avant d’être considéré comme un Igu.modotitia

primitif (Screno 1986). Nornuin ik Weishampel

(1990), puis Weishampel et ai (199J) le cla.ss’ent

provisoirement comme un Iguanodonria imrrtae

sedis. Norman (1998) a défendu l’hypothèse selon

laquelle Rbabdodon pourrait être un hypsilo-

phodontidé qui montrerait des caractères conver-

162

GEODIVERSITAS • 1999 • 21 (2)

Rbabdodon de Transylvanie du MNHN

gents avec les Iguanodontia» tout comme

Tenontosanrus d’Amérique du Nord et

Muttabiirrasmmta d’Australie. Des analyses cladis-

tiqucs réceutes' confirment l’hypothèse de Sereno

(198fi) et suggèrent que Rbabdodon se situe à la

base des Iguanodontia dans une polyroinie non

résolue avec Imontosaurns, Mutaburrasciurus et les

Euiguanodontia de Coria S'algado (1996)

(PinccmaiUc 1997 ; Weishampel étal. 1998).

LES ANCIENNES COLLECTIONS DE

DINOSAURES DE TRANSYLVANIE

Les anciennes collections de reptiles fossiles pro¬

venant du Crétacé terminal de Transylvanie sont

conservées dans les musées de Londres et de

Budapest. Une grande partie de la collection

réunie par le baron Nopesa et sc-S assistants dans

les gisements du bassin de Hapeg fut vendue à

plusieurs reprise.s au Narural History Muséum de

Londre.s (BMNH). La documentation officielle

des Archives du BMNH atteste de l’achat de rep¬

tiles fo.ssiles de la collection Nopesa en 1906,

1923 et 1924 (S. Chapman, comni. pers.). Le.s

deux premiers lots ont été vendus par Nopesa

lui-méme et celui de 1924 eût comme intermé¬

diaire le comte L. S/apary, un ministre hongrois

à Londres. La collection de 1906 inclut des spé¬

cimens provenant de Sànpetru» y compris les

types des ornithopodes Rbabdodon rohmtus

Nopesa, 1900 et Telmatosaurus ti-anssylvanicus

Nopesa, 1900 (Weishampel et ai 199L 1993),

ainsi que des restes de titanosaurc'. Le materiel

contient aussi des restes décrits à l’origine comme

appartenant à des oiseaux (Andrews 1913), mais

qui se sont révélés par la suite être de petits thé-

ropodes (voir un sommaire dans Weishampel &

Jianu 1997)t La collection de 1923. acquise pour

200 livres sterling, comprend des spécimens de

Sànpetru et Valioara, notamment les spécimens

types de l’ankylosaure Struthiosanrus transylim-

1. Nopesa utilisa aussi les noms de Limnûsaurus et

û'Onhomerus pour se rapporter à Telmatosaurus, et celui de

Mochlodon au lieu de Rhabdodon (Brinkmann 1988). Il attribua

les restes de titanosaure au genre Titanosaurus. avant que von

Huene ne propose le nom générique Magyarosaurus en 1932.

2. Il faut signaler à ce propos que Nopesa entretint un lien très

niciis Nopesa, 1929, du titanosaure Magyaro¬

saurus dacus (Nopesa, 1915) et de la tortue cryp¬

todire Kallokibotion bajazidi Nopesa, 1923

(Nopesa 1923b, 1929 ; Huene 1932 ; Gaftney &

Mcylan 1992). Les archives du BMNH conser¬

vent des documents relatifs à cet achat, notam¬

ment une liste détaillée des ossements (environ

deux cent soixante-cinq) et le prix ap|)roximatil

de chaque échantillon. D’autres fossiles, compre¬

nant des restes fragmentaires d’ornithopode, de

titanosaure et de rhéropode, ont été donnes par

Nopesa au BMNH en 1909 et 1922. Une autre

collection fut réunie par Lady Woodward à partir

des gisements de Sànpetru et Nagt'-Csula lors

d’une visite dans les propriétés de la famille

Nopesa en Transylvanie. Celle-ci, qui contient en

outre des restes de Rhabdodon^ Telmatosaurus,

MagyarosauniSy Kallokibation et de petits théro-

podes, fut présentée au BMNH en 1923'. Le

matériel conservé a Londres a été décrit par

Nopesa dans plusieurs articles, dont cinq dans la

série dédiée aux dinosaures de Transylvanie

(Nopesa 1900, 1902a, 1904, 1925. 1929 ; voir

aussi Nopesa 1902b, 1923a). Il est intéressant de

signaler qu’une collection de lames minces de

reptiles fossiles, notamment de dino.saures, fut

présentée par Nopesa au BMNH en 1925.

Certaines de ces lames ont été effectuées à partir

du matériel dinusaurien de Transylvanie (Nopesa

& Hcidsicck 1933).

Les collections de reptiles fossiles de Transylvanie

conservées a Budapest au Magyar Allami

Foldtani Intézei (MAFI) incluent principalement

des restes récoltés par le géologue Ottokar Kadic

dans les environs de V'âlioara pour le

Ungarischen Ceologischen Reich.sânstalt. Les

fouilles furent entreprises en 1914 (d’après les

étiquettes du musée) ou 1915 (selon Kadic

1917). Les principaux spécimens ont été décrits

par Nopesa en 1915. Le matériel de dinosaure

comprend des spécimens des ornithopodes

Rhabdodon robustus et Telmatosaurus transsylva-

fort avec la famille Woodward, comme en témoigne la corres¬

pondance de 1906 à 1933 entre le baron et Sir Arthur Smith

Woodward et le journal de Lady Woodward conservés aux

archives du BMNH. A. S. Woodward était le responsable du

déparîemeni de Géologie du BMNH et pendant ses nombreux

voyages à Londres, Nopesa séjourna souvent chez les

Woodward (S. Chapman, comm. pers.).

GEODIVERSITAS • 1999 • 21 (2)

163

Pereda Suberbiola X. & Taquet P.

nicus, ainsi que du titanosaurc: Miigjtarosaums

dacus. Ixs aiares reptiles représentés sont les croco-

dilicns (avec notamment le spécimen-type

diAllodiipostichus precedvns Nopesa, Î928) et les

tortues. Les fonds du MAPI contiennent aussi

des spécimens fossiles beaucoup plus fragmen¬

taires d'une collection récoltée par Nopesa lui-

même (L. Kordos, coinm. pers.). Ils proviennent

aussi de la localité de Valjora mais ne portent pas

de date de collecte. En plus des restes d'ornirho-

pode et de ritanosaure, l’assemblage comprend

un fragment de crâne d‘un théropode

Arctometatarsalia, récemment reconnu (Jianu &

Weishampel 19*^7), des plaques de la carapace de

Kallokibot'fon et des dents isolées de crocodile.

Enfin, des restes de ptérosaute mis au jour par

Nopesa dans la région de Sânpecni (Nopesa

1914) et longtemps considérés comme perdus,

ont été retrouvés dernièrement dans les collec¬

tions du MAI'l (Jianu et al. 1997a)- D’autres

restes de dinosaure ( TelrnatasaurnS} provenant de

Sânpetru sont conserves au Magyar Ncn/eti

Müzeum (MNM) de Budapest (Weishampel et

ai 1993). Ces pièces proviennent du MAFl et

font probablement partie de la collection amas¬

sée par Kadic en I914'1915 (D. Weishampel,

comm. pers.).

La petite collection Nopesa de dinosaure ornitho-

pode du Muséum national d’HLstoirc naturelle

de Paris vient s*ajouter à celles conservées à

Londres et h. Budapest. Dans une lettre adressée a

son collègue et ami Friedrich von Huene, datée

de 1924, Nopesa mentionnait la présence dans le

musée de Munich de quelques os de titanosaure

provenant de IVansylvanie ( Fasnddi-Kubac.ska

1945). Ces os été soir récupérés par Nopesa

après cette date, soit détruits lors dci bombarde¬

ments de 1944 durant la Seconde Guerre mon¬

diale. Toujours est-il qu’aujourdTiuî, aucun reste

de vertébré du bassin de lïateg iVest déposé au

musée de Munich (P. Wellnhofer, comm. pers.).

11 convient enfin de signaler que certains spéci¬

mens fossiles de la région de Hareg ont été per¬

dus. C'est le cas du matériel du dinosaure

cuirassé Onyrhosaurm bungaricus décrit par

Nopesa en 1902 (Pereda Suberbiola & Galton

1997).

De nouvelles coilecrions de vertébrés des gise¬

ments de Transylvanie ont été constituées plus

récemment grâce aux efforts de plusieurs musées

et institutions roumaines. C7csi ainsi que les

musées de Deva et les universités de Cluj-Napoca

et Bucarest ont pu récolter de nouveaux restes de

dinosaures (notamment des chetopodes et des

œufs d'hadrosaure), de crocodiles et de tortues,

mais au.ssi des poissons osseux, des amphibiens et

des mammifères multituberculés (Grigorcscu

198.3, 1984 ; Gtigorescu et al. 1985, 1994 ;

Weishampel étal. 1991 ; Jianu étal. ]9y7b). Un

siècle après la première decouverte de restes fos¬

siles. le bassin de Hateg est devenu une région

fossilifère classique en Europe et une référence

obligée pour les paléontologues qui s’intéressent

aux faune.s de vertébrés finicrétacés.

CONCLUSION

Une mandibule, deux vertèbres caudales et une

scapula de dinosaure ornithopode provenant du

Crétacé supérieur de Transylvanie font partie de

la collection Nopesa donnée par celui-ci au

Muséum national d'Hi.stoirc naturelle en janvier

1923. Apres étude, cc.s restes sont attribués à

riguanndoncia RlmbdodoUy noiammeni d’après la

présence d’un dentaire à bords dorsal et ventral

parallèles portant uniquement dix alvéoles, ainsi

que d’une scapula k bord antérodorsal droit et

montrant une forte proctnincnce sur le bord

postcrovcncrai. Ce matériel fournit des informa¬

tions historiques complementaires sur le sort des

.uicicnncs collections de dinosaures et autres rep¬

tiles fossiles de Transylvanie.

Remerciements

Nos remerciements à Mr. John Thackray

(ATchivc=:s, The NaturaJ History Muséum) pour

avoir permis au premier auteur de consulter la

conespondaiice du baron Nopesa conservée à

Londres, à Mrs. Sandra Chapman pour I accès à

la collection Nopesa de reptiles fossiles conservée

au BMNH. ainsi que pour ses renseignements

sur le matériel fossile, aux Dts Laszlo Kordos

(Magyar Allami Foldtani Tnré/et, Budapest),

Itsvan Fôzy (T'crmészeciudom;inyi Muzeum,

Budapest), M. Coralia Jianu (Muzcul Civilizatiei

Dacice si Romane, Deva), Vlad Codrea

164

GEODIVERSITAS • 1999 • 21 (2)

Rhabdodon de Transylvanie du MNHN

(Universitacea Babes-Bolyai, Cluj-Napoca),

David B. Weishampel (The Johns Hopkins

University, Baltimore) et Peter Wellnhofcr

(Baycrische Sraatssammliuig für Palaoncologic

und hisrorische Ceologie, Munich) pour les don¬

nées fournies sur les collecrions de vertébrés de

Transylvanie, enfin, au Dr. David R. Norman

(Scdgwick Muséum, Cambridge) et à un rappor¬

teur anonyme pour la relecture critique du

manuscrit. Nous exprimons aussi coure nocre

gratitude à Mme Françoise Pilard pour son aide

et a M. Denis Serrerte pour les photographies des

pièces.

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166

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts (Mammalia) from

the médial Cretaceous (upper Albian or lower

Cenomanian) Mussentuchit local fauna, Cedar

Mountain Formation, Utah, USA

Richard L. CIFELLI

Okiahoma Muséum of Natural History and Department of Zoology, University of Okiahoma.

1335 Asp Ave.. Norman, Okiahoma 73019 (USA)

rlcCa)ou.edu

Scott K. MADSEN

Dinosaur National Monument,

P. O. Box 128, Jensen, Utah. 84035 (USA)

Cifelli R. L. & Madsen S. K. 199S. — Spalacotheriid symmetrodonts (Mammalia) from the

médial Cretaceous (upper Albian or lower Cenomanian) Mussentuchit local fauna, Cedar

Mountain Formation. Utah. USA. Geodiversitas 2^ (2) : 167*214.

KEYWORDS

Syiiitnetrodonta,

Spalacotheriidae,

phylogeny,

Cretaceous.

ABSTRACT

Symmetrodont mammals, generally rare and poorly reprcsenred in rhc fossil

record, are exccptionally abundanc in the Mus.sencuchit local fauna of rhe

upper Cedar Mountain l’ormarion (upper Albian or lower Cenomanian).

Enicry County, Utah, USA. Herein we describe three new species of symme¬

trodonts (four or more arc présent in ihc launa); one îs refcrable to

Spalacotheridlums otherw'i.se known from iKc Turonian (Laïc Cretaceous),

and the other wo are referred to a new genus. With rhc possible exception of

Mktodon^ ;ill North American Cretaceous symmetrodonts are rclerable to rhe

Spalacotheriidae. Spalacochcriids are distincrly diflexcnr from more primitive

symmetrodonts .such as Kuchneotherium in jaw structure (e.g., dctachment of

postdencar)' cléments, pre.scnce of pterygoid cresc) and molar morpholog)'

and function (e.g., development of continuons mesial and distal shearing sur¬

faces). To rhis extern, they are more clearly similar to tribosphenic therians

than are archaic symmetrodonts, although they are uniquely speciaÜzed.

Somc featurcs of advaneed spalacothercs, suth as the loss of the coronoid and

meckelian groove, dcvclopcd convcrgcntly in tribosphcnidans and many

other groups, and hence represent iicrarivc thèmes in the évolution of

Mesozoic mammals. Fearures of the molars and deniary suggest chat the

Family Spalacotheriidae is a monophyleric group, with the European

SpaUtcotherium and Chinese Zhnn^heotherium forming successive outgroups

to remaining gênera. Within the family, Norlh American taxa appcar lo

form a monophyleric clade, culminailng in rhc highty specialized

Symmetrodontoidn of lhe Latc Cretaceous; Microdersou, known only by a

single upper molar from the Cretaceous of Morocco, is of ciiigiliatic alTini-

ties. Spalacotheriids were clearly présent in North America by the Aptian-

GEODIVERSITAS • 1999 • 21 (2)

167

Cifelli R. L. & Madsen S. K.

Albian and, assuming chat North American taxa Form an endemic and exclu¬

sive monophyletic group, their présence on the continent cannot be atiribut-

ed to an hypothesized inid-Creraccous interchange with Asia. Instead,

phylogencric data suggest ihcir origin trom western Europe sonietimc in the

Early Crctaccous, siipporring the hypothesis that there was some dcgrce of

faunal continuity at that time between the two landmasses, based initially on

similariries of the dinosaur assemblages.

MOTS CLÉS

Symmetrodonca,

Spalacotheriidac,

phylogénie,

Crétacé.

RÉSUMÉ

Les symmètrodGnies spaincotheriidés (Mammalux) dtt Crétacé (Albien supérieur

ou Cénomunien in férieur) de la faune de Mussenwebit, Formation Cedar

Mountain, IJtah, T/S/l.

Les mammiterciJ symmétrodontes, qui sont généralement rares et mal repré¬

sentés dans le registre fossile, sont exccptionellemcnt abondants dans la faune

locale de Musscntuchit du sommet de la Formation Cedar Mountain »

(Albien supérieur ou Cénomanien inférieur), F.meiy County, Utah, USA.

Nous décrivons ici trois nouvelles espèces de symmétrodontes (il existe au

moins quatre espèces dans la faune de Mussenciiehii) ; une espèce sc rapporte

au genre Spalacotbertdium, connu par ailleurs dans le Turonicn (Crétacé

supérieur), et les deux autres sont attribuées à un nouveau genre. A l’excep¬

tion, peut-être, de Mictodon, lou.s le.s synimétrodonte.s nordaméricaias appar¬

tiennent à la famille des Spalacotheriidac. C^es derniers dillèrent nettement

des symmétrodontes plus primitifs tels que Kuehneotheriurn par la structure

de leur mâchoires (entre autres, le détachement des éléments postdenraires et

la présence d’une crête ptérygoïde) et par la morphologie et la fonction de

leurs mohiircs (eiurc auirc.s, le dcvclt^ppcment de suifaces coupanic.s conti¬

nues mésiales et distales). Sur ce plan, ils ressemblent plus à des thériens tribo-

sphéniques qu’à des symmétrodontes archaïques, bien que présentant des

spécialisations uniques. Quelques caractères de spalacothcres dérivés, tels que

la perte du processus coronoïde et du sillon de niecke! se développent de

façon convergente chez les ïribospbenida et de nombreux autres groupes, et

constituent des thèmes répétitifs dans révolution des mammifères méso¬

zoïques. Les caractères des molaires et du dentaire suggèrent que la famille

des wSpalacotheriidac constitue un groupe monophylétique, le genre européen

Spalacotherium et le genre chinois Zbangheotherium formant les extra-groupes

successifs des autres gcjims. Au sein de la famille, les taxons nordainéricains

sembleni consiiruer un clade monophylétique qui culmine avec les formes

hautement spécialisées comnH' Symwetrodontoides du Oeracé supérieur ; les

alFinitcs de Microdersoru connu par un seule molaire supérieure du Crétacé

du Maroc, restent énigmatiques. Les spalacorheridés étaient clairement pré¬

sent en Amérique du Nord dès l'Albien-Aptien et, si l’on admet que les

taxons nord-américains constituent un groupe monophylétique endémique

et exclusif, leur présence sur le continent ne peut êta- attribuée à un éventuel

échange launique avec l'Asie au milieu du Crétacé. Ln revanche, les données

phylogénétiques suggèrent que l'origine des Spalacotheriidac se situe en

Europe occidentale au Créracé inférieur, renforçant ainsi l'hyporhèse selon

laquelle il existait, â cette époque, une ceaaine continuité faunique entre les

deux continents, hypothèse fondée iniiîalemcnr sur les ressemblances obser¬

vées entre les faunes de dinosaures.

168

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from rhc mid-Cretaceous of Utah

0 10 20 km

I_I_I

INTRODUCTION

Symmetrodonts hâve long been accorded a criti-

cal position in mammalian évolution because the

principal cusps of iipper and lower inolars form

the “reversed triangle’" pattern that is widely

bclieved to be morphologically intermédiare bet-

wecn the serially tricuspid condition of tricono-

donts and the more claboratc molars of

tribosphenic mammals (e.g., Patterson 1956).

Unfortunately, tbeir fbssil record is abysmal, and

chcy are probably the worst representcd of ail

Mesozoic mammal “groups” (in a inorphological,

not taxonomie sense) - ilie receni discovery of an

exceptional spccimen from the IvUtc Jurassic or

Early Cretaceoius of China (Hu et ât. 1997,

1998) notwithstanding. Given the fact that

knowledge of symmetrodont diversity and mor-

phologv’ is poor, it is unsurprising that they are

generally omitted from compréhensive analyses

of mammalian phylogeny (e.g., Rowe 1988;

Wiblc 1991; Wibic ôi Hopson 1993; Rougier

et ai 1996), notable exceptions being the works

of Prothero (1981) and Hu et al, (1997). In

Fig. 1. — Outerop map of the Cedar Mountain For-

k mation, Utah (inset), and field area in Emery County.

Localilies are OMNH sites that produced specimens of

C Spalacotheriidae described in the text (see Cifelli et ai

y 1999, for sections showing stratigraphie positions of

( sites).

North America, symmetrodonts were long

known from the Lafe Jurassic only, as represented

by Tinodofi Marsh, 1879 and the probably syno-

nymous Eurylambda Simpson, 1929 (sec

Simpson 1929; Crompton & jenkins 1967;

Prothero 1981). Latcr discoveries rccordcd the

survival of apparent ‘*acure-angled” spalaco-

theriids, hitherto represented uniy in what wa.s

then considered the Jurassic of Hngland, in the

Early Cretaceous of Texas (Patterson 1955,

1956) and the Lare Cretaceous of Alberta (Fox

1972. 1976, 1985) and Southern Utah (Cifelli &

Madscn 1986; Cifelli 1990). Mictodon Fox,

1984, an apparcntly relictual taxon from the

Campanian of western Canada, represents the

only record of an 'obcuse-angled” symmetrodont

from the Cretaceous of North America (Fox

1984b).

Herein we describe new taxa of intermédiare âge

between these Early and Late Cretaceous records

in North America. One of the taxa, at least, is

represented by an uniisually comprehensive

sériés, affording the opportunity to examine

variation and positional changes in the molar

GEODIVERSiTAS • 1999 • 21 (2)

169

Cifelli R. L. & Madsen S. K.

scries, and prcsenting new information on chc

dentary oF advancfd Spalacothcriidac. Finally vvc

bricfly commciu on rhc stacus and placement of

thc Spalacoihcriidae wirh respect lo other sym-

mctrodonts and to more advanced mammal

groups, and provide an hvpoiliesis of relation-

ships within Spalacotheriidae.

rhc symmctrodont.s described hercin \vere col-

Icctcd hom thc Cedar Mountain Tormation,

Emcry County, Ulah. I his unit was named for a

sériés of terrigenous scdiincnlary rocks lying bet-

ween thc LJppcr jurassic Morrison Formation

and rhc Upper Cretaceous Dakota Formation

(Stokes 1944, 1952), and is broadly exposed in

central and e;isrern Utah {Fig. l). Five units (in

ascending order) of thc formation arc now reco-

gnized: thc Buckhorn Conglomerate, and ihe

Yellow Cat, Poison Strip Sandstonc, Ruby

Ranch, and Musseiuuchit members (Kîrkland

et al. 1997). The symmccrodont spccimen.s

resulted from a conccrtcd collccting effort in a

restricted stratigraphie intervaJ of the uppermost

unit, the Musscntuchit Member. 10-20 m below

thc contact with thc ovcriying Dakota For¬

mation. The spccimens described hetein dérivé

from eight sires (Fig. 1); the vast majority w'crc

collected from OMNH locality V695. Fhc fossil

horizon ac rhis locality ts dircctly oveiTain by a

volcanic ash. Multiplci concordant ‘^^Ar/^’^Ar

decerminâtions on .sanidinc phcnocfyst.s from

this ash, and from rhc samc horizon ncarhy^yield

a date of 98.39 i 0.07 Ma (Cifelli et ai 1997);

hcncc, the fauna is indistinguishabic in age from

the Albian-Cenomanian (Eatly-Latc Cretaceous)

boundary, placcd at 98.5 ± 0.5 Ma hy

Obradovich (1993) and ac 98.9 ± 0.6 Ma by

Gradstein et i?/. (1995)- Strarigraphic sections

showing placement of thc principal fossil locali-

ties arc given in Cüfelli ei al. (in press),

The vertebrate assemblage from the upper part

of the Cedar Mountain Formation, lermed thc

Mussenruchit local fauna, is known by more

than 5000 spccimens representing about 80 taxa

(Cifelli et ai 1999). Of rhe mammap, only the

marsupial or near-marsupiat Kokopellia Cifelli,

1993 (see Cifelli 1993; Cifelli Ôc Murzon 1997),

three triconodontids (Cifelli & Mndsen 1998),

and several multicuberculates (Eaton 6c Nelson

1991) hâve been described thus far.

METHODS

Specimens were recovered using a combination

of standard quarry procedures, through which

mosr of the dentulous jaw fragments and a fcw

of rhe larger isolatcd teeth were reœvered, cou-

pied wirh a largc-scalc underwater scrcenwashmg

operation (Cifelli et ai 1996: Madsen 1996). It

is wortbwhile pointing ont chat most of the iso-

latcd reerh were recovered from rhe fine fraction

of internested screen boxes, in which the corres-

pnnding screen si/.e was 30'mesh; had only wln-

dow screen been einploycd, as is commun

practice for Latc Cretaceous rocks of tlic DS, few

of thèse spccimens (rnost ol which bave a maxi¬

mum dimension of significancly less than I mm)

vvould liavebeen recovered.

Measiirernenis were raken with a Reflex micro¬

scope. which permits non-contact recording of

point coordinates in three dimensions; minimum

standard errors are ivvo microns on the x, y-axes

and five microns on the /.-axis (MacLarnon

1989). Reflex data nre autoniatically rccordcd to

0.001 mm, and rhese data arc reproduced Verba¬

tim here, alvhough we point oui that rhis dues

not lâke into accouni measuremem error (see

Liilcgraven & Bieber 1986).

Measurenients are shown in Figure 2.

Spalacotheriid molars are cxrremely small and

fragile; the luwer molar cingula are particularly

vulnérable to breakage. In order to maximize

sample si/.e foa* lower molars, we took standard

length and wîdth measurcmenrs minus the cin-

guluiu. We rerneUsured spccimens of other rele¬

vant Spalacrjtheriidae using tlie same procedure;

measurcments of Spalacotheroidcs bridwelli

Partersont 1955 are from an epoxy cast, and

those of Synmiet>’odontoidvs canadernh Fox, 1976

are from Fox (1976, fig. 5; 1985, fig. 1). Other

measuremencs were taken by defining points at

thc apiccs of chc primary cusps (pataccmid, pro¬

toconid, mctaconid) and calcuUiing: (1) the dis¬

tances between chem; (2) thc angle (hcrcin cailcd

rrigonid angle) formed her\vccn the points, with

the protooonid at thc apex. l‘hc trigonid angle is

tacher variable, even among teeth of chc sanie

locus. Considération of tooth niorpholog)' sug-

gests that the trigonid angle decreases with wcar:

the mesial and distal faces of the paraconid and

170

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmecrodonts from the mid-Cretaceous of Utah

angle

Fig. 2. — Spalacotheriid molars showing measurement conventions; A, right lower molar, occlusal view: B. right lower molar oblique

occlusolingual view; C. left upper molar, occlusal view. Abbreviations: angle, angle lormed by lines connecting apices of paraconid

to protoconid and metaconid to protoconid; ANW, anterior width (greatest width of lower molars); L. mesodistal length; pad-med,

distance from apex of paraconid to apex of metaconid: POW, posterior width; prd-med, distance from apex of protoconid to apex of

metaconid.

metaconid (respeccively) are rather vertical ^ where-

a.s thcir opposing faces slopc downward toward

cach other, so that thc apparent ccnters of the

cusps will migiatc toward cach other as wear pro¬

gresses. Wc also attempted to cake heighr measu-

renients. However, our efforts were frustrared by

our inability to define a repearable plane of réfé¬

rencé, and the fact that wear varies considerably

from one specimen to the next. Hence, réfé¬

rencés to différence in crown heighe arc qualita¬

tive only. For upper molars, we took measure-

ments (Fig. 2) analogous to those employed for

tribosphenic therians (see Lillegraven 1969,

fig. 5). Calculations, descriptive statistics, and

tests were donc vvith Systat version 7; original

data are availablc from the senior author upon

rcquest.

Dental terminology is shown in Figure 3. The

homologies of some of the upper molar cusps of

spalacotheriids and other symmetrodonts - e.g.,

the presence of a metacone (Butler 1939;

Patterson 1956) — are unclear, and the nomen¬

clature is inconsisrent. Most w'orkers hâve refer-

red to the three primary cusps of upper and

lower molars in primitive mammals as A, B, and

C; and a, b, and c, respectively (e.g., Crompton

&c Jenkins 1968; Cassiliano ôc CÎemens 1979;

Jenldns & Crompton 1979). Crompton (1971)

regarded the metacone of tribosphenic Theria as

a neomorph, and referred lo che disiolabial of the

three primary upper molar cusps as cusp we

follow convention in referring to this as cusp C,

in order to avoid confusion with lower molar

cusp c, thoiigh we point out that similar pro-

blems exist with this term, as upper molars of

primitive marsupials and certain other tribo-

GEODIVERSITAS • 1999 • 21 (2)

171

Cifelli R. L. & Madsen S. K.

distal stylar cusp

paraconid metaconid

Fig. 3. — Dental terminology employed in this paper ; A. upper molar (M4) of Spalacotherium {modified after Simpson 1928a, fig. 34,

and Patterson 1956. fig. 12): B. upper molar of Spalacolestes', C, lower molar of Spafacolesîes.

sphenic mammals hâve a stylar cusp C (see

Clemens 197*^-. we thank R. C. Fox for poincing

this out to us). Olher auihors (Butler 1939;

Patterson 1956; Kermuck et al. 1968; Prothcro

1981; Fiopson 1997) intcrprct cusp C — which

in spalacorhcriids (where présent) is located on

the postparucrista about niidwuy bctween the

paracone and the disiolabial corner of ihc tooth

- as homologous with the tribosphcnidan meia-

cone. Füllow'ing argunients presented by

Sigogneau-Russell &: Ensom (1998), we believe

this to be probable, but follow Hu er al. (1998)

in retaining the traditional nomenclature forspa-

lacorheriids. Certain spalacotheriids also have an

unusual cusp on the preparacrista, about halfway

between the paracone and the mesolabial corner

of the tüOth, where a second cusp is generally

présent. We follow Patterson (1956) and

Sigogneau-Russell &: Ensom (1998) in regarding

the latter cu.sp as the styloconc (see also

Sigogneau-Rus.sell 1991a), so rhat the cusp lin¬

gual to it (but labial to the pajaconc) is a neo-

morph. The most rcccndy applied term for this

cusp in the middie of the preparacrista is cusp Bj

(Hu et al. 1997), and ihi.s usage is adopted

herein. Symmetrodonc upper molars also com-

monly bcar one or more cusps placcd on chc sty¬

lar shelf, distal to the ectoflexus (if one is

présent). Thac ar the corner of the tooth may be

termed, by convention, the metastyle. The more

mesially placed cusp has been referred to as a

posterior stylar cusp (e.g.. Fox 1985) or, in aiia-

logy with the siniilarly placed cusp of triho-

spheoic therians (e.g.. Simpson 1929; Clemens

1979; Fox 1984a), as stylar cusp D (e.g.,

Sigogneau-Russell 1991b; Sigogneau-Russell &

172

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Ensom 1998). Unforrunarely, the term “cusp D”

has also becn applicd to the metaîjtylar cusp (Hu

et al. 1997). In ortier to avoid confusion with

this usage or implicd homology with the similar-

ly-posirioned cu.sp of marsupials, placcntals, or

ihcrians of ‘'inetatherian-euthcrian grade," wc

refer to the cusp placed on the stylar shelf, distal

to the médian part of chc tooth but proximal to

the metasrylc (with which it should not bc

confused), as a "distal siylar cusp.”

The médial surface ol the deiitary in mammals

commonly beats a ridge, crcsi, or analogous

structure, gciierally near or ai lhe inferior margin

and located posterior to the mandibular fora¬

men, for attachment of the m. pterygoideus

medialis. We are unabic to find a standard anato-

mical term foi this structure, and various names

hâve been applied to it in the literature. Simpson

(c.g., 1926, 1928a) alternarively referred to this

structure as u "pterygoid crest" or “prerygoid

ridge,” soinecmies using botb ternis in the sarne

Work (e.g., Simpson 1929). Recent authors (e.g.,

Rowe 1988) somenmes refer to it as a "pterygoid

shelf,” and this term has become standard for

multiruberculates (e-g.^ Miao 1988; Gainbarayan

dit Kielan-Jâworowska 1995), in which tlic infe¬

rior margin of the dentarj’ is strongly inflecied

lingually. In order to promote précision in usage

and to avoid confusion in characcer srate or

implied homology (see discussion in Miao

199.5), we refer ro fhe structure in question sim-

ply as a "pterygoid crest," cxccpt wherc il is

obviously devclopcd inco a shelf, as in multi-

tuberculates, or into an inllectcd angle, as in

marsupials and some early Eutiieria (see Sanche/-

Villagra ôc Smith 1997, andbclow). At least, one

of the specics described hcicln is cburactcrizcd by

a pterygoid crest that bcars a hyperttophied,

process-like lingual extension that is unique,

so far as we are awarc. Lacking any standard

term for this feature, we refer to it as a "pterygoid

process.”

AlîBREVIA'nON.S IOR INSI ITUTIONS IN THE

TKXT

BM British Musciim, London, UK;

UK FMNH, Ficld Muséum of Natural

Hisiory, Chicago, Illinois, USA;

(il PST Institute of (ieology. Section ofPalaeonco-

logy and Srrntigraphy, Mongolian

Academy of Sciences, Ulan Bator, Mogo-

lian Peoples' Republic;

MNA Muséum of Northern Arizona, Flagsiaff,

Arizona, UvSA;

OMNII Oklahoma Muséum of Natural History,

Norman, Oklahoma, ILSA;

UALVT Uriivcrsity of Alberta Laboratory for

Vertebrare Pale4)ntology, Edmonton,

Alheria, Canada;

USNM National Muséum of Natural History,

Smiilisonian Institution, Washington,

D.C, USA;

YPM Yale Peabody Muséum, New Haven,

Connecticut, USA.

SYSTEMATIC PALEONTOLC9GY

Order SYMMETRODONTA Simpson, 1925

Cc)MMl.NTs. — rhis group was proposed by Simpson

(1925a) ro include rhen-known taxa (ali thought to be

Jurassic in age) having the three principal molar cusps

arrangecl in a triangular pattern, rherehy disringuLsh'

ing rhem from the serially iriciispid iriconoJonts,

wiih which they liacl heen prcviousiy groupecl {c.g.,

Osborn 1888, 1907). Symnicrrodonta h:ivc long been

conceivcd as u pataphyleric group (sec, c.g., Paitcrson

1956; Cas.siÜano àL Cilernen.s 1979, hg. 7-4), but tins

simple picture becamc increasingly complcx with the

discovery ol geulogicilly oldcr taxa, particularly

Rhacto-Liassic Kuihruvlherimn Kemiack et al., 1968

(sec Kermack e/ ai 1968) and, l.U'cr, Wouiersia

Sigogneau Russell, 1983 (see Sigogneau-Uussell 1983;

Sigogncau-Riissdl & Uahn 1995). Mandihles referrctl

ro Knehricoiherium. ai least, ictain a po.srdciuary

trough and attachment lacets (or ihe postdeiuar)' clé¬

ments seen in cynodonrs and primitive mammals such

as MtirganuviitliiH Külinc, 1949 ind docodonrs

(Kcrmack Musxett 1958; Kermack ci al. 1968;

Kermack er i^/. 1973; I illegravcn & Krusat 1991), and

lack derived features (silch ps a pterygoid crest) fouiid

in other mammals, meluding some iriconodoncs (c.g..

Rowe 1988; Wible 1991; Wible 6i Hopsori 1993).

Hcncc, eiihcr ihc mammalian middic car complcx or

the **revtrsed iriangle" pattern ol upper and lowct

molars evolved independciuly more than once; boih

cavscs bave been argued (sec, c.g., Allin & I lopson

1992; Roiigicr (T 1996). Prornero (1981) rcsoivcd

this probk-m hy excluding Kuchnctithcrium from

Symmetnxionta (sec aUo I lopson 1994), but his ana¬

lysis did noi incIudc non-tncrian groups, and it is

undcar how other. more rcccntly dc.scrihcd rax.i (c.g.,

WotitersiiK Zhitnghcotheriufn Hu et uL^ 1997,

Thereuodoii Sigogneau-RussdL 1989, Shaothcrtuni

Chow & Rich, 1982, Kotatherium Datca, 1981, and a

host of enigmatic taxa from the Campanian Los

GEODIVERSITAS • 1999 • 21 (2)

173

Cifelli R. L. & Madsen S. K.

Alamitos fauna i)f Argcniina: see retcreiices cited

above and Daic.i 1981; Chow & Rich 1982;

SigogneaU'kus.scII 1989: Bonaparte 1990; Sîgogncau-

Russell 1991 b; FVasad & Manhas 1997: Sigogncau'

Russell & linsoiTi 1998) would fit inco this schemc.

The situation is furchcT complicated by the laer that

some molars ot AmphÜcstidae, generally placed in the

Triconodonta (e.g., Üîinpson 1945), bave their princi¬

pal cusps arranged in an obtuse triangle, and a

relationship to SyinincTrodonra lias been suggesicd on

ihis basis (Mills 19"^!; see aUo Icnkins &c Scri;itT 1988;

and discLjsçion in Kieliyn^Iaworowska èc Dashzeeeg

1998). Fox (1985) proposée! a rrif'old classificadon ot

Symmetrodonta. including Tinodoniklac (coniaining

Kuebneothn'ium and scvcral oihcr taxa, as wcll as

Tirjodon). Kuehneathmum has heen shown to be high-

ly siniilai tu Tinodim (e.g.. Crompton ifc Jenklns

1967), but ils platcineut in ihc Tlnudotnidac is pla-

gued by flie sanie ditficulîy as irs referral ro

Symmetrodonta in general: ir rerains an extreniely pri¬

mitive javv structure, whereas in Titiodon the post-

dentary éléments were evidenily detached and a

pterygoid crest is présent (see Fruihero 1981). We qui

offer nothitig new lo solvc this dilemma and ibus hâve

not artcmpicd lo dePinc or dlagno.se Symmerrodonta.

However, there arc some data to uphold integrity ol

the “core" group, Spalacotheriidac. Pending hirther

analysis and, bopelully, nmre data Icum nie tossil

record, wo bnd it uscful ro reinin a iniditional, incln-

sive concept ot '‘synimetrodonrs” (c.g., C^assiliano &

Clemens 3979; Fox 1985). A rmmr. compréhensive

discussion of (lie prohicm in derming Symme-

trodonra. together vvitli a tliorough historical review

of relevant rax;u is given by Sicogneati-Russell &

Fnsom (1998)- McKcnria iU Bell (1997) distiibuted

ihc contents ot ibc Symmetrodonta aniong several

higher gronps within Mammalb, which rhev diagnos-

cd primarjly ou the basis pf derachmenr of acce.vsory

jaw bone.s (posrdeniary complex) trom the cranio-

inandibular joint and their association with the cra-

nium as éléments of the nuditory apparatiis. I his

arrangement implics a rcvcr.sdl for Ktiahneothifrium

which, as noted, cvidcntly retained a fuil complément

of postdentary element.s chat wcic wcll integrated with

the dentary.

Recently, the tenu “ I beria'’ lias been tortnaUy dclnied

as a crown-based raxon restricted to the common ances-

tor of marsupials, placentaU. and ail of iis dcscendant-s

(Rowe 1988). FIcrcin wc follow a more tradiiionaL

informai concept that aiso includes "Theria of meta-

therian-eutherian gradcT peramuruns, eupantotheres^

and symmetrodoncs (e.g., Patterson 1956), in récogni¬

tion of die current instability in phylogenetic interpré¬

tation of the major groups ofmaninials.

Family Sl'Al.ACOTHFRnDAL Marsh, 1887

Type GENUS. — Spalacotherium Owen, 1854.

iNcl.uni I) t,i;Ni:RA, — The type, and SpalacoiheroicUs

l’atterson. 1955; Symmetrodontoides Yox, 1976;

Spalactithcridium Cilcili. 1990; Microderson

Sigogneau'Russcll, 1991; Zhangheothenum Hu ei al.,

1997: and Spalacolates^ n. gen.

DiSTHiBi'riON. — ?Làtc jurassic through Ea-rly

Cretaceous, western Europe (Cleniciis 1963: Clenicns

& Lees 1971; Krebs 1985); Faily through Lare

Crciaceous. Nortb .America (Panctson 1955; Fox

1976); Late Jitravsic or Early C'rccaceous, Asia (Hu

et al. 1997): ?Early Cretaceous, northern Africa

(.Sigogneaii'Russell 1991b). Spalaaitherium was First

descrihed frotn ihc Purbcck beds, iradilionally regard-

cd as Upper liira.ssic (sce di.scu.ssion in (.’lernens ei ai

L979). Récent litetature increasingly refers the

nianimal-beiiriiig part n( the Puii>ct.k ro the Bcrruisian

(l.ower Crciacciujs, sec Allen (k Wimliledon 1991;

Kiclan-Javvofowska & Enaom 1994, Sîgogneau-

Russell & Ensom 1994; Ensom & Sigogncau-Riissell

1998).

Kl-VISEU I ïIAGNO.SI.S. — Sytnrnerrodonvs wiih lower

molars bcaring wcll-dcvelopcd primary cu.sps (para-

conid, protoconid. meraconid) arranged in an acute

angle, a reduced lalonid; fivc lower molars pivsciU in

Ztfanght‘othenuia, incrcasing to six or more, where

known, in orher laxâ. Uniï]ae pattern of inrerlocking

for lower molars, whereby the distal cingular tusp of

one molar is placed labial to the mesial cingnlar cusp

of rhe snccccding tooih. Upper molars piimitivcly

wnh acce.ssory cusp (Bj) on preparacrista beiween

paracone and stylocone.

COMMENTS

A more detailcd diagnosis of Spalacotheriidae

was given by Fox (1985)j bascd on rhen-known

taxa: SpciliiCuthcriuni^ SpuLicothetoide^^ and

SyjNmetwflohtoklt'i. The concept of the lamily is

broaderifd here lu include Zhanglnothcrium,

recently dcscribed from the Uatc Jura.ssicoi\ more

probably, Early Cretaceous of China (Hu ai

1997). By compari.son to icinaining spalaco-

thcMcs, /Jnmghcothmum wmld appcar to bc pri-

micive in some respects, such ,is rhe lower

niimber of mol.irs, ?bck of cotifinuoas mesial

and disral shearing surlacc.s on molars (upon

éruption), and. perhaps, fearures on the mcdial

sidc of the dentary (see below). In other respects,

such as the complété lack of cingula on the lower

molars, Zhangheothenum is strikingly atypical.

Nonethcicss, molar morphology is otherwise

similar to that of Spa'lacütherinmy particularly in

the presumably derived features cired in the dia-

174

GEODIVEflSITAS • 1999 • 21 (2)

Spalacotheriid symmecrodoncs from the mid-Cretaceous of Utah

gnosis. Wc tcntativcly follow Hu et aL (1997) in

rcferring Zhanf^heothcriiim to ihe Sp^laco-

theriidae. Cusp R, is presenr in Spalacotheriuyn

(e.g., Clemens 1963), Spalacatheroides (sec

Patterson 1956), imà Zhangheolbernim (sec Hu

et ûL 1997), rhc geologically oldcsr and, [or rca-

sons detailed below, considered by us to orlier-

wise be thc mosr primitive members ol the

Family. Wc therefore tentarively regard tlie pré¬

sence ot cusp B| to be primitive for (and dia¬

gnostic oF) Spalacotheriidae. Sigogneau-Ru.sscll

&C Ensom (1998) considered ihe loss of

Crompton (1971) s lacet A as characteri/ing

Spalacorheriidae. Ii i-s presenily uncertain as to

whether or noi lacet A is seen in Zhungheo-

theriîim, and this fcatiirc bas accordingly been

omiteed From the diagnosis, pending detailed

description of that taxon.

MierndersoHy rcprcscnced by a sitigle tootb belong-

ing lo ihe type and only spccies, M. lanronssii

Sigogneau-Rtisscll, 1991, From chc Early

Cretaceous of Morocco, vvas initially described as

a spalacotheriid (Sigogncau-Russcll 1991b), but

ils pertinence to tbc fiimilyhas rcccnrlv been cal-

lcd into question (Sigogncau-Russcll ^ Ensom

1998). We tcntativcly ineJude it Itère For thc sakc

of completcness but, bccausc it is poorty known

and of enigmatic aFfmities. inake only pa.ssing

référence to îc in the comparisons beiow.

Given the low known diversiry oF symme-

rrodonts atid their mcager reprcsentacion in rhc

Fossil record, ir contes as so(ncwbat of a surprise

that several species, collectively tepresented by

more thaii 250 spécimens, are presciu in tlic

Mussentuchit local fauna. Most of lhese spéci¬

mens are isolated teeih, ntany oFwhich are worn

or incomplète. I hist couplcd with dte Facts chat

the specicvS arc quitc similar to cach other and to

named taxa Front North America, chat thc rooch

rows include ntany molars (probably seven in the

lowcr sériés and six in the upper sériés) that are

rather simple and vary in only subdc ways front

one position to the nexc, and tliac itt> closciy

similar taxon is known hy anything close to a

complété dentition, inakes identification ol

taxon and toorli position less ihan straight-

Forward (see alsti Mills 1984). Wc gave lirsi consi-

der-ation in our analyses to the lower molar

sériés for several reasons. First, the sample of

lower molars is comparatively large, ihus permic-

ting slalistical ircatment and somc appraisal of

variability. Second, thc most morphologically

informative spccimens, hoth from the Cedar

Mountain formation and eisewhere (e.g., Fox

1976), are dentigerous mandibular fragments. A

final, most compelling rcason for giving primary

consider-ation to the lower molars is that the

holotypes ol ail spalacotheriid symniecrodonts

(excc'pL Microdersnn) include lower molars, and

most spccies arc bascd on lower molar séries or

individual teerh.

rite less numetous upper molars were rhen assi-

gned to lower molar taxa based on size and mor-

pbological considérations, and were .sorted to

locus (see beluw). judged From thc compo.silion

of the Mussetiiuchii local Fauiia (Cifclli et al.

1997, 1999) and coinparison wiih Spataco-

theriunt (see Simpson 1928a), .spalatotheriid pre-

molars are eJearly presenr in tbc existing collec¬

tion. No attempt was made co sort these

according to taxon, however, because of the diffî-

culties posed by assignment to locus; hencc, only

the molar dentition is considered herein.

I.OWTR MOLARS

Foriunarely, one of the taxa from thc Ciedai

Mountain Formation is known by a rclativcly

cnormous sample, several jaws arc included, ïtnd

as a rcsult the lasi four molar loci of rhe mandi¬

bular dentition arc represented by cccth in place.

A deiuary fragment pre.serving the amerior pan

of the molar sériés, including three loci, is known

for the closely similar Sytrnuetrodoaioidcs canU'

demh (see Fox 1972, 1976). Fhcsc spccimens,

togethet with comparison to che complète denti¬

tion pf thc somc-what ntore divergent Spalaco-

therium (see Simpson 1928a; Clemens 1963),

provide the Foundation for evaluating position of

isoiared reeth on thc hasis ol relative crown

height, dillcrcnccs in thc proportions ol thc lin¬

gual cLisps (paraconid, mctaconid), acuteness of

the rrigonid angle, configuration of thc lingual

cingulum, width to Icngch proportions, and

other considérations.

Once the lower molars were sorted to cheir relati¬

ve and approximate positions in the tooth row, it

bccamc apparent chat chrcc diagnosablc spccies,

differing in size (Fig. 4) and morphology, were

GEODIVERSITAS • 1999 • 21 (2)

175

ANW m5 ANW m3 ANW m1

Cifelli R. L. & Madsen S. K.

L m5 L m6

Fig. 4. — Bivariate scatterplots for lower molar length (L) and width (ANW) of Spalacotheriidae from the upper Cedar Mountain

Formation by inferred tooth locus. Symbols: circles, Spalacoîheridium noblein. sp.; squares, Spalacolesîes creîulablatta n. gen.,

n. sp.; triangles, Spalacolestes inconcinnus n. gen., n. sp.; diamond, gen. and sp indet.

présent in rhe sainpie; the well-represeiucd taxon represcnis a fourtli, unidentificd taxon. For the

just meniioned, knovvn by several jaws and near- ,spccic.s rcprç.scnicd by the largosi sample> il was

ly 100 isoFated molarst a .smaller .specie.s, repre- then po.ssiblc to condiici deiailcd comparisons in

sented by about 3^ isolaied teeih; and a larger order to ideinily tooth locus more prcciscly» eVa-

species, for which only thrcc lowcr molars are luace variabiliry ac cach looth position and ditt’e-

known. One problematic specimen evidently rences berween adjacent teeth (in ternis of their

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts trom the mid-Cretaceous of Utah

mea.surcmciusj, and estimatc* tlie niolar couru.

These results wcre thcn uscd for rcfincment of

identifications for thc othcr spccics. Description

of the resuit.s is deferred to thc individuol species

accounts. but some prchitoiy comnients are war-

ranted here becuuse of thc nccd to provide a basis

for comparisoii with prcviously dcscribed species,

and the iniplicalions of thc results prcsented

herein for interprcuuion of toorh position among

spccimens rcferrcd to thosc species.

As rnentioacd, the best-represented spalaco-

theriid of the Mussentuchii local fauna is known

by thc lasc four lowei molars in place, as well as

by a large samplc of isolatcd teeth. Excepi for the

lasr niolar, which is morphologicaJly distinctive,

it is not possible lo ideiuity positions of indivi-

dual posrerior inolars vvîth certainty because of

overlapping ranges of variation between loci.

Nonethcless, thc isolatcd molars fàll imo reco-

gnizable clusters corresponding to known toorh

positions, and wc consider uur identifications to

he probable, with niistaken identiry by more

than onc tooth position being unlikely.

Among the isolaved reerh rcprcscniing anterior

mular loci. one group is consistently and clearly

recogni/able. Based on known rrends in the denti¬

tion of spalacotherjids (e.g.. Simpson 1925b,

1928b; CIcmens 1963; Fox 1976), tliese teeth can

bc confidcntly idencified as ml. I he teeth ihat aie

morphologically intermédiare beiween this first

molar and rhe anreriormosi of ihose in known

cooth positions fall into vwo clusters, recognizablc

on thc basis of both qualiradve and quantitaiivc

chaiacteristics. Again,. huwevcr, because of over¬

lapping ranges of variabiliiy, individual teeth can

be identified wilh probability, not certainty,

according to position. Hence the available data

indicatc that, in rliis species at least, seven lowcr

molars wcrc présent, Rachcr than using quotation

or question marks on thc numerous spccimens

ciced below, wc simply aill attention to tlie facts

chat a complété dentition of a Norrh American

spalacotheriid remains unknown and ihat, in any

case. Identification of isolated molars (except thc

first and last, wliicli are morphologically distinc-

rive) cannor be esrablished unambiguously. even

wirh complète spccimens ac band for comparison.

The identincacions of teeth tu respective loci

should be regarded as tentative.

Of lhe remaining spalacotheriids from the

Mus.scnruchit local fauna, comparison wiili the

taxon jusi mcniioncd indicated that a second

species is represented by iccth recogniz.able as

belonging to thc fir.si six lower molar positions;

rhese will be rcferrcd to as ml-6. h is unclear as

to wheihcr an m7 was lacldiig in this species, or

is not represented in exiscing collections. The

rhird diagnosablc species is known by only three

lowcr molars, each recognizable as ro locus with

thc samc confidence as rhe morphologically simi-

lar, best-represented species.

Among described species of Non h American

Spalacotherüdae, the best known is Symmetro-

dontoidei canadensh, the holotype of which (UA

8588) consisis of an incomplète jaw with three

teeth. rhese werc idcmiticd by Fox (1976) as

probably represcnting m3-5. The basis for tins

ideniîHcation was a referred specimen, UA

12086, a mandibular fragment with a molari-

form looth considered ro be ml - The cooth of

UA 12086 bas a broadly obtuse rrigonid angle

chat, if compared co UA S588, would form a

graded sériés witli the teeth on thc lattcr spcci-

rnen, assuming that an unrepresented tooth posi¬

tion intervened beiween thc two spccimens.

Dentigerous jaws and numerous isolated teeth

similai' CO chat of UA 12086 are known from the

Odar Mountain Formation, and work in pro-

gress by one of us (RFC) indicares tliar tliese are

noi lowcr molars of spalacotheriids. Cairnparison

of UA 8588 to rite extensive sériés meniioncd

above indicates rhar the teeth in this specimen

arc mI-3. fox (1976) aiso referred an isolated

molar, UA I20S7, to Symmeirodontoldes mnit-

demis. As dcmonsirated by Fox. this tooth clearly

represents a more posterlor looth locus.

Considération of the proportional différences

between this specimen and m3 of the holotype,

together widi chc large samplc from the Cédât

Mountain Formation, suggesrs th;U UA 12087

probably is an m5 or, as Fox (1976) suggesied,

m6.

(3cher species are knowli by isolated teeth only,

and identification of looth po.siiion is more pro-

blematic. The first symmeirodont to be des-

cribed from the Crctaccous of Norch America is

Spcdncolheroides bridwelli, from rhe Aprian-

Albian of the Trinity Group, Texas. S. bridwelli

GEODIVERSITAS • 1999 • 21 (2)

177

Cifelli R. L. & Madsen S. K.

was based on a mandibular fragment wiih a

single lower molar (f*anerson 1955); scvcral

upper molars wcre larer referred to the spccies

(Patterson 1956). Patterson ( 1955) suggcstcd

rhat, as wirh Spalacotberhnv-, SpalavoTberoides

probably had seven molars, and rliac the tooth in

the hülotype (FMNH PM 93.5) is tbe ancepcnuP

timatc - i.e., ni5. Pox (1976) indicated rhat this

tooth bcst malchcs rhc second rooth on chc holo-

type of Synuneirodontoide^ catiadensis^ then iden-

tified as m3 or 4 and considered herein to be

ni2. We concur: in rerms of size and overall mor-

phology, we find PMNH PM 933 co bc most

similar to m2 or 3 SymmetrocloHtoides.

However, rite paraconid on PKINH PM 933 is

lacking. lx)r rhU reason, and because tooth locus

cannot be reasonably hypothesized without ocher

specimens belonging to the same species, the

lower dentaion oï SpitLicotheroides hridwelli tnasi

be set asidc kom comp.irisons for the tirne bcîng.

This species is said ro ditfer froni Spalucotherium

(see Patterson 1955) and ail other Spalaco-

theriidae (Fox 19^6) except Zhimgbeoîhermm în

having an incomplète labial cingulum. No otlicr

lower molars are yet known for S. bridwetlf. A

somewhav more obtuse-angled tooth (perhaps

ml), comparable in si/.e and overall gestalt to

FMNH PM 933, is now known from rbe

Cloverlv Formation (RLC, im[>ubli.shed data),

which is approxiniatel) équivalent in .ige ro the

part of* the d'rinity Group that produccd

Spidacotheroidvs (Jacobs et al, 1991). Unfortu-

nately, the labial side of this tooth is damaged,

leaving open tlie question of whether or not the

cingulum was complété,

Three spedes of Spalacotheriidac hâve been des-

cribed from chc Upper Crccaccous of Southern

Utah. SyninieirvdoiHoides foxi Cifelli <5^ Madsen>

1986, from the Wahw'cap Formation (assumed

to be lower Campanian and approximatcly équi¬

valent ro the upper Milk River Formation,

Alberta, which produced S. awadensiî), was

based on presumed m4 (MNA 4589, the holo-

type) and u referred tooth (MNA 4522) a.s.sumed

to rcprescnc m7 (Cifelli ik Madsen 1986). A new

specimen of this species. together with compari-

son to the extensive séries representing a morpho-

logically similar spçcics (rom the Cedar

Mountain Formation, suggests that MNA 4589

is an m2 and char MNA 4522 is m6 (or. less

probablv,. m5)î m4 of 5. fhxi is represented by

OMNFl 20135.

4 hc Smoky Hollow Alcmber of chc Straight

Clifts Formation, lare Furonian in age (Eaton

1991). bas yieldcd nvo spalacothcriids (Cifelli

1990). The présent comparisons, which incliide

new marerials, indicaie that the holorype of

Sjmwctrodontoides ohgodaatfis Cifelli, 1990

(MNA 5789) is probably mh (not m7 as origi-

nally thought), and that the original referred spe¬

cimen (OMNH 20381) represcncs ni4.

Newly-refen'ed specimens include MNA 6047

and 6755, tenrarively identified as m2 and m4,

respecrively: and 29523, a m.mdibular fragment

with m2 The holocype uf che diminative

Spalacothcridtuni mekeatuit Cifelli, 1990 is most

probably m2 (not m4 as originally believed);

newiy-referred OMNFl 29524, MNA 6046, and

OMNFI 29526 arc probably ml, rn4, and cor-

roded m6, rcspectivcly.

UpT'ER MO la un

The upper molars are e\4dently much more fra¬

gile and subject to breakage during the screen-

washing process; the sample from the Cedar

Mountain Formation includes only 54 catalo-

gued upper molars, as opposed to more than

200 lowers. Of tfiese, 45 upper molars pnwed

assignable tu species Iml only 31 were formally

induded in hypodignis becausc ol their greater

completcness; statistical analysis was prccludcd

by insiiffkient sainplcs. With one exception (des-

cribed utuler ?Spvalacotheriidae, indet.), the

upper molars readily fell inro three categories <m

the basis of size, as had been cstablishcd for the

lower molars. Within species, thcrc is variation

chai is cJcaiiy duc co tooth locus. Trends known

for symmerrodonts represented by dentulous

jaw'S (c.g., Spalacoiheriam, /Jkinghi'ojhenum, see

Simpson 1928b; Clemens 1963; Hu et ai 1997),

together with molars (Fox 1985.) of a species

similar ro chose from- the Cedar Mountain

Formation, pnwided rhe liMsis (or esrahlishing

which tceib were more arucriorly placed in the

jaw, and which of thèse represented rhc first

molar. For the most abundani species, ir was

then possible to .sort molars iryo discrète mor-

phological categories, and to document progres-

178

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from thc mid-Cretaceous of Utah

sivc changes chn:>ugh rhe upper .sériés. \Ve reco-

gnize six morphological categories among upper

molars of ihis species, and therefore tentarively

regard the upper molar couni a*s six. This^ accords

with rhe fact chat, in die lowet dentition, where

we believe seven molars are présent, m7 is rediic-

ed and lias a prcvallid (for shearing against rhe

distal face of IV16) but no posrvallid surface. A

differcnrial molar counr berween upper and

lower jaws is not wholly unexpecred, as the

condition îs known to occur in thc primitive spa-

lacoihcnid Zhangheotherium (see Hu et ai

1997). Simpson (1928a) regarded rhe upper den¬

tition of Peralestes (whicli we include in

SpaLicotherittni) as having seven molars. CIcmens

(1963) later showed rhat only six were présent on

one specimen, at leasr; ir is possible that molar

count varied in this taxon.

Identification of upper molars belonging ro the

most abundant spccies then served as rhe basis

for identifying loci among isolatcd teetli referable

to thc Icss abundant raxa from the Cedar

Mountain Formation, and for morphologically

similar species from elsewherc. l*ox (1985) refer-

red two upper molars to thc Aquilan spalaco-

iheriid Symnietrodimloides atnadenm and indicar-

cd that onc probably represents a more posterior

locus than the other. We find this indeed to be

the case, ont compari.son.s suggesting that

UALW 16271 is Ml or 2 and UALVP 16272 Is

M2 or 3 of this species. No upper molars hâve

yet been described foi .symmetrodonts from

Southern Utah. but matcrials in hatid suggest

that thc following arc represented: Symme-

trodontoides foxi, M4 (MNA V4653); S. oti^o-

doutas. Ml' (OMNH 29525), M2 (MNA

V6048. OMNH 29040), and M6 (OMNH

29039); Spalucotheridium njcke7)nal, M5 (MNA

V6756). Other spaka>thcriids for which isolared

upper molars bave heen described are problem-

atic bccausc only single teeth (rather thaa sériés)

are available, and they arc so different from thc

taxa considered here as to bc non-comparable,

Several isolatcd upper molars have been referred

ro Spalacorberoidei bridwM. Of these, FM N H

PM 1235 is relaiivcly long mesiodisrally and pro-

bably represents an ançerior position, as indi-

cated by Patterson (1956, fig. 1). A casr of

FMNH PM 1 133, which is just a fragment of a

tootli, suggest-s that this may have been a more

posterior molar. ’l'hc upper molar sériés is Icnown

for Zhangbeotheriu7n\ bowever. ihe prcliminary

description and available illustration (which

shows the upper molars in an oblique orienta¬

tion, Hu et ai 1997, fig. 2) permit only cursory

comparisons with remaining spalacotheriid.s. The

remaining taxon po.ssibly referable to

Spalacorheriitlae and represented by an isolatcd

upper molar is Mkroderson Ltaroussli. from the

Early Cretaceous of Morocco. As recognized by

Sigogneau-Russell (1991b), thi.s tooth is so dissi-

milar to upper molars of Synimetnukntoides (and,

by implication, to remaining taxa considered

here) rhat wc cannot hazard a guess as to its posi-

lion in the jaw.

SpalacOLESTINAE n. subfam.

Tyve genus. — Spalacolestes. n. gen.

Incluüku f.;hNhR/\. — The type, and SpaUcotberoides

I^arterson, 1955, Symmetrodontoides Fox, 1976, and

Spahicotheridium Cifelli, 1990.

DisrumUTlON. — Crciaccous (Aptian-AIhian ihrough

carlv Canipanian), Norrh America.

DiAGNOSiS. — Distingiiished from primitive spaiaco-

theiiids (Spulacütheriuw, Zhatighcothcrium) in posses-

sing the following derivcd features: molars more

acutely angled; anteriur upper molars with sirong

panistylc; uppet molars wilb preparacri.sta lowci ilian

postparaciista and with distal sndar cusp presenr,

pro.ximal lo the metastyle. Prerygoid cresc and pier)--

goid fossa, where known {Spabicolestes. Spalaeo-

tberoides) exrend anrerodorsally from mandibular fora¬

men roward alvcolar rnargin of dcniai*)'.

CoMMENTS. — As shown by thc comparisons

below, there is goud reason rt) believe that rhe

Norrh American Creraceous spalacothcriids forrn

a monophylecic assemblage with respect co

remaining members of rhe family. We fdrmalize

this rclationship by placing the Nortb American

taxa in rheir own subfamily, named for rhe besr

known genus (described below). Wirhin rhe sub¬

family, poprlv known Spalacothero/des is thc

i.ildesr and appears to retain the gr&.ite.st number

of primitive features. The remaining généra ni

Spalacotheriidae {Spalacotheriurriy Zhangheo-

GEODIVERSITAS • 1999 * 21 {2)

179

Cifelli R. L. & Madsen S. K.

theriurn), not rrcaccd in detail hcrcin, are rele-

gated. by default, co Spalacotheriinae (Marsh

1887), n. rank, which current évidence suggests

may be paraphyletic (sec below).

Spalacolestes n. gen.

Type specihs. — Spalacolestes creUilablatta n. sp.

Included SPECIES. — The type, and 5. mconcinnus

n. sp.

ErYMOlA)GV. — Spulax (Cîreek), mole, and a com-

monly used prefix for gênera oF this Family; lestes

(Cîreek), robber. plunderer, and a commonly used suF-

fix for généra oFsmall and presumably stealthy, preda-

ccous mamnrals.

Distribl 1 ION. — Albian-Ccnomaniun, western

United Srate.s.

DlAGNOStS. — Dibers Froni Spalacothcrium in having

more acutcly anglcd trigonids on posicrior rnolars and

in having lowcr molar paraconid miich lower chan

mciaconid. DiOers from Symmetrofhntoides in having

proportionarcly nairowcr posterior lowcr molar.s witn

more obtusely-anglçd trigonids and lessvr Keiglit ddfe-

rendal bcrw'cen paraconid and meiacxniid; ml diflci's

from ihat t»l Sytnfuetroçlouîoieiçs in Iraving lower, more

coniad paraconid and lowcr puraçristid. Lower rnolars

diftei (rom ihosc tif SpalacotheyifUuiti in Ir.iving a more

pronounced heighr dilïerentia! between paraconid and

metaconid. Upper moUis simibr, whcre known, co

chose oF SynjmenoclûHtoides. excepe chat Ml-2 hâve a

more bulbous-based paraconc witb a gcntly ciirving

(noi tighlly arced or foîdcd) lingual face. Upper

rnolars dllFer From tho.se ot Spalaentheroides and prtmi*

cive taxa in réduction ol rhe stylocone, lack ot cusps

Bj and C, présence of ati extremcly low pa*paracrista

(anterior loci onJy), and pte.sence ot an enkrged distal

scylar cusp. UiFFers From tbe orherwîse similar

Spalacotheridjum in having deeper irigon ba.sins and,

on posterior iippcr rnolars, patasiylc rcduccd.

Spalacolestes cretulablatta n. sp.

(Figs 6-11)

Hoeotype. — OMNH 29600, right denrary with

m4-7.

Hw^DKiM. — ’l be holotype, and the following speci-

niais:

|aws: deinaty witb m4-S, OMNH 2”421; deiuaiv

with m6. OMNH 27557.

Lower rnolars: mt. OVINH 26424, 26425, 26697,

29608. 30621. 33044, 33220; m2. OMNH 27451,

27511, 27541,32947, 33045, 33054, 33226, 33898;

m.3, OMNH 26698. 2:^591, 2763J, 33047, 33217,

33851; m4, OMNH 26419, 26420, 26422, 26704,

26708, 2"471, 27484, 27630, 30627, .30628, 30631,

33055, 33225, 33901; iii5, OMNH 26423, 2669 5,

26703, 26706, 26707. 27462, 29603, 29606, 30619,

30620, 33037, 33042. 33043. 33049, 33050, 33222,

33905; m6. OMNH 27425, 27464. 27569. 29601,

29767., 30622. 30625, 33040, 33046. 33048, 33218,

33228;, m7, OMNH 27463.

Upper muhirs: MT OMNH 26426, 33233; M2,

OMNH 26686, 2%l I, 32897; M3, OMNH 2^512,

33060; M4, OMNH 2668vS, 26693. 3061 I, 30612,

3305T 33231; M5. OMNH 25796; M6, OMNH

26691.30614, 32949.

.ADDITION’AI Kl-H-RItEl) SPECIMENS. — Incomplète

upper molar.s. locus uncerrain: OMNH 25795,

26427, 26430. 27632. 33056. 33058. 33059, 33235,

33237,33906, 33907.

LoC'AIIIIES ANE) HORIZON. — OMNH localities

\^235, V239, V695, V794, and V868 (Fig. !); upper

part ot (ledar Mountain Formation; Albian-

Ctnomanian.

HlVA-iOEOGY. -— Cremla (Latin, dim. ot creîa). chalk;

blatta (Latin), tuckroach. Allusion is lo rhe

C^recaceons age and leniarkablc, roach-like abundance

of the species.

DIAGNOSI.S. — l'hc smaller oFrhe rwo species reFerred

10 tbe genus. DiFFcrs From rhe sliginly smaller

Symntetradonwkles oli^odontos in ch.iraciers noied For

gcncric diagnosis ami in having Icss slender, antero-

po.stcriorly compresscd lowcr molar para- and rnetaco-

nids. Oiffers tiom S. foxi and S. ctwademis in generic

charactefs and in beiitg much smallcT,

COMMEN r.S AND DESCRIPTION

Loiaer malar sériés

Spalûctflestes cretiilahlatta n. gen., n. sp. is the

jTiost abüudanc therian mammal oF the

Musseniuchii local fauna and is represented hy

94 specimens ot the lower molar sériés. OFrhese,

the holotype préserves the lasC tour rnolars in

place, and tsvo other dentnlous jaws bave teeth oF

known position in the posterior part oF the

sériés. Tsolared teeth nor rcterable to these loci

can be sorted invo three groups ba.sed on crown

height, relative heighr and position of the para-

conid,, curvaiurc of the lingual cingulum, and

oiber charactcrisiit-s. Standard ineasuremenis lor

the.se teeth (labié 1) also group iiuo distinct

clusters; as nored above, wc rherefore recognize

seven lower rnolars. The most morphologically

180

GEODIVERSITAS • 19^9 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Table 1 . — Descriptive statistics for lower molar measurements (mm) of Spalacolestes cretulablatta n. gen., n. sp. See Figure 2 for

measurement abbreviations and conventions.

ANW

Pad-med

Prd-med

Angle

Range

0.989-1.016

0.750-0.856

0.730-0.750

0.559-0.716

61.771-68.440

Mean

1.002

0.801

0.740

0.652

65.106

0.019

0.052

0.019

0.126

0.072

Range

0.496-0.529

0.694-0.752

0.431-0.444

0.415-0.480

45.600-50.309

Mean

0.515

0.721

0.437

0.448

48.163

0.033

0.037

0.015

0.073

0.049

Range

0.562-0.705

0.809-0.975

0.369-0.552

0.498-0.663

42.738-48.260

Mean

0.626

0.878

0.473

0.562

45.950

0.083

0.059

0.113

0.099

0.051

Range

0.470-0.675

0.736-0.963

0.359-0.472

0.496-0.598

29.714-40.046

Mean

0.601

0.868

0.413

0.598

35.068

0.106

0.075

0.087

0.086

0.094

Range

0.387-0.612

0.742-0.876

0.362-0.469

0.501-0.610

34.071-40.857

Mean

0.533

0.811

0.406

0.563

37.656

0.107

0.046

0.085

0.055

0.063

Range

0.381-0.555

0.651-0.826

0.288-0.424

0.402-0.630

31-920-37.643

Mean

0.480

0.749

0.347

0.522

34.239

0.101

0.072

0.117

0.134

0.059

Range

0.383-0.410

0.484-0.487

Mean

0.396

0.485

0.048

0.004

—

distinctive molars, identified as mU hâve a low,

anteriorly placed paraconid, somewhat lower

Crown in general, low crown width to length

ratio, and vvide irigonid angle (grcater than 60*^).

rhe second and ihird lower molars are progressi-

vely shorter, wider, and tailcr crowned, wich a

more acute trigonid angle ( làblc 1 ; Figs 4-5) and

bases of paraconid atid inctaconid more closcly

approximated; difletences in mcans for dimen¬

sions of ml and m3 are significant at the p =

0.05 level, whcrea.s that for the trigonid angle is

not (Table 2), These trends are continued

through the fourth molar; the différences in

width and length are not .significaiit between m3

and m4, whercas fhar (or the trigonid angle is:

m4 has a mtich more aciite angle (Tables I, 2),

The fitth lower molar Ls ncarly as tall as m4, but

tooth length and width dccrease past m4, and

the différences berween mcans arc highly signifi-

cant. Values for trigonid angle overlap considera-

GEODIVERSITAS • 1999 • 21 (2)

181

Cifelli R. L. & Madsen S. K.

Fig. 5. — Proportional différences (length, wldth. trigonld angle) according to toolh locus for lower molars of Spalacolestes cretuta-

blatte, n. gen.. n sp. Symbolsicircles, ml; squares^ m2: upward pointing triangles m3; diamonds. m4; downward pointing tri¬

angles. m5: hexagons. rnS (m7 is omitted because il lacks a melaconid and lhe irlgonkj angle cannot be calculated).

bly between m4-6 (see commcnts above), and no

consistent pattern is recognizablc. On posterior

molars, thcrc is a tcndenc)' For the paracristid to

bc slightly longer than tlie protocristid, whcreas

the reverse is rrtie For Syrunietro^/onroides. l he

sixth lower molar is lower crowned and sniallcr

than mS. The last lower molar, m7) is lower

crowned yet and is by lar lhe smallcst in che

sériés. The nietaconid on this toorh is lacking

From OMNH 29600 (the holotype) and from

one isolatcd, rclcrrcd specîmen. We consider it

unlikely rhat présence of rhe ciisp is variable

within the spccies, given the distinctivcness of

m7, although it is curions that so few molars

assignable lo this locus were rccovercd, in view of

rhe large samplcs of more mesial teeth. The lack

of a funcriona! posrvallid surface on m7 suggests

chat, as in Zhangheotberhiw (see Hii et ai 1997),

Spülacolesrcs atiulMima had one fewer molar in

che upper than low^r sériés. Changes in propor¬

tions and trigonid angle ihrtîiigh rhe molar sériés

are summarized in Figure 5, and a composite res-

torarion is shown in Figure 6C, 19.

Ail lower molars are douhic-rooccd. None of the

isolated reeth préserves both roots intact, but

comparison within this sample, together wich

preserved alveoli on the jaw fragments assigned

to Spalacolestes cretidablattay indicates that the

182

GEODIVERSITAS • 1999 • 21 (2)

Spalacothcriid symmetrodonts from the mid-Cretaceous of Utah

Table 2. — Two-sample / test (independent / test) comparing mean measurements for adjacent lower molars. Spalacolestes cretula-

blatta n. gen., n. sp. First and last molars omitted because of insufficienl data (IM for samples are given in Table 1).

Length

Width

Pad-med

Prd-med

Angle

m2 VS. m3

Différence between means

0.111

0.157

0.035

0.114

2.213

Pooled variance /

- 3.514

- 5.664

- 1.111

-3.228

1.371

Degrees of freedom

Probability

0.007

0.000

0.295

0.005

0.208

m3 VS. m4

Différence between means

0.024

0.011

0.060

0.035

10.882

Pooled variance t

0.911

0.437

2.935

-1.460

7.568

Degrees of freedom

Probability

0.080

0.061

0.009

0.016

0.000

m4 VS. m5

Différence between means

0.068

0.057

0.007

0.035

2.588

Pooled variance t

3.195

3.238

0.487

2.135

-2.041

Degrees of freedom

Probability

0.003

0.631

0.043

0.055

m5 VS. m6

Différence between means

0.053

0.062

0.058

0.041

3.418

Pooled variance /

2.685

3.899

3.554

1.887

3.097

Degrees of freedom

Probability

0.012

0.000

0.002

0.074

0.007

roots were subequal in size (excepc for m7, in

which rhc distal root is sinaller) and inc.siodLstally

compîressed, witli a charactcristic .subrcctangular

cross section tliat makes edentulous spalaco-

Icstinc inandibles easily recognized us such. l’be

cingulum is complété on ail lower molars and Is

cspecially scrong on tbe lingual side of ml

(Fig. 7A, B,)> where it shows almost no flexurc;

on subséquent molars, il dexes dorsally bctween

che bases of paraconid and meraconid. 1'lie cin*

gulura descends considerably as !t extcnds pasr

the inrerstirial régions ol ihc Looth, so tbal tbe

labial pari of the crown is miich bigher dian the

lingual sidc. Mcsiolingual and distolingual eus-

pilles are présent on the cingulum-, as ibey are in

other Nortb American Spalacotheriidae; lhese arc

variable but are gcnerally salieitï, projecting

somewhat mcsially and distal ly (as well as dorsal-

ly) from the cingulum. The paraconid and meta-

conid of ml (Fig. 7At B) bave conical, well-

separated bases wKen viewed lingually; the mera¬

conid is about tAvo-rhirds the beigfit ol rhe pro¬

coconid, whereas che paraconid is less. Lhaii

one-half the height of that cusp when the tooth

is viewed lingually. On m2 (Fig. 7C, D), the tri-

gonid angle is more acure, and the bases of para¬

conid and mctaconid are mtjrc closely

approximated because the former cusp is more

posteriorly placxd* The paraconid is idacivcly taJ-

ler lhan on ml, but lower rhan un succccding

molars. The meuconid of m2 is more siender,

with H less robust base, chan on ml. The lingual

cingulum dcvelops a pronouneed lingual flcxure

on m3 (l'ig. 7h, F), and this continues on suc-

ceediiig molars. Yhc labial face of ebe protoconId

on this and succceding reerh is less rounded and

more sharply folded than on inl-2; the meta-

conid i.s nearly as rail as the pmtoconid, with the

paraconid being only about half as tall as the

proroconid, viewed lingually. Of the two basal

cuspules, tbe discal fi.c„ calonid) lends lo be tbe

more prominenr and projecting; the succccding

tooth fus ivito ilic concavity of the distal cingu¬

lum lî/nned labial lo ihc distal cingular cusp (sec

Fox 1976). By m4 (Figs 70, FI, 8, 9), the bases

of paraconid and mctaconid arc appressed, wiih

a more slendor, anteroposteriorly compresscd

appearance than on anterior teeth, when viewed

GEODIVERSITAS • 1999 • 21 (2)

183

Cifelli R. L. & Madsen S. K.

Fig. 6. — Composite lower molar sériés in occlusal (A, C, E) and lingual (B, D, F) views; A, B. Spalacoîheridium noblei n. sp.;

C, D, Spalacolestes cretulabtatta n. gen., n. sp.; E, F, Spalacolestes inconcinnus n. gen., n. sp. Tooth sériés scaled to relative size.

184

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 7. — Scanning électron micrographs, lower molars of Spalacolestes cretulablatta n. gen., n. sp.; A. C, E, G, I. K, M, lingual

views; B, D, F, H, J, L. N, occlusal views: A, B, left m1 {OMNH 28424); C, D, right m2 (OMNH 33226); E, F, left m3 (OMNH 33851);

G, H, left m4 {OMNH 26422); I, J. right m5 (OMNH 30631); K, L, right m6 (OMNH 27557); M, N, left m7 (OMNH 27463). Jaw frag¬

ments and roots eliminated where needed to improve clarity. Scale bar: 1 mm.

GEODIVERSITAS • 1999 • 21 (2)

185

Cifelli R. L. & Madsen S. K.

lingually. Lovvcr molars 5 an J 6 (Figs 7I-L» 8) are

similar, but progressivcly smallcr, with crown

height dfcreasing aftcr an apparent maximum at

m4'5. The last lower molaj* (Figs 7M. N, 8) is

distinctive in its mueh smailcr sizc; although it is

two-roored like more anterior ceeth. che meta-

conid and protocristid arc lacking, and che distal

cingulum is expanded. The posteriormosr molar

(m7) SpaliU'Oihcnnm is idso quite sinall, but ir

retains a full complément of trigonid cu-sps (see

Clemcns 1963).

The available sériés oi lowcr molars of Spalnco-

lestes cretukihlntta enconipasses a widc variccy oi

wear stages. As wear progresses, ihe V-.shapcd

notches in paracristid and protocristid hecome

rounded and U-shaped. On mf, wear is heaviest

on the protocristid, which develops a facet that

dips distally. The wear tacers on paracristid and

protocristid are rarher oblique tu the occlusal

plane in early wear, progm.ssively hecoming more

parallel lo thar plane. In advanced wear stages

(e,g., OMNH 27569, m6), the crown forms a

continuons, concave, rriangular wear surface that

dips slighrly in a mcsial direction. Obliquely

oriented striations are présent on prevallid and

postvallid faces of worn molars; thèse arc more

pronouneed and recogni/able on m4-(?, whcre rhe

mesial and distal faces of rhe teeth are somewhat

more planar than on more anterior molars. whcre

lhey arc more convex, The rini of ihe cingukun

forms a sharp ridge in unworn teciii, c.spocially

mesially and distally. With wear, small intemitial

lacets de\'clop mesiall}' and distally, and the sharp

mesial and distal rims are bcveled off into rather

fiat, obliquely oriented lacets.

Mandible

Asidc from small fragments, tlie detuary of

Spalacolestt's trciuLibLum is known from n \'0 spé¬

cimens, OMNH 29600 and 27421 (Figs 8, 9).

OMNH 27421 préservés the horizontal ramus

ventral to the level of ml, the posterior alveolus

of iti2, the base of m3, m4-5 intact, and paired

alveoli for m6-7. Most of die ascending ramus

and angular région arc missing. OMNH 29600

(the holot)pe) includes che hori/onwl ramus pos-

terior to m3 and préservés m4-7 in place. The

crown of m4 is hroketi from its b:LSC and is rota-

ted and dispiaced; minor postmortem rotation

and displacemcnl of the other molars fias aiso

occurred. The posterior and inferiur mai'gins of

the angular région arc intact, excepr for the loss of

the condyle. The posterior margin is intact for a

short distance dorsal lo rhe position of rhe condy-

le: rhe coronoid process is broken obliquely and

irs fvill extent cannot bc determined.

The horizontal ramus (Figs 8, 9) bas a veiy gracile

appearauce compared to tfiat of SpciLlcothcrii^m.

The ventral margin of the liori/oniul ramus

appears somewhat bowed in latéral view, owing

to a slightiy greater depth bcneach m4-5 dian

antcriorly or posteriorly. In dorsal view (Figs 8B,

9B), the ramus is relativcly .setaight posterior to

the icvcl ot m3; anterior to rliat tooth position, it

ciirvcs mcdially. The ascending ramus arises

about a molars Icngth posterior to ihe position

ol m7 and angles dorsally at about 45° with res¬

pect to the alveular margin ol the horizontal

ramus. The posterior pan of the jaw is aiso

remarkably gracile in appeârance, the bone beiiig

very rhin in comparison to the far more rohust

(and larger) mandible of Spalacotberium.

Froporcionately, the ascending ramus is much

longer anceroposteriorly ihan it is in Spnlaco-

thenum.

On the latéral sidc u( ihe jaw (Fig.s SC., 9C). the

masseccric foss.i is well marked and, owing to the

fotm of the angular région (sec helow) and latéral

flexurc of the anterior margin of the ascending

ramus, lias the appearunce ol being quite deep.

T here is nn labial mandibiiLir toramen présentât

the apex of the masseteric Ibssa, as thcrc is in a

number of otiier primitive mammals (c.g.,

Dasivzeveg îk Kidan-Jaworowska 1984; Marshall

&: Kiclan-Jaworowska 1992; Cifelli et al. 1998),

although there is an extreincly small nutritive

foramen in one spécimen (OMNH 29600) ai a

point somewhat dorsal tu the apex of the masse¬

teric fossa.

Tfie atuerioi margins ol the ascending ramus and

nias.scfcric lossa l1cx strongly in a lareral direction

as they rlse above rhe alveolat margin ol flic jaw.

.Sirnilarlv. the interior margin of the dciuary in

the angular région ha.s a salieiit latéral dellcction.

1 bis .strongly dcflccted angular région and latéral

flcxure ol rhe anterior margin ot rhe ascending

ramus, rogeihcr with sunilady strong featums pn

the lingual side of the mandible, give the dorsal

186

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 8. — Dentary of Spalacolestes creîulablaîta n. gen., n. sp. holotype (OMNH 29600), right dentary with m4-7, in lingual (A):

occlusal (B), and labial (C) views. Scale bar: 2 mm.

GEODIVERSITAS • 1999

PJH

Ir i

f' Æ- i

h J

Fig. 9. — Dentary of Spalacolestes cretulablatta n. gen., n. sp. (OMNH 27421), left deiitary with nr»4'5, in lingual (A), occlusal (B),

and labial (C) views. Scale bar, 2 mm.

(Figs 8B, 9B) and posterior views of the man-

dible in Spalacolestes cretulabLitta an appcarance

thac is unique, so far as we are aware, among

Mesozoic mammals.

On the lingual side ot the jaw (Figs 8A, 9A), a

strong crest, which we interpret as being for die

insertion of the m. pterygoideus mcdialis, des¬

cends posreroinfcriorly froni the junction of the

horizontal and ascending rami, beginning jusc

below the alveolar margin. This crest screngthens

to a shelf as it passes just inferior to the mandi-

bular foramen, which has double openings for

the mandibular canal on one spccimen (Fig. HB).

Fhe mandibular fonurten is comparattvely large

and faces posrerolabially, owing to the great deve¬

lopment of rhe pterygoid crest beiicath, the anre-

rior and posterior margins of the mandibular

foramen are dcvclopcd as lips that projccc lin-

gually as they descend to the pterygoid crest. A

short distance posterior to the mandibular fora¬

men, the pterygoid crest is developed as a sallent

process that thickens into a robtist tip. A deep

188

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Crctaceous of Utah

Table 3. — Measurements (mm) and descriptive statistics for upper molars of Spalacolestes cretulablatta n. gen., n. sp. See

Figure 2 for measurement définitions.

L(N)

L (Range)

1.164-1.225

0.747

0.669-0.846

0.684

0.546-0.552

L (Mean)

1.195

0.747

0.745

0.684

0.549

L(CV)

0.036

1.000

0-103

1.000

0.006

ANW(N)

ANW (Range)

1.084

0-971-0.988

0.949

0.942-1.113

0.745

0.614-0.682

ANW (Mean)

1.084

0.979

0.949

1.034

0.745

0.656

ANW (CV)

1.000

0.012

1.000

0.065

1.000

0.056

POW (N)

POW (Range)

1.152-1.168

1.028-1.038

0.871-1.090

0.962

0.757-0.888

POW (Mean)

1.160

1.033

0.994

0.962

0.813

POW (CV)

—

0.007

0-107

1.000

0.083

pocket is enclosed berween rhc pterygoid process

(lingually) and the body of thc dentary (labially).

Where présent, the pterygoid crcst commonly

exrends posteiiorly to thc condylar région in

Mcsozoic mammals (e.g., Triconodontidac,

Tinodontidae, Dryolcsiidae, see Simpson 1928a,

1929): in Spalacolestes cretulablatta, ii terminâtes

at thc posterior margin of thc proccss, ’J'he ptery¬

goid fossa is very broadly developed anterior to

the mandibuUf foramen^ Mnd in this respect dif-

fers from iliat oïZhangheotheriunt.

No meckelian groove or postdentaiy trough are

apparent, nor are scars for the coronoid or other

postdentury boues, as commonly seen in primi¬

tive mammals (Kerniack Ôe Mussett 1958;

Kcrmack et ai. 1968; Dashzeveg 6c Kielan-

Jaworowska 1984; Kielan-Jaworowska &

Dashzeveg 1989; Krebs 1991; Lillcgravcn

Krusat 1991; Nessov et al. 1994). A vestigial

trace of the meckelian groove is présent vinterior-

ly on the dentary Zhaughecftheriuw, but a

postdentary trough is lacicing. The condyle is noi

preserved, but its position is shown by a slight

rhickening of bone just ventral to the- preserved

posterior margin of the ascending ramus in

OMNf] 29600 (Pig. 8A, C). The condyle would

have been situated ac, or sHghtly below, the

alveolar margin of the horizontal ramus, lower

than in Spalacotherium (e.g., BM 47750).

Upper molor sériés

The upper molars (Figs 10, 11) are two-rooted.

They lack a lingual cingulum (although a faint

basal sweiling is variably présent), as seen in

Symmetrodonloides canadensis (sec Fox 1985),

and cusps on che pre- and postparacrista, as seen

on upper molars Spalacotherïiim (see Simpson

1928a), Spalaeothtroides (see Patterson 1956),

and Zhangheotbetiurn (sec Hu et al. 1997).

Acuieness, iransvcrse width. and height of thc

prcparacrista relative to thc posrparacrisia incrctse

trom Mi to M4. wluch is almost completely

symmetrical. The parastylar Jobe and distal stylar

cusp art promineiii on anterior molars (as they

are in Symmetrodotttoides), and decrcase through-

out the sériés. By contrasta che parastylar lobe is

more strongly developed on posterior upper

molars of Spalacotherium (see, e.g., Clcmcns

1963). The paracone is mrjst distally rccumbcnt

on Ml, decreasing in recumbency through M4,

where it is symmetrical arid crect. Past M4,

molars arc progressively smaller fiable 3), narrow-

er labiolingually, and hâve a more posteriorly

placed paracone, A contposite restoratjon i.s

shown in Figure 1 I; many of thc trends évident

in the restored séries are also seen in ihc compo¬

site of Ktiehaeotherium (sec Mills 1984).

The disrolabial parc of Ml (Fig. 10A> B) is not

preserved on available spccimcns. ’Fhe paracone

is strongly recumbent distally; iis mesial face is

rounded, lacking rhe "pinched" appearancc, wich

relatively straighi shearing surface, of more distal

tccth. A weak ridge, which develops heavy wenr

on its occliisal surface, descends almost vcrtically

from the apex of the paracone down its meso-

labial surface; this is équivalent to thc prepara-

GEODIVERSITAS • 1999 • 21 (2)

189

Cifelli R. L. & Madsen S. K.

Fig. 10. — Scanning électron micrographs, upper molars of Spalacolestes cretulablatta n. gen., n. sp.; A, C. E, G, I, K. occlusal

views; B, D. F. H, J. L. meslal views; A. B, left Ml (OMNH 26426); C, D, left M2 (OMNH 26686); E. F. left M3 {OMNH 33060);

G, H, left M4 (OMNH 30611); I, J. left M5 (OMNH 25796); K, L, right M6 (OMNH 26691). Jaw fragments and roots eliminated where

needed to improve clarity. Scale bar: 1 mm.

190

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 11. — Composite upper molar sériés in occlusal view. A, Spalacotheridium noblei n. sp.: B, Spalacolestes cretulablatta, n. gen.,

n. sp.; C, Spalacolestes inconcinnus, n. gen., n. sp. Sériés scaled îo relative size.

crista, but a crest as such is noc dcveloped until

rhis surface approaches the parastyle, where its

presence is only faindy suggested. The parastyle

is very low, being placed near ihc base of the

Crown, and is developed as a promincnc, mesially

projecting lobe» A small accessory shclf descends

lingually from the parasrj'lar lobe ncar the labial

terminus ol the preparacrista, terminating ar che

base of the crowai; there is no other htnt of a cin'

guluni, alrhough the enamel is variably swollen

on the lingual base ofthe tooth.

M2 (Hg. lOC, D) is generally similar to thar of

Symmetrodontoides. although the parastyle and

meta.style are Icss dcveloped. It differs from Ml

in havinga flattened mesial (prevallum) shearing

surface, more strongly dcveloped preparacrista,

more acute angulation, less reciimbcticy of the

paracone, and less proniinenc, bulbous parasC)'lc.

1 he postparacrisca descends ar a sreep angle from

the apex of the paracone, forming a V-shaped

norch near the base of rhat cusp. The crest ter¬

minâtes near the distolabial corner of the tooth,

not quite reaching the small metastylar cusp.

From the merastyle, a faint crest descends lin¬

gually for a short distance along the disral ftee of

the tooth, terminating near the base of rhe tooth.

This crest (perhaps a remnant of a cinguliim)

evidently formed the margin of the occlusal sur¬

face of ihc tooth, perhaps serving as a guide for

rhe corresponding IcAvcr molar shearing surfhcc:

ir is oricneed at the same angle as wear striations

lücated higher and more lingually on the same

surface of che molar (c.g., OIVINH 32897:

Fig. lÜC, D). Mesial to atul separate from the

metastylar cusp is a prominent, mesiodistally

elongate, trenchant srylar cusp, as seen in

Symmetrodontoides. This crestlike cusp extends

mesially to rhe ectoflexus, whcrc ic mccts a lower

crest de.scending distally from che région of rhe

stylocone. 'fhe lacrer cusp is noc preserved on

available specimens but, if présent, it was small.

None of the M3s in the sample is complété, but

GEODIVERSITAS • 1999 • 21 (2)

191

Cifelli R. L. & Madsen S. K.

available specîmens show a continuation of

trends estahlished in ihe uppcr molar sériés: die

trigon is more acute and dccply basincd; die

paracone has only slight recunibency; clic prc-

paracrisia is reladvcly liighcr; and rhe stylar cusp

is smaller. One broken specimen (OMNII

33060; Fig. I0E> F) préserves rhe paraconal

crests in prisdne condition; cu.spules arc lacking

froiii rhese.

M4 (Fig. |0G, H) is rhe most traii.svcrsclv deve-

loped, acLiie-angled, nearly syinmeirical cooth ot

rhe sériés; in SpaLneotberium (including

Peralestes), ir is M3 chat appears to bc most near-

ly symmctrical (Butler 1939). The paracone is

mesiodistally compresscd, wich a distinct lingual

fold, and is not recuinbenr. AJI specimens are

almosi pcrfeccly symmetricah esxepr for minor

différences in chc parasryhir and metastylar

régions. The pre- and postparacrisrae are equal in

height, enclüsing a radier deep trigon basin, with

Hat, .strap-like faccts (as dcscribed for Symme-

trodontoideSy sec Fox 1976, 1985) on their occlu-

sal surfaces. I hc labial surface of the tooth bulge.s

adjacent to dic labial terminus of the preparacris-

ta, suggesting the présence ot a small stylocone

(obliteraicd by wcar on availabic specimens).

Mesial to this, the parastyJe is nuich reduced,

forming an incoiispicuous knob at the mcsiola-

bial corner of the tooth. Fhe mera.style is similar-

ly developed; just mesial to it, ihe niesiodistally

elongate stylar cusp is [neseni along the margin

of the stylar shelL This sr>'lar cu.sp is variable in

developnieiu, being largcsc in die figured spéci¬

men (OMNH 30611; Fig. lOG, H), but is much

reduced in conipanson to more anterior niolars,

l'hc cusp descends niesially as a crest rimming

the stylar shelf and enclositig the trigon basin

labially in the région of chc cctoficxus, where

there is variably (C)MNH 26693, 30611) a sniall

cuspule présent.

M5 (Fig. 101, J) is |ess transverse and forms a

more obtuse angle than M4. It i.s sonicwhat

lower crowned as well, although rhe prC' and

postparacristac arc high relative co the apex of

the paracone, and enclose a deep trigon basin.

The paracone is more distally placed than on M4

or preceding molar.s, recalling the condition in

tribosphenlc dicrians, where the protocoiie is

more distally placed on distal molars. The trend

in die molar sériés roward réduction ol tlie para-

st}4e is complété: no trace of it rcmalns. I hcre is

a faint trace of die metasiyle ac die distolabial

corner of rhe tooth. Fhe distal margin ol dic sty-

lar shelf is fonned bv the stylar cusp, which has

rsvinried apices in the single complète specimen

(OMN'H 25796; Fig. lOJ, J). l'iiis specimen is

virtualiy unworn, and shows (again) the lack of

acces.sory cusps on the pre- and postparacrista. It

also shows thaï the stylocone, worn away in most

other specimens availabic, is prc.Sent and i.s deve-

loped as a trcncliant, mesially placed counterpart

to rhe distal sr)'lar cusp.

M6 (Fig. lOK, L) is smaller and less transverse

ihan M5. Parastylc and metastyle are lacking,

and the labial part ot the tooth is insread occu-

pied by ilie stylocone and the distal stylar cusp,

the lattcr not quicc extending to the distolabial

corner ol the tooth. The paracone is more pos-

ceriorlv placed than on M5; the distal face of the

tooth is distinctive in heiiig ciirvcd, with a

rounded distolabial corner thac differs from the

more angular appearance of preceding molars.

Spalacolestes inconcinnus n. sp.

(Figs 6, 11-13)

Hoi.otvte. — Right m4, OMNH 33903.

HvrooiGM. — The holotvpe, and ml. OMNH

33039; m3. 33897; M2, 33034: M4, 3391 1.

Log-mity an'd HOKi/oN. — OMNH localin' V868;

uppcr part of Cedar Mountain iHirmarion; Albian-

Cenomanian.

FiVMOKït.V- — U-atin), awkward, coarse,

în rcferencc to die appearance of tite tcedi wlicn com-

pared to dic daiiiiy, élégant rnorpliology gcncrally

characicri/ing smaller .species of Spalacoihcrüdac.

DlAGNX)Sfs. — J ht* larger of tlic iwo .spccles referred

to the genus; lovvvr tin)lar ciiiguluin better developed

niesiolaliially on tnl lhaii in Spalacolestes crftulublatta^

iVoin which ii also differ.s in liavitig the trigon Ixisin

incompletely enclosed ar the cctotlcxiis ol posrerior

uppcr molars. Larger than Synimetrodotito'tdes frxr,

approximately similar in si/.c to S. catiadcmis^ from

which il dirfer.s in having pioportionately narrower

lower molars and other genenc tharacterisiics.

DesCRIV I ION AND t.'OMMENTS

Teeth of S. inconcinnus n. sp. resemble those of

192

GEODIVERSITAS ■ 1999 • 21 (2)

Spalacotheriid symmetrodoncs from the mid-C'recaceous of Utah

Table 4. — Measurements (mm) of Spalacolestes inconcinnus n. gen., n. sp.; see Figure 2 for measurement abbreviations and

conventions.

Tooth

ANW

POW

Pad-med

Prd-med

Angle

1.248

0.962

1.270

0.667

0.739

43.097

1.050

1.314

0.660

0.877

39.444

1.861

1.356

1.601

1.033

1.608

1.564

—

S. cretulûblaïuî in most charactcristics» difFering

chiefly in thcir much grc.iter size (Fig. 4) and in

features probably rclaccd co size (such as ciisp

robusticity). S. inconcinnus is much less common

than cither 5, cretuinblatta or Spabjcotheridiurn

noblei n. sp., bcing represented by only fivc

molars, two from the upper dentition and three

from the lower.

The lower molars arc so similar to those of 5. cre-

tulnblnttii that only a few commerits arc warran-

ted. l'he cinguliim is more strongly dcvelopcd

than in S. cretulablatto, particularly on ml

(Fig, !2A, B). In addition, it appears that, in

.S* iticnncintais^ ihe paraconid is lower relative to

the metaconid at corresponding tooth positions.

Variability cannot bc assesscd with the sample in

hand, however, and this po.ssiblc diftcrcncc has

accordingly been omirted from the diagnosis.

Rg. 12. — Scanning électron micrographs, lower molars of Spalacolestes inconcinnus n. gen., n. sp.; A, C, E, lingual views; B, D, F,

occlusal views: A, B. right ml (OMNH 33039); C. D, left m3 (OMNH 33897); E, F, right m4 (holotype, OMNH 33903). Scale bar:

1 mm.

GEODIVERSITAS • 1999 • 21 (2)

193

Cifclli R. L. & Madscn S. K.

Fig. 13. — Scanning électron micrographs, upper molars of Spalacofestes inconcinnus n. gen.. n. sp.; A, C. occlusal views; B, D,

mesia! views; A. B. left M2 (OMNH 33034); C. D. right M4 (OMNH 33911 ). Scale bar; 1 mm.

The only upper molar of 5. inconcinnus in which more bulbous, with a more pronounced labial

the roots can be seen is OMNH 33034 (M2; bulge, than in S. cretulablatta\ a vveak cresi, bare-

Fig. 13A, B), where cwo are présent. A faint swel- ly hinted at in the latter species, extends lingually

ling on the postparacrista, aboiir two-thirds of into ihe trigon basin Irom the base of the stylar

the distance front paracone to mctasryle, suggests cusp. The présence of a small stylocone can be

the possible présence of a ctisp on rhis crest, but contlrmed on M2 of 5. inconcinnus. On M4

breakage in this région prccliides judgmenl on (Fig. 13C, D), ihc para.slylar région and distal

this point. The crest descending lingually front stylar cusp arc more bulbous than in S. cretiüa-

the metastyle on M2 is stronger but shorter than blanUy so that the ectoflexus is deeper, although

it is in S. cretulahLttta. The distal stylar cusp is the distal stylar cusp of 5. cretulablatta variably

194

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Crciaceous of Utah

projecrs farther labially than in thc single known

M4 of S. inconcinnus. Howcver, thc crcsts des-

cending to the ectoflcxus from the styloconc and

discal stylar cusp arc very weak, so that the trigon

basin is not cnclosed labially, as iï is in S. cretula-

hLittü. OMNH 33911 includes ihe M4 embed-

ded in a hagmcnr ol the maxilla. The labial

margin of the toorh is oricnted at a high angle

with respect to thc latéral sidc of the maxilla (the

parastyle is near tlic latéral margin ot thc maxilla,

whercas the distolabial corner of thc tooth is

some distance from ic), suggesiing chat the ros-

trum flared latcraJly in this cegion (l-îg. 13C).

Genus Spal^icotheridium Cifclli, 1990

Type SPECIES. — spalacotheridium mekennai Cifelli,

1990.

Inci.UDED SPECIES. — The type, and

Spalûcotheridiîim noblei n. sp.

Distribution. — Albian-Ccnoniani.in throiïgh

’l'uronian, Utah.

RrAaSED DIAGNOSIS. — Spalacotheriids differing Irom

other members of the family in rheir small size (maxi¬

mum length and widrh niea.suremeius ol molars gene-

rally Icss lhan 0-75 mm), l.owcr molars differ from

those Spalaauhcrium in being more nearly symme-

trical ana acutely an^led; from Spalavothcroides in

having a complcie Tibial cingiilum; and from

Synmutrodonioiden and Spalacolenes in being lower

crowned, with paraconid and metaconid suhequal in

development and uf approximately équivalent hciglit,

only .slighrly lower than the protoconid, and in having

posterior molars that are proportionately narrower

and hâve mare obtuse rrigoniu angles. Upper molars

distinct from Spiihnorhemda in ihc piescncc of a lar-

ger distally pluted srylar cusp and more promineni

parastylar hook (anrerior loci), the kck of cusps Bj

and C, and thc extremely low placement of the prepa-

racri.sta (anterior loci). Upper molars dilfer Irom those

of Sthdacolcstes in having a .shallower trigon basin, and

in tne présence of a prominent parastyle on M6.

Comme,NES

This genus was originally bascd on a single inolar

of the type and then only specie.s (Cdelli 1990).

The recovery of anorher specics, repnesented by a

much more extensive sample from the Cedar

Mountain Formation, upholds thc morpho-

logical distinctiveness of thèse tiny symme¬

trodonts. Lower molars of Spalacotheridium are

generally similar to Spalacotheroides, except for

the described lack of a labial cingulum in the lai-

ter, but available materials Spalacotheroides do

not permit comparison betwcca thc two. in

concrast, thc upper molars of the two taxa arc

quiie different. By comparison with Spalaco-

therium, Spalacotheridium appears co be primitive

wirh respect to Syfnmetrodiniîoides and Spulacu-

lestts \y\ the features cited in the diagnosis.

Spalacotheridiuin noblei n. sp.

(Figs 6, IL 14-16)

Hourm'E. — OMNH 25828, left m4.

HypoimEiM. — rhe hointype, ond thc tollowing isoJa-

red teerh:

Lower molars: ml, OMNI! 25609. 3.5038, 33205,

33221; m2, OMNfî 30623. .53219; m3, OMNH

27261, 29605, 30630. 32948. 33041.33229, 33900;

m4, OMNH 25794, 26421, 27424, 27441, 27593,

29766. 30626, 30629, 32946. 33215. 33902;

27258. 27629. 29607, 29653, 33052, 33053, 33224,

33899; m6, 29602.

Upper molars; M), OMNH 26429; M2, OMNH

33061; M3, OMNI 1 .50618, 3.5895; M4, OMNH

26689. 26692, 30617. 33232; M5, OMNH 27595.

3.3912; M6, OMNH 27'i61.

ADDITIONAE REEERRED SPECIMENS, — Incomplète

upper molars, locus uneertain: OMNH 26687,

33236.

Dh.alitie.s and HCmiZÜN. — OMNH locaiitics

V235. V239, V240> V695. V696, V80L and V868;

upper part of Cedar Mountain Formation; Albian-

Ceiioinanian.

F.i vMOiûtfV. — For the .Samuel Roberts Noble

Foundation oFArdmore, Oklahoraa, in récognition of

its support for the Okiahoma Muséum of Natural

Historv'.

Diagnosis. — Differs from the most similar species,

S. mekennai, in having smaller (espccially in Icngth)

lower molars, wirh more acurc trigonid angle on ni4.

COMMKNT.S AND DESCRIPTION

S. noblei is rnrher similar to S. mekennai and is

distingnishable bccausc the sample of lower

molars is sufficient to show thaï the few known

spccimens of the latter fall oiitside thc range of

size variation in 5. noblei. The holotype of

5. mekennai (MNA 5792), idcntified as m2, has

GEODIVERSITAS • 1999 • 21 (2)

195

Cifelli R. L. & Madsen S. K.

Table 5. — Descriptive statistics for lower molar measurements (rnm) of Spalacotheridium noblei n. sp. See Figure 2 for measure-

ment abbreviations and conventions.

ANW

Pad-med

Prd-med

Angle

Range

0.888

0.692-0.722

0.532-0.626

Mean

0.888

0.710

0.579

1.000

0.018

0.115

Range

0.447-0.482

0.475-0.601

0.359-0.398

0.396-0.441

49.781-54.643

Mean

0.465

0.538

0.379

0.418

52.212

0.053

0.166

0.073

0.076

0.066

Range

0.421-0.539

0.633-0.795

0.340-0.411

0.416-0.511

42.789-47.581

Mean

0.490

0.711

0.393

0.460

45.490

0.087

0.090

0.076

0.072

0.042

Range

0.411-0.559

0.559-0.745

0.318-0.389

0.402-0.562

33.018-41.155

Mean

0.485

0.678

0.358

0.490

38.105

0.080

0.073

0.064

0.093

0.075

Range

0.409-0.465

0.608-0.720

0.348-0.360

0.424-0.492

37.875-39.004

Mean

0.446

0.671

0.354

0.452

38.439

0.043

0.067

0.024

0.055

0.021

0.352

0.537

0.270

0.400

36.642

length and width proportions siniilar to those oF

m3 in S. Noblei, which is considcrably larger, but

the trigonid angle of MNA 5792 is much greater.

When coniparcd to m2 of 5'. nohleU on the other

hand, MNA 5792 differs grcatly in its propor¬

tions, falling near the maximum for width and

the minimum for length (Tables 5, 6), Ail molars

of S. noblei arc sliorter than m4 of S. mckennaly

which has a relatively obtuse trigonid angle, des¬

pire its tooth position.

The lower molars of Spabtanheridium noblei dif-

fer from chose of Spolacolestcs and Symrnelro-

dontoides in being proportionateJy lower crown-

ed, wirh somewhat more obtuse trigonid angles at

corresponding tooth positions. The most ncarly

complété ml is ÜMNH 25609, which lacks only

the tip of the metaconid and parts of the cingu-

lum adjacent to chat cusp (fig. l4A. B). The

paraconid appears to be relatively taller, with a

broader, more robust base than is the case in

Syinmetrodontoides or Spalacolestes^ although it is

Table 6. — Upper molar measurements (mm) of Spalaco-

ttieridium noblei n. sp. See Rgure 2 for définition of measure¬

ments. Where more lhan one spectmen was measureable a

range is given: sample size for each appears in parenthèses.

Tooth

ANW

POW

1.175

—

0.896

0.646

0.743

0.917-0.957 (2)

0.537-0.551 (3)

0.628-0.827 (2)

0.754-0.838 (3)

0.649

0.795

0.746-0.838 (2)

0.435

0.548

0.441

196

GEODIVERSITAS • 1999 « 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 14. — Scanning électron micrographs. anterior lower molars of Spalacotheridium noblei r\. sp.; A. C, E. lingual views: B. D. F.

occlusal views; A, B. left ml (OMNH 28429); C. D, right m2 (OMNH 33219); E. F. left m4 (holotype. OMNH 25828). Scale bar:

1 mm.

still lower than the metaconid. As in ail other

North American Spalacotheriidac, ml is morpho-

logically disiinctivc l>y viriuc of the anicrior pla¬

cement of ihc paraconid. Lengch to width

proporcion.s appcar lo change ihrougli the molars

sériés about as in Spalacolestes crctuLiblaïUi,

known by a much larger santpic; chc teeth arc

lower crowned clian in Symmetrodontoides or

S[fnLu'oleste<. The second niolar (Pig. l4C, D) is

considerably shoncr than the first; absolutc widih

increascs rhrough m4, wirh m5-6 being sequen-

tially shorrer and narrower than m4 (Table 5).

On m2 and succeeding ceeth, the paraconid is

subequal ro the metaconid and both citsps arc tal-

1er relative to the [irotoconid than in SpaLicolestes

or Symmetradontoides. blowcvcr, the paracristid

dips slightly lower in Its médian notch than does

the protocri-stid. I he cingulurn Is complète and,

as in Spalacolestes and Symmetrodontoides, bears

prominent cuspules at the mesio- and disto-

lingLial corners of the tooih. Also as in thosc taxa,

the Crown is much taller labially than lingually,

and as a resuit, dic cingulurn descends noriceably

as it proceeds labially from the inrerstitial régions

of the rooth. On m4 (pig, ]4E, F), the metaconid

is somcwhac more lingtially placcd than the para¬

conid, so ihat the protocristid is slightly longer

chan the paracristid — a condition reminiscent of

what is seen in Syînmctrvdontoidcs^ although riot

so extreme, and the posterior molars never achie-

ve rbe remarkablc transverse expansion seen in

rhat genus. The dental formula cannot he establi-

shed with ceriainty. Identification of OMNH

29602 (Fig. 15C, D) as m6, however, scems

probable bccausc of its .small si/.c. low crown

height, and proportions, fliis specimen appears

to hâve an interstitial weai facet on the distal cin-

gulum, süggcsting the présence of a seventh

molar; we tentativcly regard the lower sériés to

include seven molars.

GEODIVERSITAS • 1999 • 21 (2)

197

Cifelli R. L. &c Madsen S. K.

Fig. 15- — Scanning électron micrographs. posterior lower

molars o! Spalacothêridium noblei n. sp.: A, C. lingual views; B.

D. occlusal views; A. B. left rn5 (OMNH 27629); C. D. left m6

{OMNH 29602). Scale bar. 1 mm.

Wear on molar cu.sps and crcst.s* prcvallid and

posrvallid .surfaces, and cingula is similar ro thaï

seen in SymmcfrodojitoùJes and Spalacolestes (sec

Fox 1976: Cifelli & Madsen 1986; and descrip¬

tion above, see aiso Croinpton tt ai 1994 for

discussion of the relatioaship betwecn apical

wear, shearing surfaces, and enamel microstruc-

ture). Strap-like faccts dcvelop along the dorsal

surfaces of paracrisiid (whcrc it develops earliest

and most strongly: c.g., OMNH 27424) and

prorocristid, joining ar rhe protoconid; as wear

proceeds, these gradually form a triangular,

concave facet with an emarginaced base (cotres-

ponding ro the notch betwecn the bases of para-

conid and metaconid). An unusual variant is

OMNH 33899i a heavÜy worn m5 in which

wear is sironger on the labial than lingual side of

the tooth, with the resuit that the paraconid and

metaconid are taller than the protoconid.

Upper molars (Figs 11> 16) are generally similar

to chose Spitlacolestes vrvtuLxbLitta and will be

dcscribcd only whcrc chcy dilTcr or providc addi-

tional information. Ml (Fig. I6A. B) i.s slightly

worn, but shows that the styloconc was lirtie

dcveloped or ah.sent. The parascylar shelh which

descends lingually from rhe mesolabial corner of

the tooth, bas a more planar surfnce th;in in

Spalacokstes cretulablatta. This may bc rclaced to

differing wear on the available spccimcns: the

parastylar shelf beats a wear ficct that is conti-

guoLis with the prcvallum shearing surlacc, sug-

gesting thar the shcll may hâve sen^ed as a guide

for the occluding lower molar. No complété or

lightly worn spécimens of M2'3 (Fig. I6C) are

available; as far as can be deierinined, they show

the progressive narrowing of ihc paraconc and

réduction of paiastyle and metascylc seen in

5. cmukblam (Fig. 1 i ); the metâ.srylc appears to

be less developed on corresponding teeth than in

that .species. The irigon hasin is not nearly as

deep as in 5. cic'lukbldtta or Syfnntetrüdiy?noides\

this is particularly noiiccableon M4-5 (Fig. I6F-

1), in which ihcrc is littic relief on spécimens that

are only lightly worn (c.g.. OMNH 26692,

27595). As with Spalacolestes and ^yntme-

trodomoi4^s^ M4 is the most transverscly deve¬

loped of the upper molars, and is almost perfect-

ly symmetrical. M6 (Fig. 16J) is impressive by

virtue of its minu.scule .size, probably being -smal-

1er relative to MS than in SpaLtcolestes. Il is simi¬

lar in having a ciirved distal surface and n.mnded

metastylar région, but differs in being less trans¬

verse and in having a more prominent, projec-

ting parastyle.

?Spalacotheriidac gen. & sp. indet.

(Hg. 17)

SiM-CIMKNS. — Anterior lower molar, probably m2,

OMNH 33896 (OMNH localiry V868); posterior

upper molar, perhaps M6, OMNH 29612 (OMNH

localiry V695).

COMMENTS .AND DESCRIPTION

Two specimens (one upper molar and one lower

molar) from the upper Cedar Mountain

Formation cannot bc referred to any of the three

species descrîbed herein, and rhus document the

présence of at least one more species of non-

198

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 16. — Scanning électron micrographs, upper molars of Spalacotheridium noblein. sp.; A, C, D, F, H, J, occlusal views; B, E, G,

I. mesial views; A. B, right Ml (OMNH 26429): C. right M2 (OMNH 33061): D, E, right M3 (OMNH 33895): F, G. lefî M4 (OMNH

26692): H. I, left M5 fOMNH 27595): J, left M6 (OMNH 27461 . Scale bar: 1 mm.

tribosphenic Theria in the Mussencuchit local

fauna. If locus is corrcctly interpreted, the teeth

appear to corne from animais of rarhcr differenr

size; henccv rhey cannot be referred to rbe same

species bascd on présent knowledge.

The lower molar (Fig. 17A, B), tcntatively iden-

tificd as ml. is intccmediatc berween ml and m2

oF Spalacoiestes or Spalavatheridium in terms of

length-width proportions and general appearan-

GEODIVERSITAS ■ 1999 • 21 (2)

ce: the paraconid is somewhac displaced anterior-

ly and in this respect is reminiscent of a spalaco-

rhcrîid ml (although ir is not nearly so anteriorly

placed as in ml of ail taxa known from the

North American Crciaccous), yet lliis cusp is

better dcvcloped, the crown is tallcr, and the rri-

gonid angle more acute than is gcnerally seen on

that tooth. The.se features suggest that it i.s m2,

implying that the specimen may represent a

199 I

Cifelli R. L. & Madsen S. K.

mesiodisial brcadth and low heighi of thc para-

cont relative co rhe rest of che crown> and pos¬

sible projecting parastybu région (ihc looth is

brokcn), suggcst that il represciits a posccrior

locus, probâbly thc last (M6, if thc dental formu¬

la was as hypothcsizcd for Spalacolesies). The

metastylar cusp is promtnent, and there is a

second cusp (C) on rhc postparacrisra, about

midway borwcen thc paracone and thc metastylar

cusp, and separated from the latter by a slight

notch. A weak crest extends into thc trigon basin

from thc base of thc paracone. The prcpai'actista

is slightly lowcf chan thc postparacrista, and

bears à cusp (B|) that is somewhat doser co the

stylûconc (which is brokcn) chan thc paracone.

Discal co die scylocone is- a smati but trenchant,

mesiodistally elongate cusp chat rims the labial

margin of the trigon basin. A crest extending dis-

tally from this cu.sp does not quitc rcach the

metastylar casji, leaving thc trigon basin open

distolablully.

The présence ol cusps B, and C’ on ihc pre- and

posiparacri.scae suggests tliar thc tippcr molar (like

thc lowcr) represents a more primitive taxon tivan

naincd spccics ol Spalacotheriidac Ironi thc mcdial

and l.atc C'rccaccous of North America, and are

rescmblanccs to taxa such as Sfhdacotheyoides (sce

Patterson 1956. fig. \)^ Zhangheotberunn (sce llu

et al. 1997^ fig. 2), and Spalacotherium (see

Simpson 1928a, fig. 34): but it dearly düfers Jrom

upper inolars of dicsc as wcll. indecd» ihe presence

of a cresr extending kbially from che paracone is a

siniilarity to dryolestoids. Lacking any rcasonable

basis for comparison, we defer further comment

on rhese puzzling specimens pending recovery of

additionaJ materials.

Fig. 17. — Spaiacotherüdae gen. and sp. rndet, A, B. left m2?

{OMNH 33896) in occlusal (A) and lingual (B) views: C. right

M6? (OMNH 29612) in occlusal view. Scale bar; 1 mm.

more primitive spccics than others known from

the mcdial and Latc Crcraccous of North

America.

The upper molar (OMNH 29612; Fig. 17C) is

strongly dksimilar to chose ed Symmetrodontoides,

Spidacotheridhiniy Jnd Spalacoleste.^. The absence

of a metastylar projection, curvaturc of thc post¬

paracrista and rounding of the metastylar région,

Genus Symmetrodontoides Fox, 1976

Type sSPECIES. — Symmetrodonloides canadensis Fox,

1976.

Inclltded SPECIES. — The type, S. foxi Cifelli &

Madsen, 1986, and S. oligodofitos Q\ic\\\, 1990.

DlSTRinu TION. — Turonian rhrough early

Camp:inian. western North America.

Rl-VI-Sl-J) DI/UINOSls. — Large spalacotheriids differing

from other niembers oF rhc family in having propor-

tionately broad, acutely-angled posterior lower molars

200

GEODIVERSITAS • 1999 ■ 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

(m4-6). Differs from SpaLirotheridhon in hnvini; lin*

gually pbced pdraconid itid hcîghi difïcrcnnaî bel*

ween paraconid and rneiaconid; diffcrs from

SpaLicolrste^ in h.iving more pronouneed heieht diffe-

rential bciwccn paiaconid and nietaconid; difrers from

boih in having i.iller paraconld un ni], with raUer

paracristid. Ml-2 differ from rhose SpaLicolestes znà

SpaLicotheridium in having a paracone with less bul-

bous base and lingual face tightly arced or foldcd» not

genily curving.

COMMKNTS

Syynmetrodontoides is not présent in rhe Cedar

Mountain Formation; its contents and diagnosis

are included lierein only to pruvide a basis for

comparison. More extended diagnoses of

Syjnmetrodontoides wert given by Fox (1976,

1985), based on comparison with SpiiLicothenum

and rhe single lower nudar of Spalacotheroidcs.

The discovery of additional taxa from the

Cretaceous of North Atnerica shows that certain

feaCLires cired in the earlier diagnoses of

SymmeîrodufUotdes, such as the présence of a

labial cingulum and rhe progressive change in

trigonid angle of the lower molars sériés, hâve a

broader distribution rhan previously known.

DISCUSSION

Fragmcntary as they are, new materials from the

Cedar Mountain Formation of Utah add sub-

stantially to knowledge of symmerrodonr diversi-

ty and morphology in the North American

Cretaceous. Bccausc many taxa are based on frag-

menrary, offen non-comparable remains: because

their teeth (upon which most taxa are based) are

of ratber simple construction; and because they

remain poorly known in general, wc do not

believe that existing data arc sufllcicnt for syn-

thetic considération of symmerrodonr pbylogeny

in the context of early mammal radiations.

Accordingly, we restrict our treatmeiit of rela-

tionships to discussion of character distributions

and their possible implications for affinities of

and within the Spalacotheriidae, summarizing

these data in a traditional hypothesis of relation-

ships within the fîmily.

Sigogneau-Russell & Fnsc>m (1998) hâve present-

ed a detailed treatment of molar characteristics

in vSymmeirodonta (induding many enigmatic

taxa not treated herein), and we hâve relied oix

their work in compding the following compan-

sons. Thèse auchors aiso point ont that, in a

numher of respects, molars of spalacotheriid

symnietrodoncs are dilïicult to distinguish froni

chose of certain dryolesioid eupaniotlieres:

indeed, it is po.ssÜïle iliat dryolestoids arc more

closcly related lo s-palacotheriids' than has heen

generally believed (see, e.g., Sigogneau-Russell

1991a). In fulfilling our intent to fotus on

Spalacotheriidae, we cannot attempt coniprelien-

sivc coinparisons hcrcin. We aeknowiedge the

exisieOcc of a nuJiibcr of similaritics in rhe

molars of advaneed spalacothenids and certain

dryolcsdds,. such as the presence of a distal srylar

cusp and U hooklike parascylar lobe, mcsodistal

compression of the crown, and oiher features

cited by Sigogneau-Russell & F.nsom (1998), os

well as the markedly lower placement of lower

molar labial cingulum than lingual cingulum. At

the présent State of knowledge, we believe ihar

referral of ihc taxa considered hercin from the

North American Cretaceous [Spulacotheroide^,

Symmetrodontoideÿy Spalacotheridiuin, Spulaco-

lestes) to .Spalacotheriidae Is more compelling

than to Dryolcstidae, based on features of the

dentary (e.g., peculïar development of the ptery-

goid crest and masseteric flange) and lower den¬

tition (e.g., cxcreme réduction of the calonid and

characteristics of the roots, sec Butler 1939).

Based on size, inorphological appropriateness

and dissimilarity to other known éléments of the

Musscntuchit local fauna, and (cspecially) rcla*

tive abundance and distribution among rhe

known localitics. we believe therc is vam'shingly

little doubt as to the référencé of upper and

lower dentitions to the respective species describ-

cd herein.

Dentary

rhe structure of the dentary and associated post-

dentary" boucs has figiircd promincntly in discus¬

sion of the origin and carly difTcrcntiation of

mammai.s (e.g., Crompton &: jenkins 1979:

Kemp 1983). Mandibics ascribed to Kitehneo'

therium suggest that a full complément of at-

cached postdentary boues was retained (Kermack

et al. 1968), as they were in Morgamicodon (see

GEODIVERSITAS • 1999 • 21 (2)

201

Cifelli R. L. & Madsen S. K.

Kcrmack ^////. 1973), Marsh, 1881 (sec

Kermack & Musscti 1958), and Haldanodon

Kühne & Krusac, 1972 (scc Lillcgraven &

Krusat 1991). In tlie.se taxa, the postcromcdial

face of the dentary beats a prominenc postdenc-

ari'^ trough, overhiing by a ridge, exicndiiig ante-

riorly from the condyle. l lie postdentaiy txougli

housed the articular. prearticular, .surangulat, and

angular; attachmenr lacets for the coronoid and

s'plenial are gcnerally aiso visible, more anteriorly

on the dcniar)' (e.g., Kcrmack et ^4L 1973, Hg. 7;

Lillcgraven ôc Krusat 1991. bg. l4). 3 hc mcckc-

lian groove cxcends anrcriorly from the mundibu*

lar foramen and is connuenr wirh ihc

postdentary trough; in Mor^anucodon ir housed

the anterior part of the prearticular (Kcrmack

et al. 1973), whereas a splenial is associated wirh

(or overlies' part of) the meckelian groove in

Haldanodon (see Lillcgraven ik Krusat 1991).

Sevcral orher poorly understuod or arthaic mam-

mais retain an csseiirially siinilar condition,

although the trough and ridge arc noi as well

devcloped (e.g., Shuotheriam Chow & Rich,

1982; Ansktribosphefun Rich et al. 1997). In

mosr remaining mammals, the meckelian groove

(where présent) is separaied from the mandibular

foramen and the postdentary trough and corres-

ponding ridge arc Iost> suggescing detachment of

the main body of postdentary éléments from the

dentary, although arcachment at thcir anterior

extremit)' evidently persisted in sonu eupanro-

theres, such as Amphilhernim RUinville, 1838

and Peramm Owen, 1871 (see Allin ik: Hopson

1992). The condition may be .similar in rhe Lare

Jurassic or Larly Oetaceous symrnetrodonr

Zhangheotheriarn^ which has a pr4)minent mecke¬

lian groove and scars for the coronoid and splc-

nial (Hu et ai 1997), and, possibly, another

élément in addition co the dentary (R. C. Lox,

pers. comm.). A small coronoid apparent!)' per¬

sisted in numerous rnamnvallan groiips. First

reporled among '‘paruotheres" by Krebs (1969),

facets suggesring pre.sencc of rhe comnoid hâve

been reported in the dr}'olasiaid Henkelotherhim

Krebs, 1991 (see Krebs 1991), varions Iricono-

donis including Phatadatheriitni t3wcn, 1838

(BM 112) and Cjobkonodon 'Woîxxnov^ I9’^8 (see

Jenkins & Schafl 1988), the cutherian

Prokennalestes Kielan-Jaworowska & Dashzeveg,

1989 (sec Kiclan-Jaworowska k Dashzeveg

1989), paulchoffatiid multirubcrculatcs (Hahn

1977), and the spalacotberiid Spalacotherium

(BM 47750). Pcrsistence of rhe meckelian groove

(or a remnant of il), which may hâve housed

rcmnaïus of onc or more postdentary éléments,

is even more widespread (scc discussions in

Benslcy 1902; Simpson 1928b; Kcrmack et ni

1973). Among symirietrodoius, the meckelian

groove is présent in Knehaeaiberium (see

Kennatk ét al 19681, Tinodoa (see Simpson

1929), Zhangheodiernim (sec Hu et al 1997),

Sbiioiberium (scc Chow k Rich 1982), and

SpaliU'othenurn (see Simpson 1928a). 3 he pré¬

sence and form of the meckelian groove hâve

been used in interprering rhe phylogeny of

Mesozoic mammals (Luo 1994; scc also Hu et al

1998). Among the aforementioned ta.xa, the

meckelian groove is redueed anteriorly in ail; in

Zbitngheotberiam^ what remains is subparallel co

ihc axis ol the dentary, whereas in KnehiteO'

tbenum. ilnodoru Spalacotlmimn-, Ttnodon, and

Shuotberium the meckelian groove converges

toward rhe ventral margin of the dentary anre-

riorly. This ktrer State is presumed to be more

derived (Luo 1994), but the signitlcance of this

distribution for symmetmdonts is unclcar. Both

rhe coronoid and meckelian groove are lacking in

Spulacolestrs, and avaÜablc e\'ideïKc suggesLs that

rhe meckelian groove, ac leasc, was lacktng in

Symmetrodantüides. Hence, both of thèse fe-atures

were lost wirhin Spalacotheriidae, assuming

monophyly of rhe family.

In primitive mamm.als such as morganucodon-

tids and Kuebneotheriinn, the ventral margin of

the dentary is emarginated posteriorly (as ic is in

advaneed q'nodonts). w'hcrc the postdentary clé¬

ments arc positioned (sec. e.g., Kermack et tü.

1968, 1973; Gamharayan & Kiclan-Jaworowska

1995). The évolution of an angular regii»n and

procesN on the dentary of more derived mammals

is imcenain becxuise (»f dispiited homologies and

differing criteria on which récognition of these

characrers are based (see excclicni discussion in

Wible 1991). An anteriorly placed process is pré¬

sent in cynodonrs (Sues 1986) and certain primi¬

tive mamjTials, such as Morganucodon (see

Kermack et al 1973), Docodon (scc Kcrmack &

Mussett 1958), and Haldanodon (see Lillcgraven

202

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Crctaceous of Utah

&C Krusar 1091). Rowe (1988) considercd the

loss ot the anteriorly placed process as an ad-

vanccd feature characterizing multitiiberculates

and thcrians. Patterson (1956) disagreed with

identification of rhis feature as a true angular

process, pointing ont the dilfercnce.s in hoth

position and inferred function (see aiso Proihcro

1981). In Dhinctherinm |enkins et ni, 1983 (sec

Jenkins ettil. 1983), this anteriorly placcd pro¬

cess occurs togerher with une ihar is more poste-

riorly placed, near the condyle, snpporring

Patterson (1956)s suggestion that the former is

not homologous witli the angular process found

aniong therian mammals (jenkin.s et ai 1983; sce

aIso Crompton Luc 1993). Sues (1986; see

aIso Gambarayan & Kielan-Jaworowska 1995),

however, suggested that the angular process of

trirylodont therupsids and certain therians is

homologous, indicaiing that rhe continuous ven¬

tral margin ol lhe dentary as seen in taxa such as

symmetrodonts could have been lormed through

fusion of the two proce.sses seen in Dinne-

therium, We can offer nothing in i[»e way ol new

data to résolve this issue. It is worth pointing out,

however, that ail students who have commented

on the macter have observed thaï symmetrodonts

(e.g., Knehneotheriunh Tmodoru Spalacotheriiim,

Zhangheotherium) lack an angular process or, for

that matter, any ventral or postcrovcniral expan¬

sion of the dentary in the angular région (c.g..

Simpson 1928a, 1929; Prothero 1981: l lu et al.

1997); an angular process is aiso lacking in trico-

nodontids and gobiconodontids. This is distinct-

ly different Irom the condition (whcre known)

arnong eupanrotheres and peramurans (e.g.,

Amphitherium^ Pernmtts, Dryolescidac: sce

Prothero 1981) and among tribusphenic mam-

mais. Wc tentativcly recognize the development

of a posteriorly placed angular process as a de-

rived feature characterizing eupanrotheres and

tribosphenldans (Trechnotheria of Prothero

1981 ). This feature is absent in Spalaadestes, as it

is in Spalacotherium and ail other synimc-

trodonts.

A pterygoid crest or pceiygoid shelf (Miao 1988;

Rowe 1988; Gambarayan & Kielan-Jaworowska

1995) on the mcdial surface of the mandiblc

may be related to increased importance of the

pterygoid muscle in mandibuiar adduction,

translation, and rotation (e.g., Oron ôc

Crompton 1985). The pterygoid crest is lacking

in primilive taxa such as Morganticodnn (in

which rhe médial pterygoid muscle i.s inicrprercd

lo have been small, Crompton &l Hylandcr

1986) and Ktichneotherium. Prothero (1981)

cited the présence ofa pterygoid crest as a synapo-

morphy of Symmctiodonia (including

Spalacotheriidac and Amphidontidac), but this

crest enjoys a considerably broader distribution,

being présent in cupantoihercs (c.g., Laole.<tes

Simpson, 1927; Amblotheriiwî Owen, 1871; sce

Simpson 1929: 63, 68), triconodontids

(Simpson 1928b), gobiconodontids (jenkins &:

SchafI 1988), mullituberculates (Miao 1988;

Gambarayan &c Kiclan-Jaworowslu 1995), and

rribo.sphenitl.ins (Kielan-Jaworowska &

I9ashzeveg 1989, fig, 2Ü; Sanchez-Villagra &

Smith 1997). as wcll as symmetrodonts (except

Kiiehneotheriurn). Rowe (1988, 1993)’s Therüo-

morpha includes a pterygoid shelf as a diagnostic

character (see discussion in Miao 1993).

Allhough the pterygoid crest (and its continua¬

tion onto the angle, whcre présent, ol the therian

dentary) may be developed as a médial shclt

(e.g., multitubercLilates, Marsh 1880;

Gambarayan Kielan-Jaworowska 1995) or

inturned process (c.g., somc Crctaceous

Eulheria, Kielan-Jaworowska et al. 1979), the

condition among marsiipials (commonly termed

an inflccted angle) has been suggested to be a

derived characrer unique to the group (Sânehez-

Villagra & Smith 1997). However, usîng the

définition these authors provide (‘‘a medially

inficcted angular process may be defined as onc

that projects inward (lingually) at about

90 degrces with respect to the dorsoventral plane

of the mandibuiar ramus,” Sanchez-Villagra &

Smiili 1997; 120), we observe an inflected angle

ro be présent iu Prokenmilestes (e.g., GI PST 10-

5C. 10-6).

In Spalacotherium., the pterygoid crest originates

just below the mandibuiar foramen. The crest is

incompletely preserv'cd in available specimens, so

that its full extern cannot be detennined.

However, it was greatly expanded and, perhaps,

developed a medial process in chc région of the

mandibuiar foramen, an unusual feature among

early mammals. This clearly was the case with

GEODIVERSITAS • 1999 • 21 (2)

203

Cifelli R. L. & Madscn S. K.

Spalacolcste^^. Here the preryj;oid crest begins jusr

bclovv thc aivcolat niargin at ilie juncribn of horb

zonral and ascending ramij dcsccndlng postcro-

ventrally ro ihc rcgton of rhc mandibular

foramen» whcre it is dcveloped as a large, cuiving

proccss that encloses a small pocket. The strong

developmein ofihc pterygoîd cresi inro a process

near the mandibular foramen is unusual among

early mammals; it is lacking in oïlicr symmetro-

donts such as Tinodon and '/Juingheatherium.

'rhe condition in Spulacolesfesy in vvbicb thcre

cicarly is a largCs reflcctcd proccss and the picry-

goid crest cominues amerodorsally» is undoub-

rcdly unusual and apomorphic. An edcntulous

dencary tliar we mfcr to Spakicotherofdcs bridwelli

(FMNH PM 1025) préservés sorne limited

information on the médial side oi thc jaw in dus

species. The prerygoid crest clearly originared

just below thc alvcolar margin and exrenJcd

posterovcntrally, as iit Spalacolcstes, The spécimen

is considerably abraded in the vicinity of the

mandibular foramen, and ir is unclear whether

the pterygoid crest was developed into a proccss,

as in the taxon from Urali. To our knowledge,

the only other Meso/oic mammal wltli a prery-

goid crest that extends antcrodorsally and nearly

reaches thc alvcolar margin ol thc dentary is

Prokeiinalestes (sec Kiclan-Jaworowska 6c

Dashzeveg 1989, t'ig. 20).

A related and noieworrhy feature on the médial

side of tlie dentary in Spalacolestts is the grcat size

of the pterygoid fossa (we arc indebted to Z. Luo

for suggesting tliis co us). In most Mesozoie

mammals, such as triconodonfids (sec Simpson

1928a) and gobiconodonrids (scc jenkins &

Schaff 1988), the pterygoid fossa extends ante-

riorly ro about the levcl of rhe mandibular fora¬

men; in Sptllacoltsies^ thc pterygoid fossa extends

well anterior to the mandibular foramen. Among

spalacotheriids, the presumed primitive condi¬

tion (pterygoid fossa terminâtes anteriorly adja¬

cent to the mandibular foramen) is présent in

Spalacothcrium (sce Sirnpson 1928a) and

Zhangheotheriuïn (sec Hu e7 al. 1997), whcreas

the pterygoid fossa extends farther anteriorly in

SpalacotherouUi (1 MNH PM 1025). I he pos-

terior part uf the dcncary is not known in other

taxa; noiicthelcss, this distribution suggests thc

possibility that the derived condition may repre-

sent a shared, derived feature of North American

Spalacolheriidae.

1 hc ventral margin of the dentary in the angular

région of Spalacotherjum is reflcctcd latérally into

a ‘‘widc. eiflected, flange-like massctcric crest," as

vSimps'on (1928a: 102) noied. flie same is rrue,

to a gieatct degree, in SpaLtcolfstes. Alrhough the

posteroventral margin of ihe dentary ha.s some

laier.il reflection in certain rriconodonts (l iopson

1994) and gohiconodontids (Jenkins & Schaff

1988), it is not significantly developed in other

symmerrodoius or other Meso/.oie mammals,

and the degree of reflection seen in SpaLuo-

ilmium and (espccially) SpaLnolvues is unitsual.

The massecer muscle tnserts along the vcnrial

and lacerai side of the denrant' in rhts région, and

the unusual condition in spalacotherüds is likely

related to the dcs^elopmcnt or configuration of

this muscle (thc mas.seieric fossa is aiso railver

deep in Spalat oihcrhim and Spalacolestcs).

Lacking ihe skuli. functional interprctaiion is

limited. However, rhe mandibular symphysis was

probably unfused, as suggested by RM 47748,

referred to Spalacotheriam. The mandibic of

Zbaagheotherhim was aIso unfused (sec Ilu et al.

1997), as is thc case wirh most early mammals

(CTompton Hylander 1986). It is likely that

mastication in spalacothciiids involvcd signlfi-

cant componenrs of mandibular rotation and

latéral translation: tlte massctei serves to invert

thc dorsal border ol thc hcmimandiblc on which

it inserts, and thc pterygoid serves to evcrc it

(Oron C’rompton 1985; Ciompcon 1995).

Kcncc, ihere is reason to believe that thc unusual

condition ol thc pterygoid crest and thc latéral

reflection of the posteroventral margin of thc

dentary in spalacorhcrlids arc fimctionally related

features, and that they may intlicate .some un-

usiwl aspect (perliaps strong development) of

latéral translation or rotational movement in the

masticaiory cycle of chose mammals, as suggested

by considération of their shearing surfaces (see

aiso Patterson 1956: 57; Crompton 1971;

Sigogneau-Russell & Lusotn 1998). As has been

described for lower molars of Nortti Anïeiican

Spalacotfieriidac (Fox 1976; Cifelli & Madsen

1986), upper molars o'f SpaLuotheridiuTn and

Sptdacalestes bear obliquely oriented wear stria¬

tions on both prevallum and postvallum surfaces

204

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Fig. 18. — Microwear structures in Spalacolesies cretulablatta

n. gen., n. sp.; bottom, mesial view showing prevallum shearing

surface on left M4 (OMNH 30611); box indicates location of

enlarged photo, above Scale bars; top, 0.1 mm; bottom.

1 mm.

(Fig. 18). The relative contributions of latéral

translation and rotation of the mandible în the

masticarory cycle caniior be determined, although

il is likely that both played signifîcant rôles.

Lowf.r MOLARS

We bclieve it highly probable that the lower

tooth séries of Spalacolesies (and> tcniatively,

Spalacotheridium) included seven molars, as in

Spalacotheriuni (see Simpson 1928a; Clemens

1963); four itre présent in Tinodon (see Simpson

1929), six in Zhangheatheriuvi (the described

specimen is a juvénile and it is possible that not

ail molars were erupted, sec Fiu et ai 1997), and

three lo six in Kiiehnenthcriiim. Bascd on siudy o(

edentuloas jaws, the lower part of ihis range wa.s

thought to bc most probable for Knehneotherium

(sec Kcrmack et ai 1968). Study of isolatcd teeth

suggests that as maiiy os six molars rnay have

been présent in somr individuak (Mills

the most complète alvcolar row known for the

genus shows that at least fivc were présent on this

specimen (Gill 19^4). As noccd by Patterson

(1956), rhe molar cotint in symmetrodonts is

correlated with angulation between principal

cusps: “acute-angled” symmetrodonts arc charac-

terized by a high number of molars. On this

basis, rcmaining ntxa from ihc North American

Cretaccous (SpaLtcotheroides^ Symrnetrodontoides)

probabl}^ aiso had a higli number of lower molars,

seven being rhe Jikdy number. judged by compa-

rison to Knelmeatherimn and orher outgroup taxa,

the aciite aitgulation and higli molar count of spa-

lacothcriids represenrs thederived condition.

Individual molar charactetistics aJ.so set North

American Cretaceous taxa and SpalacoTherium

apart Irom Kuebricocherium, Thiodon, and certain

other symmcirodoiit-s. In Ktiehncotherium (and

probably Wotttersia. as well), as in morganuco-

dontids, précisé occlusion is dépendent on deve¬

lopment of extensive wear facets on corres-

ponding upper and lower molars. In spalaco-

rheriids, however, matching molar surfaces fit

more precisely (Crompton ^ l lylandcr 1986;

Crompton 1995), as they do in rribosphenic the-

rians. In Hnodoriy as in Kuebneotberium. separate

wear lacets devekïp on occluding surface.s of the

principal cusps^ whcreas iii spalacoihcrtids,

strong crescs are developed between chc piotoco-

nid and lingual cusps, and these form continuous

prevallid and postvallici shearing surfaces

(Crompton üsC Jenldns 1967; Crompton 1971).

The condition in Zbanghmberiiim is tincertain,

but it bas been described as differing from other

spalacorheriids in having roundedt cortical cusps

that lack connecting cresrs (Hu et ai 1997)>

strongly suggesring that ir is similar to Tlrwdon

or Kuelnieotherium in lacking continuous shear¬

ing surfaces.

GEODIVERSITAS • 1999 • 2l (2)

205

Cifclli R. L. &c Madscn S. K.

The cingulum and cingular cusps alsn are

variable among .symnierrodonts. In Kuviutea-

therhim and Tinudon, oïdy a lingual eingulum is

présent on rhe lower molars, the distolingual

(talonid) cusp Ls moderatelywell developed, and

two mesial cingular cusp.s are (>resen! (Crompron

Jenkiiis 1967). A eingulum is laeking Ironi

lower molars of /.luxnght'Othatum (see Hu t't /?/.

1997); it is incomplète labially, at Icast, in mosc

othcr primitive mammals, sucli as Woutersia (scc

Sigogncau-Russell & Hahn 1995). Morgauo-

codon (sec Kcrmack er al, 1973). and Amphi-

dontidae (sec Simpson 1929; Trofimov 1980). lu

spalacothcriids, a .single mcsial cusp is présent

and the talonid is soniewhat snullcr than it is in

Tiuodon and Kitehneorheyinni (Crompion ik

jenkins 1967). In addition, ail spalacoiheriid.s -

with the exception ot Zhangheotheriuni and the

apparent exception of Spaliicotheivides (sec above)

- have a complète labial eingulum on the lower

molars. In vill of thèse Ratures, Spalacotheriidae

appear to he derived wirli respect to remaining

symmernulonrs (see Sigognc.tu-Kiissell & Ensom

1998 for di.scus.sion ot the di.strihuiion and varia-

bilir>' in lhe development ol the lower molar cin-

guhim among .svmmcfrodoius).

Spalacf>theriidae also nppear to he utuisual

among Meso/oic mammals in the nianncr in

which rheir lower molars interlock (we are grate-

ful to Z. l uo for poiïtiing this ont to us). In

morganiicodoniids and docodonts, the distal-

most molar cusp, cusp d, fit.s bcivvccn cusps h

and e ot the succccding lootfi (sce, c.g.,

Crompton ^ jenkins 1968); prcsutnably modi-

fied arrangemenrs are présent in othcr taxa, siich

as Kiiehneothcrlum. gobicoiiodoiuids-, and crito-

nodoniid.s (see Luo 1994; Kielan-Jaworowska &

Dashzeveg 1998). In spakicotheriids, rhe distal

cingiilar cusp (which appears to bc homologous

with cusp d) is placed lingual ro the mcsial cin-

gular cusp (which uccupies j similar position to

cusp c and may bc honiologous with it) of the

füllowing tooch, and the mcsial cingular cusp fies

into an embayment of the eingulum char is labial

ro cusp d of chc preceding tuuth.

Within Spalacotheriidae, taxa from the North

American Crccaccous appear to bc advaneed with

respect to Spalacotherium in having more acurely

angled, higher crowned lower molar trigonids;

SpiiluLoîhcridium is rhe lea.se extreme in this

regard (rhe condition is unclear in Spalaco'

theroides, but it is prohably safe to say lhat its

lower molars are more acutely angled than are

chose of Spulacotheriunt). In spccic.s of Spalaco-

and Symnmyodontoidts^ the pafact)nid and

paracristid are di.siiiictly lower than the nieta-

conid and inetacrisfid on ail lower molaf-S (except

m7, which in Spalucalvsit's at lca.st. lacks a meta-

conid); in spccics of ^ymmctrodontoidcs^ the

height différenciai is accentuaied. The cusps are

subequally developed in Spalacathcridium,

Spidacotherium (sec Simpson 1928a), and Ihw'

dtny except on ml, whcrc the paraconid is shglit-

ly smallcr (c.g.. USNM 2131). In Woutersia (see

Sigogiieau-Russcll & Hahn 1995 ), hy coiurxst,

the paraconid is the laller of the two; in

Kitelnieotberium (sce Kcrmack et ai 1968; Mills

1984), the condition is variable and chc cusps

should bc consideivd subcqual in development.

VV'e concur with Sigogneau-Russell & Knsom

( 1998 ) in helieving that paraconid and meta-

coniil were primitively sul)equal In SpahiLa)-

theriidae (as thoy arc, for example, in /hangheo-

theriurn),, and concludc that the condition in

Spaincolestes and Sytametrodontoides is derived.

Both of the last-nieiuioned genera arc characte-

ri/.cd hy tlie présence of posterior lower molars

th.'ir are labiolingually exp.iiided, ow'irig to a rcla-

tively clongalc paracristid and lingually placed

paraconid,

Vm n MOI AHs

As with the lower molars, chc upper molars of

Spalacotheriidae are more acute-anglcd than

tliose of primitive taxa such as Kuehneotherhm

(sce Kcrmack et al. 1968) and Tinodon (.lec

Simpson 1929). Up(»er molars of

Synnnetrodontoides, and Spahiaithcridium, at

least, arc more acutc-anglcd (assumed ro repre-

sent rhe dertved condition) than rh(»se of

Spalacotherium oi Zhangheorherium. The single

upper molar of Spiilacotheroides ilhisrrared by

Patterson (1956, fig. 1) prohably represents one

of the mesial loci; other specimens (e.g., I*MNH

PM 1133. 1236) are distinerly more acute.

Mitroderson (sce Sigogneau-Russcll 1991b),

which resemblcs North American taxa in several

other respects, is also rather acurely angled. The

206

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

paracone on upper molari of SpaLicotherium and

Zhangheotheriiinu like chat of North American

Spalacotheriidae, is extremely tall relative to the

condition seen in most outgroup taxa (e.g.,

Tinodon, Kuebncolherhmi). We suspect chat para¬

cone height will prove to bc a useful charuccer foi-

or within vSpalacothcriidac whcn ics distribution

becomes better kiu)wn.

Récent studies (e.g., Hu et td. 1997; Sigogncau-

Russell & F.nsom 1998) hâve followcd I^actcrson

(1956) in idcntdying the inesiolabial cusp on

upper molars oi SpulaiothcrHiifi (lacking at

mesial loci, see Butler 19.^9) as the .stylocone, or

cusp B; hence, the more linguaJly* placcd cusp on

the preparacrista is a necnnorph by comparison

lo outgroup taxa such a.s Kuehneotherhim. This

cusp (ccrmed B, by Hu et ai 1997) is aiso pré¬

sent in Zhangheotheriurn and Spalacothernides

(cxcept the mesial molar, FMNH PM 1235,

figurcd by Patterson 1956): is ic primitive for

Spalacoihcriidac, or a derived feature characceri-

zing these threc general Üther fcacures, such as

the présence of fc\vcr than .six upper molars and

the ?lack of continuons shearing surfaces, suggesr

chat Zhanghvotherlum rcpresencs the sister-taxon

to rcmaining Spalacocheriiclae, in curn providiug

somc tentative cvidcpce tliar the presence of rhis

cusp may be primitive for the Kimily. Under this

interprétation, rhe absence of cusp B, in North

American Spalacotheriidae (cxccpt Spalaca-

theroides) would represenr a loss. Piîttcrson

(1956) alsü suggested that the mesialmost srylar

cusp of SpaliKotheroides, which lies mesial to the

preparacrista, is a neW cusp not présent in

Spalacotherhini, and tluu it represerus one of

several, Independenr acquisitions of a parasiylar

cusp in therian mammai.s. Interprétation of this

fearure is probletnatic. A parastylar cusp is lack¬

ing in Zhangheoihenurn (sec Hu et al. 1997). The

single knovvn upper molar of Tuwdon (YPM

13637) lacks the parasttHar région of the roodi,

airhough Crompton (1971) restored [inudvti

with a parastylar cusp. A cusp in this position is

présent in Kuehfieotheriuyft (sec Kcrmack cl al.

1968). Regardless, rbe prominenr,. hookÜke para-

srylar région seen on anrerior rnolar.s ut

Symmetrodontoides (see Fox 1985), Spalaco-

theridiuitti and Spalacolestes would appear to be

an advanced condition by comparison to remain-

ing taxa (.sec Sigogneau-Russell & tnsom 1998).

Of ihe remaining upper molar cusps, cusp C is

well niarked in outgroup taxa such as

Kiichricotherium and Tinodon, and is présent in

/.hangheotheriuniy Spalavatheriurn^ MterodersoHy

and Spalaeotheroides. This suggests that the

absence of cusp C in taxa from tlie médial and

Lare Crecaccous of North America represents an

advanced condition. As described by Patterson

(1956), Spalacoïbcriddcs has rwo discolabially

placed styiar cusps (a metasryle and orie thaï is

more mesially placed at the margin of rhe styiar

shelf), whereas only one is présent in Spalaco-

therium (see Clemens 1963: 376). A single disro-

labial cusp (merasryle) ix seen in KtielmeotherJuni

(see Kermack et al. 1968), Microderson (sce

Sigogneau-Russcll 1991b), ànd Zbangheolherhwi

(cusp "D” ol flu et ûL 1997). Posiiional cvidence

favors homology of che distalmosr cusp (meta-

style; see above) on upper molars of these taxa,

suggesting that the more mesial cusp of

Spalacotheroide.< is a neomorph. Taxa (rom the

médial and Late Cretaceous of North America

hâve rhe disrolahially placed metastylar cusp and,

more mesially, a mesiodistally expanded cusp al

the margin of tlie srylar shelf. Wc tentacively

regard the latter as hornoJogous with the mesial

ol ihc cwo discal stx'lar cusps in Spalucothewides

(which wc interpret to bc scylat cusp “D" of

Sigogneau-Russell & Ensom 1998), and its pré¬

sence as an advanced fearure char.aeteri/.ing

North American Spalacotheriidae. The condition

in Spalacnlestesj Spalacotheridium, and

Syrnmetrodontoïdes^ in which the mesial of the

two cusps is strongly dcvclopcd on mesial

molars, represents a more derived condition. A

strongly dcvclopcd distal srylar cusp is also pré¬

sent in the cnigmade Thereuodon from the F^arly

Cretaceous of Morocco and England, which is

othcrwisc so different as to be non-comparable

(see Sigogiieau-RusscÜ 1989, Sigogneau-Russell

&: Ensoni 1998).

In Spalncoibcroides, as in remainiag North

American Spalacotheriidae, the preparacrista is

markedly lower than the postparacrisia, particu-

larlv on mesial molars. judged by comparison

with linodon and Ktiehneotberumi, this appears

to bc a derived condition, but the distribution of

the character is difFicult to déterminé from avail-

GEODIVERSITAS • 1999 • 21 (2)

207

Cifelli R. L. & Madsen S. K.

able litcracurc. The preparacrista is cicarly lower

than the poscparacrista in MurodersoHy whcreas

the crests appear ço be subeqüal in iie\'eIopment

in Spalacotherium (sec Simpson 1928a, fig. 34)

and Zhaugheorherium (scc Hu ej al. 1997, fig. 2).

As notcd bv box (1985), the styloconc is so small

in Syninmrodoutoides that il is indistingnishable

in tecth that hâve bcen subject to even niodcratc

wcar. i'he stylocone is also siTtalI in Spalacolestes

aiid SpalacotherUiuw. ComparLson wich remam-

ing Spalacothcriidac {SpnlacatheriKm, Spalaco-

rheroideSy ZhangheorhiriimAy as we!l as Kuehneo-

iherium. Wouterslay and Tmadon (in which the

stvlocone is prominent), indicaccs that the stylo-

cône has probabty iindergone réduction in the

taxa from the médial and Lare Creraccous of

North America (see also Sigogneau-Russell &

Ensom 1998).

Comparison of upper molar shape and propor¬

tions in Spalacotheriidae ïs probleniatic because

sériés are kiii)\vn for so fcw taxa and because, as

shown above, coronal profile and degree of sym-

metry varies according to tooth position. Ml/l

Spalacothcriron h cons'idcrably smaller than

succceding molats. whereas in SpaUcotheridiinv

and Spabfüleitei. Ml/l is rcbtively much larger

(e.g., Figs 6. Il, compare with Burler 1939,

fig. 7). Insufficienr bas'is for comparison leavcs us

uneertain as to the significance and pobrity of

this fcaturc. Spalucolheridium, Symmetro-

dontoides, atid SpaLicolesies have an M4 that is

rcmarkably symmctrical. Pasc this tooth position,

molar size decrcases, and rhe lasr molar is signifi-

cantly smaller than its prcdccessor. Comparison

to Spalacotherium suggests char thèse inay be

advanced conditions, hi Spalacotherium^ poste-

rior molars seqiicnrially develop a [urascylur lobe

that, on the last tooth, prujeas sirongly. A small

parastylar lobe is présent on posrerior molars of

Spalacothcndium but lacking in Spalacolestes and

Symmetrodoutaidei, suggesting réduction in rhe

larter t\vo rnx;t. Ail ihree Norrh Américain généra

differ from Spalacotherium in having the para-

cône progtessively placed more disially on posie-

rior molars (vagucly recalling a similar shKi in

rhe protocone of (josterior upper molars in iribo-

sphenic mammals). In terms of crown relief.

Spalacolestes and Symmetrodontoides differ from

Spalacotheridium in having a relatively deeper tri-

gon basin. Microdersoti also appears to hâve a

deep trigon basin, but this appearance is due to

the fact that the paracone is extremel)' lall in this

taxon. In remaining Spalacothcriidac, the crigon

i.s much shallower rhan in Spalacolestes and

Symmetrodouioidesy so we interpret a shallow tri¬

gon basis as niost probably representing the pri¬

mitive condition.

ReLATIONSHII'S and CONClUniNt; REMAJCKS

Ib sumniarize» rhe limited data in hand süggcst

that reccntly described Zhatigheotherinw is primi¬

tive in .several respects, and represenrs rhe ÿisrer-

taxon to remainmg Spalacotheriidae (Fig. 19).

Spalacotherium^ in turn, evidenfly is the sister-

taxon ro Norrh American Spalacotheriidae. (We

have omitted MkrodersoUy known by a single

upper molar, from our phylogeny. As notcd

above, this Moroccan taxon resembles one or

anorher of the North .American spalacorheriids rn

-several respects, bui ilie signiiicance ol ihese

resernhlancc.s caiinot bc evaluated with data in

hand; sec discussion in .Sigogncau-Ru.ssell

1991b.) Of ihc Norili American spalacothcriids,

tiic geologically oldesr, Spnlacotheroides hridfoelli

from rhe Aprian-Albian, appears to retain the

mosT number of primitive fearures, sucli as the

présence of ciisps B| and C on ilie upper molars

(assuming that the presence of these cusps is pri¬

mitive for Spalacotheriidae). 5. hridweUi is un-

usual wichin the familv in that rhe labial

cingulum of lower molars is apparently incom¬

plète (see Fox I9'^6). If this contÜtion is correctly

interpreted and not simplv a matter of préserva¬

tion (the holorype, FMNll PM 933* includes

the only known lower molar of rhis species, and

ir iiiriy be abraded). dien we belteve 5. briduelli

CO be auiapomorphic in this respect. Of temai-

ning généra, neirher Spalacotheridium nor

Spalacolestes (nvo specics each) is characterized by

known synapomorphies, uniess theslight élonga¬

tion of rite paracristid on lower molars of

Spalacolestes cretulahlatra and S. incoucinnus

represenrs a shared dci îved character. Symmelro-

dontoides (ihrce spccie.s} is the geologically

youngcsi and most advanced member of the

faniily, characterized by labioHngually expanded

posterior lower molars.

Some members of rhe Musscntuchit local fauna,

208

GEODIVERSITAS • 1999 • 21 (2)

Spalacorheriid symmetrodonts from the mid-Cretaceous of Urah

Fig. 19. — Hypothesis of relationships among Spalacotheriidae: Kuehneotherium included as outgroup, Microderson omitted becau-

se of insufficient data; additional North American species {Symmeîrodontoides oligodonîos, Spalacoîheridium mckennai,

Spalacolestes inconcinnus) nol included lhey are poody Known and, based on présent knowledge, add no detail to the phy •

logeny presented. Characters at nodes isee discussion in tekt): 1. (Spalacotheriidae) molars acutelv angied; ?gain extra cusp (6l of

Hu e! al 1997) on upper motars? single mesial otngular cusp on rower molars rgduce distal cingular cusp (talonid) on low&r molars;

unique lower molar interiocking mechanisrn. whereby ihe distal dngular cusp ol one molar is placed labial lo (he medal cingular

cusp of lhe suoceeding tooth: 2. six ot more uppsr and lower molars présent; ?pterygoiil crest sirongiy developed m région ot mand»

bular foramen (known only for Spalacothenom and Spalacol^stos). continuous prevallum and postvalium shearing surfaces on

molars; '?po5ti3rointerior border çi dentary efnected (known only for Spalacothaduoi and $pala<^oleski$). ?labial ciogulum complété

on lower molars (lacKing in Spalacotherpides). 3. (Spalacofestifiay) meckelian groovo lost; pterygo'd crest extends anterodqrsally (o

near occlusal margin of dentary, witti plerygoid fossa exfendinq well anlerior to mandibular foramen (known only for

Spalacotheroidffs. Spa>acole6fef(\: niolars hiaher crowoed, more aculely angled; upper molars wtth parastyle isee Patterson 1956),

distal slylar cusp. ano preparacnsta fower tfian poslporacnsla (antenor loci): 4, upper molars wlih reduced stylocone. distal styiar

cusp eniarged sîronç hooklike parastyie (arrterior loa}, M4 stroogly symmetricâl, C cusp lost: MG reduced. with paracone posierioriy

placed (condilioo uncedam in Spalacoïhero*de6, Symmelrodontofdes), 5. ?cororioid facel lost (condition ynknpwn in

Spalacotheroides. Spalacoîheridium. SyirtmeUodonloidesY, upper molars with deep trigon basin, reduced parastyle (distal looi; condn

lion unknown in Spatacoineroides): lowor molars with paraconld and paracristid lower than metaconid and protocristid, respectively:

paraconid of distal lower molars linguaily placed. with paracristid dislinctiy longer ihan protocrisiid; upper molars willi cingulum com«

plete linguaily; 6. height differential between paraconid and metaconid on posterior lower molars pronounced; posterior lower molars

broadened; ml with taller paraconid and paracristid; M1-2 more acutely-angled, with less bulbous paracone base and more tightiy

arced or folded lingual face to paracone.

norably several groups of dinosaiirs, appear to

rcpresenr éléments of a mid-Cretaceons immigra¬

tion event frotn Asia (Cafelli et itl. 1997). Origin

of other taxa is more problcmaric. North

American Cretaccous triconodontids. for

example, appear to rcpreseni a monophylctic

group, but their origin within the known Jurassic

diversity of the family (both New and Old

World) is uncertain (see CifelÜ et al 1998)^ If

North American Spalacotheriidae are mono¬

phylctic, as wc SLiggest, then their biogcographic

tics antedate ihc hypothesized mid-Crctaceous

interchange, xs they would icpresent a group rhat

wa.s c.stahlished on the continent by rhe Aptian-

Albian, at least. Preliminary studics of somewhat

older (Barremian) dinosaurs from Utah

GEODIVERSITAS • 1999 • 21 (2)

209

Cifelli R. L. & Madsen S. K.

(Kirkiand et ai 1997) and ciscwhcre in North

America (Norman 1998) suggcsi a link with chc

pcnecontemporancous or slightly oldcr Wealdcn

assemblage of western F.urope. SpaLicotheriunh

rhe suggested sister-taKon to North American

spalacotheriids, occurs in rhe Late Jurassic or

earliest Creracci)us (Purbeck> sec Clcmcns et al

1979; Allen & Wimbledon 1991) ro Early

Crctaceoüs (Wealden) of England (Clemcns &

Lees 1971) and Spain (Krebs I9S5)> providing

corroborative support for rhe suggestion of

faunal continuity beween North America and

Europe prior to the Aptian-Albian (see, c.g.»

Norman 1998).

The Mussenruchit locil fauna includes the mOvSt

diverse assemblage of symmetrodonts ktiown

from North America, with at lea.st four species

présent, ’l here is no clear temporal irend in

North American spalacorheriid diversity: one

species is knovvn from rhe Apiian-Albi.in

(Patterson 1955, l‘‘*56), one From the

Cenomanian (). G. Eaton, pers. comm.), two

from the ’îlironian (Cifelli 1990), and one each

from two local launas of the early ("ampanian

(Fox 1976; Cifelli Ôi Madsen 1986). In view of

ihc tiny she of most knpwn species, part of diis

may wcll be due to collecting biascs, but ir is

notable that spalàcoiheriids ure not only diverse

but extremely abundaiu in the Mus.sentuchit

local faunu, whcre they vastly outnumber ail

orher mammals except muliituberculates: evi-

dently they were a nithcr successFul group in the

mid-Cretaceous of central Utah. Tnterestingly,

the distribution of species is decldedly nûn-

random in the upper part of the Cedar

Mountain Formation. Of the tliirty-two sites

sampled extensively for microvertebrates, only

eight (Fig. I) yielded remains of Spalaco-

theriidae. The overwhelming majotity of .spéci¬

mens rcfcrablc to Spalacolestes creiulablnita was

recovered from a single, heavily-sampled site,

with fewer numbers from three other sites.

Spalacotheridium tiohlei, though less abundant, is

far more extensive in distribution: it is known

from seven sites, and is quitc rare at ihc major

locality thaï produccd such a wcalih of .spcci-

mens rcfcrablc to S. cyclithihltttut. S. incomirtmis,

on the other hand, is a rare species known from a

single, poorly sampled locality - it is not présent

at the most lieavily sampled site or, for dtat mai-

ter, anywhcrc cIsc. Given chc face char ail sires are

locared in a narrow stratigraphie intcrval and are

belîeved to he essenrially isochronous (Cifelli

et ûL 1997, 1999), we considrr it uniikely that

rhese différences are temporal in nature. Ail of

the sires occur in fluvial overbank dcposics.

Although ihc dcpositional sccting appears to he

raihcr similar berween sites, wc attribute ihc dis¬

tribution of Spalacothcriidac in rhe upper part of

ihc Cedar Mountain Formation as being duc to

différences in habitat preference among species,

with Spnlitcolcstes cretulablana and, particularly,

5, inconcmfiUS’. being characterized by a far gneater

degree of habitat specificin^ than was evidently

ihe case for Spalacotheridium noblei.

Acknowledgements

We acknowledge with hearrfelt thanks the numer-

ous individuals wbo generously suppÜed us with

iinpublishcd information, access ro spécimens,

advice, commenis on un earller version a( ihis

manuscript, or olher help: W. A. Clcniens, Jr.,

Bob Hmry, Susan Evans, Richard C. Fox. Jacques

Gauthier, Jerry Hooker. Jim Flopson. Zofia

Kiel.in-jaworowska, Jay Lillegraven. Zhcxi Luo,

Bill Turnbull, and, most especiully. Denise

Sigogneau-Russell, who pruvided much help in

ail of the categories listcd. h is a plcasure to

thank the Judd fumily of Castic Dale, Utah. for

their cuntinuing and unfailing help to field par¬

ties rhrough the years; to Beth Larson, Raiidy

Nydam. Nick C/.aplewski, and Kent Smith for

their heJp in field work; and to Tom Rasmussen

and rhe Bureau of l.and Management for their

coiitinuing coopération. Scanning électron

micrographs were doue by Cindy Gordon at the

Samuel Roberts Noble Electron Microscopy

Eaboratory, University of Okiahoma, and Randy

Nydam hciped with digital inrivtge processing.

Figures 2, 3, (k and 1 I were drafted by Nick

C/aplewski; Figures 8, 9 were prepared by Glenii

Traughber; and Figure I was drafted by Coral

McCaljistcr. Comparative study and critical

iiucrcaciion with collcagues was greatly facili-

taicd by a Professeur Associe appointnicni for

R. L. Cifelli at ihc Laboratoire de Ralêoniologic,

Muséum national d’Flisroirc naturelle, Pans, and

the support of Drs Philippe Faquet, Philippe

210

GEODIVERSITAS • 1999 • 21 (2)

Spalacotheriid symmetrodonts from the mid-Cretaceous of Utah

Janvier, and Christian de Muizon is acknowled-

ged wiih hearcfeli thanks. This rcscarch was

conducted under the folltjwing grants to RLC:

National Géographie Society 5021 92 and

National Science Foündaiion BSR 8906992,

DEB 9401094, and DEB-9870173.

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Composition en acides aminés d’os de

mammifères fossiles de deux sites du

Plio-Pléistocène d’Angola. Comparaison

avec la conservation de la phase minérale

Hélène DAVID

Laboratoire de Paléontologie, Université Paris-XI, Bât. 504, F-91405 Orsay cedex (France)

Laboratoire d’Anthropologie, Université Bordeaux-I,

avenue des Facultés, F-33405 Talence cedex (France)

Yannicke Dauphin & Pascale GAUTRET

Laboratoire de Paléontologie, EP 1748 du CNRS, Université Paris-XI.

Bât. 504, F-91405 Orsay cedex (France)

Martin PICKFORD

Laboratoire de Paléontologie, UMR 8569 du CNRS, Muséum national d Histoire naturelle.

8 rue de Buffon, F-75231 Paris cedex 05 (France)

Chaire de Paléoanthropologie et de Préhistoire, Collège de France,

11 place Marcelin-Berthelot, F-75005 Paris (France)

Brigitte SENUT

Laboratoire de Paléontologie, UMR 8569 du CNRS, Muséum national d’Histoire naturelle,

8 rue de Buffon, F-75231 Paris cedex 05 (France)

David H-, Dauphin Y., Gautret P.. Pickford M. & Senuî B. 1999. — Composition en acides

aminés d'os de mammifères fossiles de deux sites du Plio-Pléistocène d’Angola.

Comparaison avec la conservation de la phase minérale. Geodiversitas (21) 2 :215-227.

MOTS CLÉS

Plio-Pléistocènc,

Angola,

Theroùithecus^

os,

FTIR,

acides aminés.

RÉSUMÉ

La comparaison des paramètres microsiructuraux et de composition (phases

organique et minérale) d'os de mammifères actuels et fo.ssilcs (Plio-

Plcisiocènc d’Angola) montre le caractère diltérentiel de la diagenèse. Malgré

une conservation microsrructiiralc globalement bonne, les compositions chi¬

miques des phases minérales sont modifiées (enrichissement en Ca, appau¬

vrissement en Mg). Très peu de matrice organique e.st conservée. Les phases

organiques solubles er insolubles sont également diversement uliérécs, la

composition en acides aminés de la phase .soluble semblant mieux conservée

que celle de la phase insoluble.

GEODIVERSITAS « 1999 • 21 {2}

215

David H., Dauphin Y., Gautrer P., Pickford M. &c Senut B.

KEYWORDS

Plio-Pleist(Xcnc.

Angola.

Theropiîhci'us^

Bas,

IxMK-,

l-l'lR,

amino-acids.

ASBTRACT

Composition in amino-acids offossil rnammalian bones f'rorn two Plio-

Pleistocene Angolan sites. Comparison with thepréservation uj the minerai phase.

Compariscm of the microstructural paranicters and composition (organic and

minerai phases) of rntpclern and lossil mammal bones (Plio-Pieiscocenc of

Angola) sliow ihe dilfercntial characier of diagcncsis. [>cspiic cxcellcnl pré¬

servation ni the nncrostrucrurc, rhe chcmical composition of ihe minerai

(ihases has hccn modihed (enriehed in Ca, rcdiiction of Mg). The amount of

preser\'ed organic matrix is greaily roduccd. fhc amino-acid composition of

the sttlubif organic phase appears to be better presctvcd lhan that of rhe-in¬

soluble phase.

INfRODUCTION

L’abondance des phases organiques dans l’os

actuel semble un facteur favorable à leur conser¬

vation chez les fossiles, et les données relatives à

leur composition devraient être très abondantes.

Cependant, de telles informations demeurent

ponctuelles cat, malgré cette circonstance appa¬

remment favorable à des analyses extensives, les

techniques analytiques de la phase organique

nécessitent la déminéralisation de To.s. Or, ceci

implique la destruction du seul paramètre géné¬

ralement étudié par le paléontii>loglste, et consi¬

déré encore uctuellejncni comme le plus, sinon le

seul, réellement important et inlormatif : la mor¬

phologie. Second point qui limite considérable¬

ment la portée des données sur les os fossiles : les

études sont très spécialisées, soit par la technique

utilisée (diffraction X, immunologie...), soit par

le composant choisi (phase odnérale, composi¬

tion de rostéocalciiic...). Bieti que Pos soit un

matériau complexe, rares sont les travaux prenant

simultanément en compte les p.iramètres de la

phase minérale et de la phase organique. Enfin, à

cause de la spécialisation de plus en plus poussée

des laboratoires et de la complexité croissante des

techniques analytiques, la comparaison de la dia-

genèse des phases organique.s et de celle de la

phase minérale demeure un problème rarement

abordé. Les données sur les phases organiques

fossiles sont donc encore réduites, malgré leur

potentiel informatif très vaste (apport à la recons¬

titution de la phylogénie, du paléoenvironne¬

ment, histoire de la fossilisation, compréhension

des processus de formation des sires...). La diver¬

sité des âges et de la géologie des sites fossilifères

est telle que des « lois » régis.vanr la fossilisation et

tous les phénonèmes connexes ne pourront être

établies qu’à partir de très nombreuses données.

Une étude préalable ayant mis en évidence la

présence de sucres dans un asiragale de bovidé

récolté dans un site angolais plio-pléistocène

(David e/ al, 1996), l’analyse de ce spécimen a

été poursuivie afin d’obtenir des données sur la

diagenese comparée des phases minérales et orga¬

nique.s. Lin .site fossilifère voisin, de même âge

mais de scdimencologic différente, a fourni des os

de primates, susceptibles tic fournir des informa¬

tions supplémentaires sur la fossilisation et la dia-

genèse.

TRAVAUX ANTÉRIEURS

I.c.s travaux sur la structure, la minéralogie et la

composition de fus actuel sont trop nombreux

pour être cités. Ires rapitlemcnt, les auteurs ont

abandonné les analyses globales au profit d’une

caractérisation des com|>osant.s protéiques i.solés,

pour en déterminer la composition ou le séquen¬

çage. Paniii les paramètres caractéristiques des

phases organiques, les acides amines semblent

av(jir été parmi les premiers étudiés, notamment

h cause dos particularités de composition du col¬

lagène. Or, cc dernier constitue environ 9(» % de

la phase protéique de l’os. Toutefois, dès 1965,

Glimcher &C Katz mettaient en évidence les pro-

216

GEODIVERSITAS • 1999 • 21 (2)

Composition en acides aminés d’os de mammifères du Plio-Pléistocène angolais

blcmes poses par la solubilisation plus ou moins

grande du collagène dans divers solvants et

acides.

Chez les fossiles, les premières analyses détaillées

semblent dues à Abelson sur du matériel

dévonien. La comparaison de la composition en

acides aminés de spécimens appartenant k des

taxons variés, et venant de sire^s d âges différents,

montre une grande variabilité (Armstrong &C

Halsiead Larlo 1966 ; Dungwortli çt ai 1974 ;

Wyckoff & Davidson 1976 ; Davidson et ai

1978 : v*on Rndr & Orrner 1982 ; Cohcn-Solal

et al. 1987). En réalité, cette variabilité doit être

supérieure a ce qui ressort de rexamen de la littc-

rarure. car seuls les résultats w po.sitils », c'est-à-

dire les analyses dans lesquelles les acides aminés

ont pu être identifiés, sont généralement publiés.

Or, certains sites fossiles fournissent des os conre-

nant encore de la pha.se organique, mais les

spectres d’acides aminés ne sont pas roiijours

ititerprétables (Monigelard et ai 1997 ; Dauphin

1998). De plus, dati.s la plupart de.s cts, seule la

phase organique insoluble (assimilée au collagène

malgré les variations pouvant être dues au pro¬

duit utilisé pour la déminéralisation de l’os), est

prise en compte. Outre le collagène, l’albumine

et rostéocalcine sont les protéines le plus Souvent

identifiées chez les fossiles ou sublossiles (Tuross'

étal. 1980 ; Lowenstein 1981 ; Huqef/y/. 1985 ;

Montgclard 1992). Il s'agir en lait des protéines

les mieux caractérisées dans l’os actuel car il y

aurait au moin.s deux cents protéines non colla-

geniques (ou NCP) (Delmas et ni 1984). L’acide

y-carboxyglutamique, longtemps considéré

comme caractéristique de l'ostcocalcine, a été

recherché chez les fossiles, ibutclois. les huerpre-

tations qui découlent de Sa présence ou de son

absence quant h la « bonne conservation de Los

doivent être pondérées. D'une parc, l’ostéocalci-

ne est parfois en quantité négligeable, comme

dans l'os humain. D’autre part, dams l’os adulte,

routes les protéines non coUageniques sont déjà

fortement dégradées ( Ibrminc 1988), ce qui res¬

treint forcement la probabilité de sa conservation

chez les fossiles. Enfin, Tacidc y-carboxygluta¬

mique a été extrait et identifié dans les squelettes

de coraux, et est probablement également présent

dans les tests de mollusques (Hamilton & Zerner

1983).

MATÉRIEL ET MÉTHODES

Matérif.l

Les spécimens actuels de références sont consti¬

tués par un tibia de bœuf (congelé et sec), un

humérus de Pftpnf anubis (Cercopithecoidea

d’Ouganda, ayant séjourné en forêt pendant une

durée indérerminée) et du collagène commercial

de type I (Sigma).

Les deux sites fossdileres, situés sur le plateau

d'I lumpata (sud de l'Angola) sont datés du Plio'

Pléi.stocènc. Le fragment d’astragale de bovidé

provient des découvertes effectuées lors d'une

campagne de FAngola Palacontology Expédition

(Picklord et.nl. 1992, 1994). Il a été récolté dans

des remplissages de fissures de la carrière de

Cangalongue 111, composées de brèches gros¬

sières, contenant de nombreux fragments de sta-

lagmires recimenrés par des travertins. Les

fragments de cotes et de vertèbres de

lUeroInthecm (primates Cercopirhocoidea) vien¬

nent du gi-semeru de Tchiua, dont les remplis¬

sages sont formés de brèches à grains très fins.

C]ctre fonnarion a été interprétée comme du

guano calcifié de chauve-souris.

Il convient de signaler que la diversité des os ne

peut guère provoquer de biais majeurs dans la

comparaison, compte tenu du niveau d’observa¬

tion utilisé dan.s ces analyses.

MiThodf.S

Microstructures

Des cassures brutes et traitées ont été observées

au microscope électronique à balayage. Les os

actuels ont été soumis a des protcolyscs enzyma¬

tiques, afin d'éliminer partiellement l’abondante

matrice organique qui tend à masquer les struc¬

tures. De la trypsine et de l'alcaluse ont été utili¬

sées car elles sont peu spécifiques. Les surfaces

polies dex os fossiles ont etc li%èremcnt décalci¬

fiées à l'acidc formique.

Cornpoùîton globale

La composition minéralogique et la présence de

phases organiquex ont etc dcicrminccs par spec¬

trométrie infrarouge à transformée de Fourier

(FTIR). Après déconramination des polluants

organiques à Lhypochloritc de sodium, les os

sont rincés à l’eau Milli-Q et séchés à température

GEODIVERSITAS • 1999 • 21 (2)

217

David H., Dauphin Y., Gautret P., PickFord M. & Senut B.

ordinaire. Ils sont en.suite finement broyés. Les

poudres d'os nn^langees à du KBr ont été analy¬

sées sur un spectromccre FLIR Bcrkin

Elmer 1600, équipe d’un accessoire à réflexion

diffuse (DRIFT). Le nombre de balayages est de

soixantc-quane (soit un remps d'analyse supé¬

rieur à quatre minutes par spectre), dan.s une

gamme de longueurs d'onde allanr de 450

à 4000 cm ' (Dauphin 1993). l e système

est maintenu sous uimosplièrc d'a>:oie afin de

réduire les bandes dues aux CO, et H^O atmo¬

sphériques.

Analyse chimirjue élémentaire

La composition chimique a été déterminée par

microanalyse localisée (spectrométrie dispersive

en énergie ou EDS). Le système utilisé, Link An

10000 couplé è un microscope électronique

Philips (SFM 505) (Université Paris-XI-Orsay),

possède un programme .spécialement conçu pour

l'analyse dex surfaces rugucascs. Par une prépara¬

tion adaprée des spécimens, il esc aisé d'identifier

les divers composants d'un fossile (tissu, sédi¬

ment). La prépararion des échantillons, les

conditions d’analyse et le traitement statistique

sont similaires i ceux précedemmenr décrits

(Dauphin 1997). Dans les spécimens actuels, les

teneurs de ccrntiirs éléments chimiques sont infé¬

rieures à la limite de dereenon de la microsonde

(Fe ou Mn p,ir exemple). Toutefoi.s, les valeurs

obtenues sont indiquées, car ces élçmcms sont de

bons indicateurs de la diagenèse, leurs teneurs

dans les fossiles étant souvent supérieures à la

limite de détection de la microsonde.

Compositiofi en acides aminés

Les os ont été décoiitaminés i l’iiypochlurite de

sodium, puis nettoyés pendant quelques minutes

aux ultrasons afin de décoller les particules et

débris divers qui pouvaJent subsister, notamment

sur les fossiles. Apres rinçage à l’eau Milii-Q, ils

ont été séchés à température ambiance.

Les poudres résultant du broyage ont été décalci¬

fiées à l’acide acétique sous un pH constant de 4.

Les phases solubles (MOS) et insolubles (MOI)

ont été séparées pat celUrifugation. La pha.se

soluble a etc de.ssaloc pat ultrafiltration avec de

Peau Milli-Q sur une membrane dont le seuil de

coupure esr de 3 kDa. La phase insoluble a été

dessalée p.u centrifugations successives dans de

Peau Milli-Q. Les phases soluble et insoluble ont

été lyophili.sécs.

Apres hydrolyse dans une solution FfCl 6N pen¬

dant vingt-quatre heures à 1 10 "C sous atmo¬

sphère d\izote, la dérivation PPPC a etc eUcctiiéc

car elle permet la détection des amines secon¬

daires. La composition en acides amines a été

obtenue par chromatographie HPl.C en phase

inverse, sur une colonne Nucleosyl Cl H, avec un

élucni d’acide orthophosphorique et NaOH à

pFI 6,4 et un gradient d'acéionirrile. Le détecteur

est réglé à 254 nm. Rappelons que Phydrolyse

utilisée detruir Jes acides aminés soufrés (cystéine

et méthionine), ainsi que le tryptoplmne. Kaeide

Y carboxygluramique, qui nécessite une hydrolyse

particulière de type alcalin, n'a pas' été recherché,

les matrices organiques exiraires des os fossiles

étant en très faibles quantités.

Points isoclectriqncs

À partir des compositions en acides aminés, on

peut calculer le point Isoélecrrique moyen (pl)

d'une phase organique (Sillero Üc Ribeiro 1989),

ce qui permet d’estimer son degré d’.tcidité, Dans

la formule utilisée par ces auteurs, les acides

aspartique- glutamique, la cysréine libre, la ryro-

sine, Phisridinc, la lysine et Parginine sont pris en

compte. Cerre rnethiule indirecte présente l'avan¬

tage de pouvoir erre appliquée sur le.s matrices

soluble et insoluble, ce qui n'est p;is le cas des

métho<les directes chromaiographique (chromato-

foenssing) ou électropliorétiquc (isoelnrric foms-

sing). plu.s précises car elles fournissent

Pcnsemble des points isoélectriqucs de.s divers

composés. De plus, le calcul à pariir de.s pour¬

centages d’acides aminés permet de connaître le

pl des phases solubles même lorsque les quantités

recueillies sont minimes.

RÉSUUI’ATS

CON'l RO\ ni-, l’flAT DK CONSERVATION

t.l-NEKAI K DLS OS

L’observation de la microsiructure est une pre¬

mière étape dans le contrôle de Pétat de conser¬

vation d’un fossile. Elle est récemment devenue

cruciale, car certains champignons contiennent

218

GEODIVERSITAS • 1999 • 21 (2)

Composition en acides aminés d’os de mammifères du Plio-Pléistocène angolais

Fig. 1 — A, Surface e)(lerne altérée dans une cassure ancienne de l’astragale du bovidé fossile de Cangalongue. Échantillon non

traité : B. vue d un fragment de vertèbre de Theropithtscus, Cercopithécûidea (Tchiua), après un nettoyage insuffisant pour détruire

un éventuel sédiment présent dans les cavités (acide formique 5 %. 15 s) : C, surface altérée d’un fragment de côte de

Theropithecus. Cercopithécoidea montrant l’os spongieux après disparition du périoste : cassure traitée à l acide formique 5 %,

15 s ; D, coupe oblique montrant la pâroi des trabecules darrs l’os spongieux d'un fragment de côte de Theropithecus : même spéci¬

men que Fig. 3 : E, détail de la précédente : F. lamelles osseuses dans une coupe oblique d'un fragment de côte de Theropithecus :

cassure traitée à l'acide formique 5 %, 15 s. Échelle ; A, 160 pm ; B, 3 mm ; C, 90 pm ; D, F. 100 pm ; E, 10 pm.

GEODIVERSITAS • 1999 • 21 (2)

219

David H., Dauphin Y., Gautret P., Pickford M. & Senut B.

Fig. 2. — Spectres infrarouges des os actuels {Bos et Papio, Cercopithecoidea) et fossiles (bovidé et Theropithecus) montrant que

les os fossiles sont conservés en apatile, et la diminution de leur quantité de matière organique.

des acides aminés jiisqu^ici considérés comme

caractéristiques du collagène (hydroxyproline et

hydroxylysine) (Celcrin.e’/1995).

Les sections diversement oricnrcc.s réalisées dans

l’os de bœuf actuel observé au microscope élec¬

tronique iî balayage apres une protéolyse enzyma¬

tique montrent la structure lamellaircj le système

haversicn ci les z.onc.s en « contreplaqué >>• (David

et ai 1996). Macroscopiquemeni ahéré (Fig. lA),

le fragment d'astragale de bovidé de Canga-

longue ne comporte pas de remplissage secon¬

daire dans les cavités naturelles de l’os, ni de trace

indiquant l'aciiviic de micro-organismes (David

étal. 1996). Les risques de contamination en

matrice organique d'origine exogène apparaissent

ainsi réduits. Les cavités des vertèbres de

Theropithecus ne sont pas comblées (Fig. 1 B). .Sur

les fragments de cotes, les alterations de l’os péri-

ostique (Fig. 1 C) permettent d’observer l’os

spongieux .sous-jacent, ainsi que des' structures

lamellaires dans les zones plus internes. Des

lacunc-S ostcocytiques sont parfois présentes. La

disposition des fibres de collagène minéralisées

est conservée sur les parois des trabécules

220

GEODIVERSITAS • 1999 • 21 (2)

Composition en acides amines d’os de mammifères du Plio-Pléistocène angolais

Na Mg Al S Cl K Mn Fe Sr

SÉDIMENT

Cangalongue Tchiua

Na Mg Al S Cl K Mn Fe Sr

Fig. 3. — Teneurs en éléments majeurs : P et Ca, des os Fig. 4. — Teneurs en éléments mineurs des os actuels et fos-

actuels et fossiles, et du sédiment encaissant. siles, et du sédiment encaissant.

(Fig. ID, E). La structure en lamelles est égale¬

ment visible (Fig. IF).

Composition olobai.e ei hémen iaire

Les .spectre.s infrarouges montrent que dans les

deux siles, les os sont en apatite. avec des modifi¬

cations modérées (Fig. 2). Termine ÔC Posner

(1966) ont rnis au point un mode de cilcul de la

crisrallinitc (sp/hnng fiactioii) de Tos à [ïarrir des

intensités relatives des bandes du doublet v4 PO.j.

Le taux moyen de cristaHinirc de l'os atteint 0,10

pour le bœuf actuel et 0,11 pour Pdph ; celui du

bovidé fossile est supérieur à 0,1 4 tandis que chez

Theropithecus il est égal à 0,08. Les variations du

taux de cristallinitc des os actuels dépendent de

l’âge de l'animal : faible chez l'animal jeune, il

sera plus élevé chez un animal âgé. Chez les fos¬

siles, ces variations originelles sont généralement

masquées par les modifications diagénétjqucs.

Que la cristallinité augmente ou diminue, la

aiusc de CCS v-ari-itions reste la plupart du temps

indéterminée : disparition de l’os amorphe, aug¬

mentation de la phase crisralline aux dépens de la

phase amorphe, ou augmentation de la cnstallini-

ré de la phase initialement déjà cristalline.

Slutman et al. (1965) ont montré que la position

et rintensiié des bandes vl, v3 et v4 dépendaient

du type d apatite : hyJroxyapatitc, chloroapatitc

et fluoroapatite. Si F est normalement en quantité

insuffisante dans l’os actuel pour altérer les fré-

GEODIVERSITAS • 19&9 • 21 (2)

221

David H., Dauphin Y., Ciautrer P., Pickford M. & Senut B.

Fig. 5. — Composition en acides aminés (%) des phases insolubles (en haut) et solubles (en bas) des os actuels et fossiles de

Cangalongue et de Tchiua. Les compositions du collagène et des protéines non collagéniques (NCP d'après von Endt & Ortner 1982)

sont figurées.

quences de ces bandes, il est généralement admis

quil est en quantité importante dans tous les os

fossiles. Toutefois, d’après ces critères, les os fos¬

siles ne sont pas enrichis en R

Les différences de composition des sédiments

(l’un calcaire, l’autre phosphaté), n’apparaissent

pas dans les spectres, ce qui confirme l’absence

d’un remplissage important des cavités osseuses.

La phase organique, encore présente chez les fos¬

siles (amides A, I et II), y est modifiée en quan-

222

GEODIVERSITAS • 1999 • 21 (2)

Composition en acides aminés d’os de mammifères du Plio-Pléistocène angolais

NCP Bos Theropithecus

Collagène 1 Bovidé

Fig. 6. — Points isoélectrlques moyens calculés d'après les

compositions en acides aminés par la méthode de Sillero &

Ribeiro 1989.

tiré (bandes moins intenses) et en qualité (absen¬

ce de certaines bandes).

Les os fossiles sont tous deux enrichis en Ca

(Fig. 3). Iheropithecus étant en outre plus riche

en P que le bf)vidc. Le rapport en poids Ca/P de

l'os de bœul actuel esc voisin de 1,9b, celui de

Papio atteint 2.02. Chez les fossiles, ces rapports

sont supérieurs : 2.08 pour Theropithecus et 2,30

pour le bovidé. Les teneurs en éléments mineurs

ne montrent pas de ntodifications importantes.

Malgré la grande différence de quantité en Fc des

deux sédiments (Fig. 4), cette disparité aest pas

transcrire dans les fossiles puisque les teneurs des

os sont similaiccs dans les dcttx sites (Fig, 4). Le

sédiment de Cangalongue est calcaire, et plus

riche en Mg et AJ que Tchiua (Figs 3, 4).

Composition pn acides aminés

Phase organique insoluble (Fig. 5)

La composition de la inarricc insoluble extraite

de Fos de bœuf esc identique à celle du collagène

commercial de référence (type 1), La matrice

insoluble du bovidé fossile est très différente de

celle de Fos actuel, mais la plupart des pics a été

identifiée. Les teneurs en glycine, hydroxy-

proline, prolinc et alanine sont plus faibles que

dans Factuel, les teneurs en tyrosine, valinc et

isoleucine sont plus élevées. Deux pics non iden¬

tifiés sont présents. La composition de la matrice

insoluble de Theropithecus est beaucoup moins

claire : de nombreux pics ne sont pas identi¬

fiables avec ceititude et le spectre n’est pas repré¬

senté dans la Figure 5. Dans la zone correspon¬

dant à la succession thréonine, alanine, histidine

et proline, on rrouv'e seulement un large dôme.

Les deux piçs non identifiés chez le bovidé exis¬

tent également.

Phase organique soluble (Fig. 5)

À titre de comparaison, la composition des pro¬

téines non collagéniques (NCP) est figurée (Von

Endt & Orrner 1982). Peu abondantes (10 % de

la matrice organique de l’os : Hauschka 6c Wians

1989). CCS protéines non collagéniques sont

cependant tres variées. Nombre d'entre clics ne

sont pas encore identifiées, et certaines sont

connues seulement particllcmcnr. La composi¬

tion de la phase srduble extraite de l’os de bœuf

actuel est relativement similaire ,a celle de la

phase insoluble ; cette similitude est partielle¬

ment due a la mise en solution partielle du coILt-

gène (Weiner Ôc Bar-Yo.sef 1990). Par rapport à

la composition des protéines non collagéniques

(NCP), la matrice .soluble du bœuf est trop riche

en hydroxyprolinc et en glycine, trop pauvre en

Icucine et valinc.

La matrice soluble du bovidé fossile se caractérise

par une très forte teneur en phenylalanine et une

baisse sensible des quantités de glycine cc

d’hydro.xyprolinc. Pratiquement tous les pics

sont identifiables. La matrice de Iheropithecus est

également riche en phenylalanine, glycine, alani¬

ne et proline, sans atteindre toutefois des valeurs

comparables à celle du bovidé. Les teneurs en

hydroxyprolinc sont nullcs. En outre, le spectre

est très clair et tous les pics sont identifiables.

POINTS iSOF.UCl RIt.?aKS (pl) MOYENS DES

MAI’KICES

Les matrices organiques de Los. collagènes et

NCP sont acides (Fig. 5). Il en esc de même pour

les matrices extraites de Los de bœuf actuel. Si le

pl moyen de la phase soluble de l'os de bœuf est

.similaire à celui de.s NCP. le pl de la phase inso¬

luble du bœuf est légèrement plus acide que celui

du collagène de type L l.a phase sotublc du bovi¬

dé fossile est devenue un peu plus acide que

Fin.soIuble. Quant à U phase soluble du

Theropithecus, elle atteint un pl supérieur à 6

(Fig, 6).

GEODIVERSITAS • 1999 • 21 (2)

223

David H., Dauphin Y., (.îautret P., Pickford M. & Senut B.

DISCUSSION

Ces deux exemples sont bien évidemment insuf¬

fisants pour en tirer des conclusions définitives

sur les processus de fossilisation des os. D'une

part seuls deux sires sont pris en considération,

d'autre pan les paramètres analysés sont trop peu

nombreux. Cependant, ptnir cliaque composant,

ils sont sulfisanrs pour révéler cenains points

communs et ccrtainc.s dilférenccs, avec les études

publiées.

Composi tion globale

La composition globale des os fossiles des deux

sites de Cangalongue et de Tchitia montre que

les os y sont conservés en apatitc. dbutelois, la

diagenèse rfesi [vas absente [îuisque des para¬

mètres tels que les teneurs en C:i, en R la crisralli-

nité et le.s teneurs globales en matrices

organiques sont altérés, fia composition des

phases organitjues est également altérée. Inès peu

de matrice organique a (>u être extraite des os fos¬

siles des deux sites. Ainsi que le fait remarquer

Glimcher (1993) the uriginiil volume occu~

pied by the ornante nuUtix rnusi hâve heeit repLued

hy new inorganic rmiterinl (aystuts Ufid possihly

(imorpi)ou^ udid phthcs) formed during the pe}'iud

of fossilizutioh iifid ujter the deuth of the tVÙHhd

[...] it ivems unteasorntble to ussume that ihe atom

and ion conttiiuents ifi the minerai phase of the ske-

leton and tooih risuns ofthe fossil specimen ave

those which werc présent ai the timc ofthe animais

death. » L'utilisation abusive des paramètre.s geo-

chimiques pour la reconstitution des paléo-

environnements doit une fois encore être signa¬

lée, que ces paramètres dérivent de la phase

minérale ou de la phase organique. Lune des ten¬

dances actuelles en cc domaine consiste a « rem¬

placer » les données issues de l’analyvSc de la phase

minérale par celle de la phase organique, qui

serait moins sensible à la diagenèse. Or, dans la

plupart des cas, cette matrice organique est très

altérée en qualité et en quantité. Et lorsque sa

quantité semble voisine de celle des os actuels,

elle est au moins partiellement exogène

(Montgclard et al. 1990 Dauphin 1998). Dans

le cas des fossiles d’Angola, seules les analyses de

la composition en acides aminés et les données

qui en découlent (pi moyens) ont été étudiées.

Les interprétations sont donc limitées, car de

nombreux paramètres restent inconnus : masses

moléculaires, composition en acides aminés et pi

de chaque composant par exemple. En fait, si a

priori l’abondance naturelle des phases orga¬

niques dans Tos semble être un facteur favorable

â leur conservation, elle est surtout un des fac¬

teurs principaux de leur destruction. D\ine part,

les cellules contiennent de nombreuses protéases

qui sont libérées à la mort de I animal, contri¬

buant ainsi à la dégradation rapide et à la des¬

truction de la |)hase organique. [3’autrc part, les

bactéries, renconrram un milieu nutritif riche,

sont très actives.

Cl'lNSEKVATlON niTFF.RENT'lELLK DES PHASES

organiques

Dans les spécimens angolais, d’après leurs com¬

positions en acides aminés, les phases organiques

ilisolubic.s sont plus altérées que les phases

solubles. Paradoxalement, les compositions en

acides aminés des phases solubles du bovidé fos¬

sile et du l'hcropithecus sont plus proches de celle

du collagène que Ic.s phases insolubles des nicmcs

spécimens, l-a comparaison avec les données de

la littérature nesi pas [mmedune, notamment à

cause de la diversité des techniques d’an.ilyses

disponibles pour les acides aminés. Il est en cflFcc

pratiqut-nicnt impossible d'obtenir, avec une

icule hydrolyse et une seule dérivatii>n, un

speertv contenvinr tous les acides aminés.

Toutefois, une telle similitude a déjà été signalée

dans des os suhiossilcs (Hedges et al. 1980).

D.ins cctic élude cependant, les amines secon¬

daires' (dont fhvdroxvproline, l'un des cléments

les plus caractéristiques du collagène) ne sont pas

mentionnées, fl est prolvible qu’au cours de la

divtgenèse, les fibres de collagène se fragmentent.

Ce phénomène peur être comparé aux procé¬

dures expérimentales permettant tle couper les

longues fibres en peptides, couramment prati¬

quées sur le collagène de type l. C'es techniques

ont pour but de rendre possible rutilisaiioti des

nicthodo.s classiques d'analyse des phases solubles

(chromarogtaphic liquide* et électropboièse

noîanuncnc) sur une pliasc initiale insoluble

(Rossi et al. 1996).

224

GEODIVERSITAS • 1999 • 21 (2)

Composition en acides aminés d’os de mammifères du Plio-PIéisrocène angolais

On peut noter la similitude entre les bovidés

actuel et fossile, qui tous deux, présentent une

phase soluble très riche en acides aminés caracté¬

ristiques du collagène. Le collagène .solubilisé par

les processus de décalcilication apparaît donc

moins sensible aux altérations diagénétiques que

le collagène mni solubiü.sé. Ces diflérences de

comportemcm pourraient êta- ducs à son degré

d'association avec la phase minérale. On peut

rapprocher ces observations de celle de Masters

(1987), qui avait déjà reconnu la conservation

diiférenrielle des composants de l’os, les NCP

étant les mieux conservées.

RÙlii nu .SÉDIMENT

L^c nombreux làcccurs interviennent pendant la

fossilisation des restes squelettiques ; en outre, les

processus diagénétiques sont permanents encre le

stade de l'enfouissement et la découverte du fos¬

sile. Parmi ces facteurs, la taille des os est ^ consi¬

dérer, Rien qiie les études sur le sujet demeurent

limitées, la composition chimique éicmciuaire de

la zone externe d’un os de grand mammifère est

plus modifiée que ses zones internes (Williams âc

Marlow 1*187 ; Williams &C Potts 1988).

Uinfluence de la composition du sédiment est

dans ce cas évidente, puisque le périmètre externe

des os est enrichi en éléments chimiques très

abondants dans le sédiment environnant.

Toutefois, l’influence du sédiment est variable

selon les sites car bien que le sédiment de

Cangalongiic soit beaucoup [dus riche en Pc que

celui de î'chîua, les os ont à peu près les mêmes

teneurs en Fc. Une étude détailIcc de l’évolution

des teneurs en Fe de l'extérieur vers l’intérieur

des os n a pas été faire. L’évenruel remplissage de

la cavité médullaire pcLir*augmcnrcr les mtjdifiai-

rions diagénétiques.

Enfin, en dépit d’un sédiment dont la composi¬

tion est plus proche de celle de l’os à Tchiua quà

Cangalonguc. le.s phases organiques du Thero-

pithecus %ox\x. plies modifiées que celles du bovidé.

Tl faut noter que les dimensions de.s fragments de

vertèbres et de cÉiles du Theropitbtxu^i éXMwt net¬

tement inféiicures à celles du fragment de bovi¬

dé. L’absence de données détaillées sur les

sédiments et le contexte géologique des deux sites

ne permet pas de faire des comparaisons

détaillées.

Il convient donc de noter que, en dépit d'une

microstrucrufc et d'une minéralogie conservées,

la diagenèse est présente dans ces os fossiles. Il

semble important d'insister sur le fait que les

composés minéraux et organiques d’une pan, et

les phases organiques solubles et insolubles

d'autre parc, ont dc.s réactions différentes aux

processus diagénétiques. Ainsi, dans le cas des

fossiles d’Angola, les données peuvent apparaître

contradictoires : la composition en acides aminés

indique une diagenèse plutôt modérée dans le cas

du bovidé, alors que resdmation de la quantité

de sa phase organique montre une forte diagenè¬

se. Il appâtait donc de plus en plus que l’état de

conservation des phases minérales ou organiques

ne peut être établi à partir de l'analyse d'un seul

type de composé. Et ce d’autant plus que, rappe¬

lons le, certains produits considérés comme

caraCTérisriques de l’os (collagène et acide y-car-

boxyglutamique par exemple) sont maintenanr

connus tl.ms des taxons variés.

La connai.s.sance de la suite d’événements qui

préside à la formation d’un gi.scmcnt de fossiles,

ainsi que la compréhension des altérations diagé-

nétiques, ne pourra erre atteinte que par des

série,s d’analyses progressives, allant des caractères

les plus généraux (composition minéralogique»

présence de phases org,iniques) jusqu’aux plus

détaillés (composition en acides aminés ou en

monosaccharides de chaque protéine par

exemple), Le problème majeur de telles études

est que les fraciittnncmeius successifs nécessaires

à une telle idcntincation impliquent que ces

phases organiques soient conservées en quantité

suffisante dans les os.

Remercjcrnenrs

Les auteurs tiennent à remercier les nombreuses

personnes, en France (Prof. Y. Coppens et

P. Taquet) cc en Angola (Dr S. Aço, Mission

française de coopération à Luanda ; Instituto

Nacional do Patrimonio Cultural ; Museu

Régional de Huila., l.ubango) qui ont permis la

réalisation de la campagne de terrain en Angola

en 1990. Les commentaires détaillés du Prof M.

J. Glimcher (The Childrcn’s Hospitaf Boston)

ainsi que ceux de deux autres rapporteurs ano¬

nymes ont été très utiles pour Pamélioration de

ce manuscrit.

GEODIVERSITAS • 1999 • 21 (2)

225

David H., Dauphin Y.. Gautret P.. Pickford M. &C Senut B.

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GEODIVERSITAS • 1999 • 21 (2)

227

New stem giraffoid ruminants from the early and

middie Miocene of Namibia

Jorge MORALES

Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales,

CSIC, José Gutierrez Abascal 2.

E-28006, Madrid (Spain)

Dolores SORIA

Departamento de Paleobiologia, Museo Nacional de Ciencias Naturales,

CSIC. José Gutierrez Abascal 2,

E-28006, Madrid (Spain)

Martin PICKFORD

Chaire de Paléoanthropologie et de Préhistoire, Collège de France.

11 place Marcelin-Berthelot. F-75005. Paris (France)

Laboratoire de Paléontologie, UMR 8569 du CNRS, Muséum national d’Histoire naturelle

8 rue de Buffon, F-75231 Paris cedex 05 (France)

Morales J.. Soria D. & Pickford M. 1999. — New stem giraffoid ruminants from the early

and middie Miocene of Namibia. Geodiversitas (21)2; 229-253.

KEYWORDS

Ruminantia,

Giraffbidea,

Cl i inacoceratidac,

Miocene,

Namibia.

ABSTRACT

Al rhc early and middie Miocène localitie^ of the Sperrgebiee, Namibia, new

material of climacocerand ruminants bave been collected reccntly. From

Elisabethfeld we describe new material belonging to Propalaeorys austroafri-

canus Stromer, 1926, togeiber with a new genus and species of cHmacocera-

tid Sperrgi'hiciomvryx ivardi n. gen., n. sp., a species witbout frontal

appendages close to Propalaeoryx and ro primitive early Miocene European

ruminants sucb as Andegtvneryx. From the locality of Arrisdrift, wc dcfinc

another new genus and species of climacoccradd witb frontal appendages,

Oraugemeryxhendey n. gcn.. n. sp., characterized by iis compic.x tincd frontal

apophyses. Comparison of the dentition and posrcranial skclecon of ebis

genus and ihose of Spnrgebietoyneiyx suggests a close phylogeneric relation-

ships beiween them.

GEODIVERSITAS • 1999 • 21 (2)

229

Morales J., Soria D. & Pickford M.

MOTS CLÉS

Ruminantia,

Giraftbidea,

Climacoceraiidac,

Miocène,

Namibie.

RÉSUMÉ

Nouveaux ruminants girajfoïdes du Miocène ancien et moyen de Namibie.

Dans les localités du Miocène inférieur et moyen de la Sperrgebier en

Namibie, de nouveaux restes de ruminants C'iimacoceraridae ont été récem¬

ment récoltés. Un nouveau matériel provenant d’EIisabethfeld appartenant à

Propalaeoryx austroafritanus Stromer» 1926 est décrit, ainsi qu'un nouveau

genre et une nouvelle espèce du Climacoceraridae, Sperrgelfietomer)>x wardi

n. gen., n. sp,, espèce ne possédant pas d’appendices froniaux* proche de

Propalaeoryx et des ruminants primitifs du Miocène inférieur européen,

comme Andegameryx, A Arrisdrift, nous décrivons un autre genre et une

autre espece de Cliniacoceratidae avec des appendices frontaux, Orangemeryx

hendeyiy qui sc caractérise par ses apophyses frontales complexes. Les compa¬

raisons entre la dentition et le squelette postcrânien de ce genre et de ceux de

Sperrgehietomeryx suggèrent une proche parenté phylogénétique entre les

deux.

INTRODUCTION

This is rhe second report on the ruminants of the

early and niiddle Miocene of Namibia collected

by the Namibia Palacontology Expédition. The

lirst paper dealt with the small bovid

Namibiomeryx seriuti Morales et ai, 1995. In this

article we describe the giraffoids from the samc

région.

Iwo new gênera of giraffoids rccovcred from the

sites of Elisabethfeld and Arrisdrift in Southern

Namibia (Fig. 1) rcvcal a grcat deal about the

origins of this superfamily of ruminants. The

new climacoceraiid giraffoids [ack frontal apo¬

physes, and occur in ihe early Miocene deposits

at Elisabctbtcld and oiher sites in the northern

part of the Sperrgebict. Sperrgebietumeryx is clo-

sely relatcd lo primitive latc Oligocène European

ruminanr.s such as Andegit}7îeryx Ginsburg et al.y

1994, and lies close to the root of the group

which subscquently dcvelopcd apophyses, the

Giraffoidea. Sperrgebietomeryx occtirs in the same

strata as another sperrgebietomerycine, the genus

Propalaeoryx Stromer> 1926.

Basal middie Miocene deposits at Arrisdrift hâve

yielded abundant remains of a new genus of cli-

macoceratid, OrangemeryXy a climacoceratine

Fig. 1. — Geographica! location of Arrisdrift (early mIddIe

Miocene) and Langental and Elisabethfeld (early Miocene) in

the Sperrgebiet, Southern Namibia.

230

GEODIVERSITAS • 1999 • 21 (2)

Miocene giraftoids from Namibia

with tincd apophyses. Exannination of the skull

and postcranial skclcron oi this genus and diose

of Sperrgebielonicryx suggests chat chc Arrisdrift

species may wcll bc the descendant of the

Elisabcthfëld onc.

l'hc transition from primitive pecoians to climaco-

ccratids with apophyses rhus appears to hâve

occurred in Africa subséquent to colonisation of

this continent by pecorans frorn F.urasia.

GEOLOGICAL SETTING

Ei.isahkthfeld

The early Mioccnc site of Elisabethfeld (Stromer

1926) occupiez onc of a séries of prc-Miocenc

vallcys which osed to drain into chc Atlantic

Océan from the région of the present-day Namib

Sand vSca. As a resuit of a worldwide risc in sea-

levcl during the early Miocene the transient sédi¬

ments in the vallcys stopped moving, and hirther

sédimentation occurred in the drowned valleys.

Ar Elisabethfeld» f]ne-grained red limey silts

accumulatcd in a plain that lay between the

Grillental and one of ics Southern nibutanes

which had eut thiough Proterozoie rocks, l'hcse

red silts, which arc often overprinted with pedo-

génie featurcs, were inciscd and then buried by

green silts, sands and conglomérâtes. These Hu-

viatile beds are ovcriain by a fine-grained palaeo-

dune sequence (Greenman 1966, 1970; Corbetc

1989). Unconlormably overlying the early

Miocene sédiments is a two métré ihick traverti-

ne which Iras invaded the upper portion ol the

aeolianite. Fragments of thi.s travertine hâve been

incorporated into a yoiinger set of aeolianites

which crop ont exrensively m the area, often

filling palaeo-valleys eut into ihc early Miocene

sédiments.

At the base ol onc of the green, pebbly-sand

channci infillings cropping ont as a low diff and

immcdiatcly overlying chc basal red limey silts,

the partial skclcron ol^ a ruminant was observed

by Drs J. Ward and I. Corbett in June, 1993.

The specimen was phorographed and leh />/ situ

for later excavation. In Augusi, 1993, Drs

M. PicDord and B. Senut visited the site with

J. Ward and excavated the skeleton. It was évi¬

dent that at least 1 cm of sédiment had been

removed, principally by sand-blasting, since the

photugraphs had been taken rwo months pre-

viously. A mandibic with the cheektecth in place

in June had eroded so diac only the ventral mar-

gin of the jaw Was lefr in Augusi.

The associated fauna indicatc-s that the

Elisabethfeld skeleton is i>f early Miocene âge.

The sire corrélâtes broadly with the localities of

Songhor and Koru. Kenya, and is thus interpret-

ed to be about 20-21 Ma old (Faunal Set I of

Pickford 1981).

Arrisdrift

The site oi Arrisdrift occurs in a latéral channel

of the Proto-Orange located about 1 km east of

the presem-day channel of the river. Fossiliferous

sédiments lie at an altitude of about 4l m above

mean seadevcl, infilHng a low channci carved

into the Gariep Group of I,ate Protero/oic âge.

Fhe cb.ajinel is filled with a coniplcx cut-and-fill

sequence of sédiments raiiging in grain si/c trom

conglomerares ro clays, the latter representing

clay-drapes deposited during periods when the

Arrisdrift channci was eut off from the main

river. During periods of high water, the channel

would be active, so that numerous scour and ftll

épisodes occurred, and can be seen in superposi¬

tion in the excavation.

During periods of low water Icvel, the channel

was effectively isolated from the main srream and

would bave been a quiet pool of water. ’Fhis

channel lay close to sca-lcvel, as indîcated hy the

présence of serpulid worm tubes in abundancc,

even to the extent of forming serpulid reefs.

Ibday thèse invertebrates livc in brackish water

in estuarine settings. I here can be little doubt

that al the beginning of the Middlc Mioccnc,

somc 17-3-17 Ma ago, sca-lcvel was somc 41 m

above présent day Icvcis.

The site of Arrisdrift, like the carlier ones in the

norchern Sperrgebier, owes its formation to the

world-vvide risc in sea-level that occurred at the

end of the early Miocene, which caused rhe

back-ponding of sédiments widiin ihe Proto-

Orange valley. The difïcrcncc in âges of chc tossÜ

sires in ihc northetn and Southern Sperrgebict

indicate that chc rise in sea-lcvcl was relarively

slow, the highest stand being reached some 2-

3 Ma later than the onset of rising sea-lcvcls.

GEODIVERSITAS • 1999 • 21 (2)

231

Morales J., Soria D. & Pickford M.

Fig. 2. — Atlas of Spengebietomeryx wardin. gen., n. sp. (EF 37'92); A, dorsal view: B. ventral view. Scale bar: 20 mm.

This scénario is confirmée! by the discovery of

early Miocene manimals ar Aucha.s, another

Proto-Orange deposit, in sédiments 32-37 m

above present-day mean sca-level, or .some lU m

lower rhan the deposiis ai Arrî.sdrift.

By the rime that sea-lcvels droppcd again later in

the middlc Mioccnc> the Proto-Orange river had

abandoncd some of ils mcander loops and was

following a Icss sinaons course rowards the coast.

When incision occnrred lollowing lowering ai

base-levcls, the early and niiddie Miocene sédi¬

ments deposiied in many of the abandoned loops

were left high and dry.

rhe fauna associated with Oningemeryx is early

rniddle Miocene in âge, correlaring closcly with

Ruropean zone MN4 (De Rruijn et ûl. 1992)

and with rhe sites of Ruiuk and Maboko in

Kenya, assigned to Raunal Set PIII (Pickford

1981). It is probably about 17.5-17 Ma

(Pickford 1994).

SYSTEMATIC PALEONTOLOGY

Suborder RUMINAN'RIA Scopoii, 1777

Superfamily ClRAl-l’OIDKA Simpson, 1931

Family CUMACOCER/Vl IDAH Hamilcon, 1978

(= ClimacOCERIDAE Hamilton, 1978)

DiagN(')S1S. — Ruminants of medium to large size

charactcriscd by rhe rendency — in relation ro orher

runiinanrs of the saine age - for élongation of (he

hccIn (incliiding ihf atlas) and limbs. Distal epiphysis

nf ihe nieiar.ifsal wirh open gnUy. Dentition wiili a

clear hypsodont tendency. Palaeomery’x fold in lower

molars moderatc or suppressed. hypoconid i.soiated

and lobe ol md simple.

SpERRGEEIETOMKRYCINAE n. subfam.

Type genus. — Spengebietomeryx n. gen.

DlAüNtrsis, — Clîniacoccratidac without crânial pro-

fiihcrancev Cranîum wiih wide fronials, sagiu-il crest

and niichals strongly dcfmed, Dcniirion moderarely

hvpsodont. Premolar sériés elongate. l-o\ver molars

wiih strong styhds and rnodcr.ne paleomeryx fold.

Upper molars with very strong stvIüs and laie union of

the internai lobes with the outer wall.

Sperrgebïetomeryx n. gen,

T^TE SPECIES. — Sperrgebietomeryx mardi n. sp.

Diagnosis. — The same as for the type species.

Sperrgebietomeryx tvardi n. sp.

cf. Strogulognathus sansaniensis V\\\\o\ (Scromer 1926).

232

GEODIVERSITAS • 1999 • 21 (2)

Miocene giraffoids from Namibia

Fig. 3. — Occlusal view of the lower dentition of fossil ruminants trom the Sperrgebiet. Southern Namibia: A, right m3-p2 of

Sperrgebieîomeryx wardir\. gen., n. sp. tEF 37'93) from Elisabethfeld; B, right m3-p2and alveolusof pi, Propalaeoryxaustroafrica-

nus Stromer (EF 3’93). from Elisabethfeld; C, left m3-p2 ol Orangemeryx hendey) (AD 1521). tram Arrisdrift; D. left m2-p3 of

Orangemeryx hendeyi n. gen.. n. sp. (AD 654'94-), (rom Arrisdrift- Scale bar' 20 mm.

Hoi.OT\’PE. — EF 37’93, skull, manclibic and associa-

ted arias, with parts ol ihe verccbral culuinn and hind

limbs (Figs 2, 3A, 4, 5» IDF-H), honsed in the

Muséum of the Gcological Survey of Namibia,

Wiiidhoek.

T\TE EOCALITY AND AC.E. — Elisabethfeld, Namibia.

F'arly Miocene.

ErVMOinGY. — Sperrgi'biets German name for ihc

"foibiddcn teiriton'* on accoiint of rhLs nanie.applicd

to tlie niaimmd Area of .Southern Namibia; rntryx,

Grcck for deer. The specics name lionours geologisi

Dr John Ward who found ihe holotypc.

OlAGNOSI.s, — Medium-sued giraffoids, premoJar

séries long and gracile. Lower p4 wirh simple meta-

conid, directed posteriorly, anlerior wing wiihoiu bifur¬

cation. P2 and p2 iicarly ihe .vainc .si7.c a.s P3 and p3.

Dibt-trtl N'TlvM ijiA(;nosi.s. - Spi^ngehittomeryx

from Propalaeoryx an.'droafricanus'hy iis smaller sîzc,

by the more primitive niorphology of the p4 and P4,

and the loss of p 1 . It differs from Walangania africana

{Whitworth 1958) by irs larger .size, the more gracile

premolars and the more primitive construction ol the

p4 and IM. Ir difTers from Prolibyîherhtm

1961, Giraffîdae and Climacoccratinae by the absence

of craniaJ protubérances which arc présent in the laiter

three groups.

Desckiim ion

The skull (Figs 3-4) is wcll preserved, although

eroded on dic anterior part of chc lefr sidc, and

slighily crushed dor.sovcnrrally. In ventral view

the skull présents a cicarly primitive morpholog}^

comparable u> thaï of Dremolberium (Sigogneau

1968). The auditor)' région is of primitive tj^pe,

with ihe siyloid process locaccd berween the mas-

toid process and the cympanic huila, the larter

boing modctarcly inflaced, while the exrernal

auditoty^ meatus is prominenr and almosr circular.

d'he basioccipiral i.s relarively wide, with .strong

poaterior and anterior tubcrclcs lor muscle inser¬

tions. The width of the insertion zone for the

GEODIVERSITAS • 1999 • 21 (2)

233

Morales J., Soria D. & Pickford M.

Fig. 4. — Sperrgebietomeryx wardi, n. gen., n. sp., holotype skull, Elisabethfeld green sands, northern Sperrgebiet, Namibia, early

Miocene; A. dorsal view; B, ventral view; C, right latéral view. Scale bar: 20 mm.

234

GEODIVËRSITAS ■ 1999 • 21 (2)

Miocene girafFoids from Namibia

Table 1 . — Measurements (lenglh. width, in mm) of the upper dentition ol Sperrgebietomeryx wardi n. gen., n. sp. from Elisabethfeld

(EF 37’93), Propalaeoryx austroafncanus from Elisabethfeld (EF 4‘93) and Orangemeryx hendeyi n- gen., n. sp. from Arrisdrift

S. wardi

r EF37’93 1

EF 4 93

P. austroafncanus

EF200‘93 EF20r93

AD273

O. hendeyi

AD283'94

AD334’95

LMM-PP

68.0

LMM

39.5

38.2

41.5

63.0

LPP

31.0

LM3

13.5

13.0

14.0

21.6

WM3

12.8

12.3

18.2

LM2

13.6

13.5

15.1

21.4

WM2

15.5

15.2

21.9

LM1

13.5

12.2

13.0

21.0

WM1

13.0

12.2

20.0

LP4

10.0

9.5

10.5

11.4

13.1

WP4

10.0

10.5

10.3

15.9

LP3

10.0

10.7

11.5

WP3

9.5

9.2

13.2

LP2

10.4

12.5

14.1

LP2

8.9

11.8

masseter muscles is remarkable, and reveals their

strength. In dorsal view, the width of the frontals,

the strength and heighr of rhe sagittal crest and of

the ntichal crest arc ail notable features of the

skiill. The zygomatic process of the Iromal is very

prominent. The frontals arc relatively widc,

although they appcar wider on account of the

dorsoventral compre.ssion ihat affects the skull.

There is no lacrimal f'ossa, and cherc may hâve

been an ethmoidal fenestra although the préser¬

vation of this part does not allow of certainty in

this matrer. The external occipital protubérance is

very strong and projeers posceriorly. The supra-

occipital has a wcll-marked crest.

Upper Jentîtmi (Ttbie 1, hig. 4B)

Molars with strong parasrylcs and mesostyles; in

the M3 the metastyle is also strong. The internai

Table 2. — Measurements (length. width, in mm) of the lower dentition of Sperrgebietomeryx wardi n. gen., n. sp. from Elisabethfeld

(EF 37'93), cf. Strogulognathus (Stromer 1926): Propalaeoryx austroafricanus Irom Elisabethfeld (EF 3'93) and Langental

(1926-507, holûtype), cf. Strogulognathus sansaniensis from Langental (Stromer 1926) and Orangemeryx hendeyi n, gen . n. sp.

from Arrisdrift.

S. wardi

EF 37’93

cf.Strogulog.

P. austroafncanus

EF3’93 1926-507

O. hendeyi

LMM-PP

75.0

82.0

90.6

95,0-109.0

100.6

LMM

44.0

48,2

52.8

58.0-65.5

62.1

LPP

32.0

34.9

37.0

34.6-45.5

38.6

Lm3

18.5

18.0

21.5

21.2

24,2-29,0

26.7

Wm3

8.0

8.5

9.0

9.5-11.5

10.6

Lm2

13.6

13.0

15.5

17.0-21,5

20.2

Wm2

8.9

9.0

7.9

9.4

10.2-13.0

11.9

Lm1

12.5

14.8

15.2-20.5

16.7

Wm1

7.9

7.0

8.8

9.3-12.0

10.5

Lp4

12.2

12.5

13.0

12.5-15.5

14.5

Wp4

5.7

7.7

6.5

8.0-10.0

8.9

Lp3

10.9

12.0

12.4

11.3-15.1

14.1

Wp3

5.5

6.5

6.0-8.2

7.3

Lp2

9.5

10.2

10.5

9.0-12.5

9.9

Wp2

4.1

5.0

4.3

4.0-5.7

4.8

Lp1

5.0

6.0

GEODIVERSITAS • 1999 • 21 (2)

235

Morales J., Soria D. & Pickford M.

Fig. 5. — A D. Sperrgebieîomeryx wardi n. gen., n. sp., Etisabethfeld green sands, northern Sperrgebiet, Namibia. early Miocene;

A. holotype skull in left latéral view; B-D, right mandible; B. occlusal view of cheek dentition: C, lingual view; D. buccal view;

E-G, Propalaeoryx austroafricanus, Stromer, Elisabethfeld red silts, northern Sperrgebiet, Namibia, early Miocene, right mandible;

E, buccal view: F. lingual view: G, occlusal view. Scale bars: 20 mm.

GEODIVERSITAS • 1999

Mioccne giraffoids from Namibia

lobes fiise late with each oiher, the incerlobular

column is sioall. M2 is largcr than tlic othcr rwo

molars, which arc approxinlaicly the same size.

The prcmolârs arc long, including fhe which

posscsscs a scrong anterior sryle with cinguliim

round the protocone. P3 and P2 aLso possess a

strong anterior style and a well-defined anterior

lobe [close morphologicaily to the prcmolârs of

Bosclaphini (Bovidac)].

ÎAnver denthwii ( Table 2, Fig, 5B-D)

'The stage of wear prevents rïiuch being obseiwed,

es[>ccia]ly in the mulars. The third lobe of m3 is

relativcly small. T hc prcmolârs arc long and gra¬

cile. 1 hc close morphologiçal similariry berween

the premolars is notable, cspccially between p3

and p4. Both prcmolârs bave simple metaconids

dircctcd posreriorly and the anterior wing wirhout

bifurcation, while the exrernal incision is modcratc.

Discussion

d1ïe existence of two ruminant spccies in the

early Mioccne of the northern pan of the

Spcrrgebict was noted bv Stromer (I926) on the

basis of fossils from Langcnial and Elisaberhfeld

assigned by him lo cf. Strogidognathm samanien-

sis Filhol, IS70 and Propal^wryx nustroafiiavius

Stromer, I926, respectively. The dentitions assign¬

ed to Strogidognathiis vvere slighily smuilcr ihan

those assigned ro Propnlneoryx austroalriccttnts^

biir were otherwise similar to ihem, which is why

they hâve subseqiienrly been pooled with those

of P. atdstroafriawus (Hamilton Van Couvering

1976).

Genus Propalaeoryx Snomtu 1926

Tyi'K Sl’HCiHS. — Propalacoryx austroafricamis Stromer,

1926.

Holcti ypi:. — 1926-507, mandiblc.

Type ixicali py, — Elisabcthfcld, Namibia.

New CO) na riOKS^ — FJisabcthtdd, Namibia: EF

3’93, riglu inandibic (Table 2, Fig. 5E-G); EF 4’94,

fragment ol right maxilla with damaged P4-M3; EF'

200^93 righi P4; EF 200r93 lefi P2 ( l'able 1).

Description

The mandiblc has lost the ascending ramus and

the symphyscal portion. The horizontal ramus is

robusr and préservés the alveolus for pi which is

iiniradiciilate The lower molars possess a mode-

rate paieomeryx fold, most marked in the mF

The metastylid is sirong and isolatcd. ’Fhc pos-

terior wing of tire hypoconid i.s wcll scparaicd

from the encoconid. d'Iie basal pillar is of mode-

rare size. The hypoconuhd of the m3 is simple

and of moderate size. 1*he p4 présents a bifurcate

anterior wing and complex metaconid positioned

in front of the protocomd and fbrming an inci-

pient internai wall. There is a deep vernc.il inci¬

sion in the poscerior part of the external wall.

The p3 is much sirnpier. It also has a biFtircate

anterior wing, but tlie metaconid Ls a simple crest

directed backwards. 'Fhe p2 is smallcr than p3

with a simple anterior wing. The dimensions (in

mrn) of the spccimcn arc as follows: molar séries

48.2; premolar séries (without ihc pl) 34.9: m3

2F5x:8.5;m2 13x7.9; ml 12.5 x 7; p4 12.5 x

7.7; p3 12 X 6.5; p2 ! 0 2 x 5; p I (alveolus) 5x2,

The maxilla fragment is hadiy abraded so tirât

rhe internai lobes of the fi.uir teeth P4-M3 hâve

been parriy desrroyed, so tliat only the external

lengrh of rhe recrli can bc measured, as lollows

(in mm) : M3 14; M2 154: Ml 13; P4 10.5. In

the ihree molars rlic parastylc and mcso.stylc arc

well-dcvelopcd and cxtcmally wclFdcfincd. In

the M3 tlierc is, in addition, a strong merasryle

which is also wcll defined cxternally and which is

Linitcd by a basal cingiilum to the other styles.

The P4 is elongated with strong parastyle and

metastyle.

The isol.ned P4 (11.4 x 10.3 mm) has an exter-

nal wall similar to rhat in rhe specimen described

above. The protocone is surrounded by a rclaii-

vely strong cingulum.

The isolatcd P2 (Icngth 12.5 mm) is missing its

protocone. Fhe parastyle is large and globular.

The paracone is well marked exrernally and i.s

joined to the parastyle by a smooih cingulum

which continues to the posterior margin.

D1SCU.SSION

The Elisaberhfeld Propidaeoryx mandible differs

from that of Sperrgebîetomeryx by its larger size,

the presence of a well-developed pl and premo-

lars which are more robust and complex. The

p4s are particularly different, those of

GEODIVERSITAS • 1999 • 21 (2)

237

Morales J., Soria D. & Pickford M.

Propalaeoryx Stromer, U)26, pos.scsMtïg ;i hifur-

cate anrerior vving, a strong meraconid vvhich

forms an incipieni lingual walK and a deep exter-

nal incision. In ail these characters this specimen

is close to the holotypç mandiblc of Propalaeoryx

austroiffiicantis, although tlic latrcr is slighrly

larger and ha.s a simplcr mctaconid in its p4. The

new jaw f'rom Klisabethfcid has hypsodont

molars and, as in the hoJorvpe, rhe palaeomeryx

fold is weak, being strongesf in ihe m 1.

Table 3. — Measurements (in mm) of the postcranial skeleton of Sperrgebietorneryx wardi n. gen., n. sp. Irom Eiisabethfeld and

Propalaeoryx austroafricanus from Eiisabethfeld and Langental (Stromer 1926). Abbreviations: APO. antero-poslerior diameter;

TD, transversal diameter; c.c., corpus calcanei; t.c.. tuber calcanei: m., maleolus: s., sustentaculum.

S. wardi

Stromer, 1926

P. austroafricanus

RADIUS

EF2r94

EF 23’94

EF24’94

EF41’94

Length

214.0

Proximal APD

15.9

16.0

17.7

Proximal TD

Distal APD

27.9

19.4

28.0

31.5

Distal TD

25.6

25.1

S, wardi

P. austroafricanus

S. wardi

Stromer, 1926

HUMERUS

EF 36'94

EF 22’93

SCAPHOID

EF 23’94

10a

lop

lOy

Distal APD

28.0

30.0

Antehor Height

13.1

13.0

12.5

14.8

Distal TD

28.6

34.1

APD

17.2

18.5

19.0

20.5

S. wardi

Stromer, 1926

S. wardi

Stromer, 1926

SEMILUNAR

EF 23*94

FEMUR

EF 37’93

Anterior Height

12.4

12.0

Proximal TD

50.0

52.0

Proximal APD

16.3

Head APD

22.1

20.0

Proximal TD

13.5

14.5

Head TD

28.1

27.0

S. wardi

Stromer, 1926

METACARPAL

EF35'93 EF23’94

EF24’94

TALUS

EF 37’93

18a

18b-p

Length

207.2

205.0

Latéral Length

31.4

31.0

35.5

Proximal APD

17.3

17.8

Médial Length

29-9

Proximal TD

21.4 22.1

Latéral APD

18.3

Distal TD

23.8

Distal TD

20.2

19.0

21.2

S. wardi

P. austroafricanus

S. wardi

Stromer,1926

CALCANEUM

EF 37-93

EF 36’93

TIBIA

EF 37'93

16a

16P

Length

70.9

Length

257.0

c.c. Length

49.0

50.3

Proximal APD

46.0

t.c. APD

18.4

20.2

Midshatt APD

17-8

t.c. TD

17.8

18.4

MidshaftTD

20.7

c.c. APD

18.9

18.3

Distal APD

23.5

24.0

c.c. TD

8.6

9.9

Distal TD

29,0

31.0

30.0

m. APD

25.9

s. TD

21.3

238

GEODIVERSITAS • 1999 • 21 (2)

Mioccne giralToids from Namibia

S.wardi P.austroafricanus Stromer, 1926

1PHALANX

EF 24’94

EF 27’94

EF 28'94

13a

Length

34.7

39.8

43.5

42.0

Proximal APD

13.7

15.2

16.2

Proximal TD

12.2

14.3

13.9

13.0

Distal APD

9.4

9.5

9.9

Distal TD

10.1

12.1

11.5

S.wardi

Stromer, 1926

Il PHALANX

EF 24'94

13p

Length

20.4

21.5-25.0

Proximal APD

12.0

Proximal TD

10.2

Distal APD

10.7

Distal TD

7.9

7.5-9.0

The upper dentiiion of Propalaeofyx austroûfi'ica-

nus was hitherto unknown, and thc attribution

of the new Elisabcthfeld üpccimcn to this species

is based on its discoverv locus, its size (largcr

than Sperrgebietomeryx) and its more robusc pre-

molars.

POSTCRANIAL SKELETON OF Sperrgebietomeryx

AND Propaheoryx

Vertébral column

Articiilatcd wilh thc holotype skull of Sperrge¬

bietomeryx wardi therc was thc atlas (Fig. 2) and

rhrec cervical vertebrae (the axis, V3 and V4).

The arias is notable for irs élongation, being

almost as long as it is wide, and in this respect

resemblcs the atlas ol scvcral antclopes such as

Gazella dama Fallas, 1766. 'l‘he margtns oi the

wings are virtually parallcl and srraight. The axis

and the other vertebrae are poorly preserved,

only V3 being complété and revealing that it too

was elongated.

Lirnb bones

Elisabethtcld has yieldcd some thirty ruminant

limb bone-s, in addition ro the partial skeleton

found wilh the holotype skull and mandiblc of

Sperrgebietomeryx wardi. Many of the specimens

liavc been sand^blastcd and some arc broken, so

JC is often difficult to assign rhem raxonomicully.

Bccausc of thc uncertainty in idcntifying to

which species the bones belong we describe the

S.wardi

III PHALANX EF 24’94 EF 34’94

Plantar Length 22.3

Dorsocaudal D 17.4

Dorsoptantar D 13.7

specimens together. but .suspect that the larger

specimens belong to Propalaeoryx austroajricanus

Stromer, 1926, wTile the others probably repre-

sent S. Witrdi (Table 3).

Specimens £E 22’93, a distal humérus

(Fig. lOE), EF 4r94, a proximal radio-ulna,

EF 36’93, thc body of a calcanéum, and

EF 28^94, a firsr phalanx, atc assigned provision-

ally to P ûtistroafrimnus.

The following specimens are assigned to

S. wardi'. EF 36’94 and EF 22’94, dist;jl humerai

epiphyses; EF 21 94, a complété radius;

EF 23'94, arriculared juvénile radial epiphysis,

carpus and proximal end of mctacaipal;

EF 24^94, articuluted radius, mctacaipal and

phalanges: EF 35’93, complété metacarpal;

EF 37’93. proximal half of a fémur, tibia, talus,

calcanéum found with the holotype skull, man-

dible and atlas; EF 27 94, First phalanx;

EF 34*94, third phalanx.

Humérus

S. wardi has the radial fossa limitcd by a tubero-

sity which rcaches the latéral épicondyle, and is

very large because the capitulum has a moderate

vertical development and proximally dues not

ascend grcatly, w'hile disrally it stays ai ihe same

level as the trochlear groove. Fhe humérus of

P. austroafricaniis is similar but il is sligbtiytvider,

bas a médial condyle which is less well-developed

proximodistally, and has very strong relief in the

médial épicondyle

GEODIVERSITAS « 1999 • 21 (2)

239

Morales J., Soria D. & Pickford M.

Fig. 6. — Orangemeryx hendeyi n. gen., n. sp., frontal apophyses from Arrisdrift, Southern Sperrgebiet, Namibia, base of the middie

Miocene; A-C, holotype frontal with base of left apophysis (AD 595'94); A. médial view: B, frontal view; C. latéral view; D, E, lower

part of apophysis latéral views (AD 130); F, mid-part of apophysis with base of bifurcation towards top of frame (AD 131). Scale bars:

20 mm.

GEODIVERSITAS • 1999

Miocene giraffoids from Namibia

Radius

The proximal epiphysis is the samc in the two

species, but in P. austroafiicanus it is slightly big-

ger and ihe latéral tuberosiry is scronger. The dia-

physis in EF 2r94 (Fig. lOG) Ls gracile, and the

articulai' Faccts of ihe distal epiphysis in ail the

spécimens bave ihe t}'pical morphology of early

Miocene ruminants: thac of rhe pyramidal is

small and horizontal, whereas thaï for ihe sca-

phoid is largcr and more elevated than rhar of

the semilunar and tlierc us a platform for the ulna

contact.

Carpus

d'he matcrial is articulated making it difficult to

observe ail the morphological détails (Fig. lOF).

The semilunar is subquadrangiilar and the distal

lacerai facer is miich widcr than rhe médial one,

aiso irs morphology and measurements are iden-

tical to those of cf. Strngulogruiihm saHsantensis

Filhol, 1870, cited bv Stromer (1926, pl. 40,

fig.4).

Metacarpal

This bonc is long and gracile (Fig. lOH), the dia-

physis having a flat posterior surface, and the

proximal extrernity being narrow with respect to

the anrero'posierior diameter. l'hc facct for the

magnotrapezoid is very large compared to that

for the unciform. The distal pulleys bave wcll-

devcloped keels posterioily.

Femur

The proximal half is preserved but both trochan¬

ters arc broken. On the posterior surface, below

the Icsser trochanter, thcrc is a roughencd triangu-

lar area dclimiccd by two crests which continue

parallel to the length of the diaphysis as in the

extant giraffid species Okapia johnstoni (Sclater

1901).^

Tibia

This bone is also long, gracile and straight. The

tibial crest is long, rcaching to mid-shaft of the

diaphysis. On che posterior surface next to the

popliteal line there is another line parallel to it

and somewhat shorter, also as in Okapia. The

médial distal groove is wide and shallow. In distal

view the anterior and posterior margins of the

epiphysis are markcdly concave and the wall

which séparâtes the trochlear facets is short antero-

posreriorly.

Astrapdus

Corresporiding lo the morphology of the distal

tibia, lhe dépréssion hciwecn the two condylcs ol

the proximal end Ls deep and Ls also asymmetri¬

cal. In the médial condyle thcrc is a strong postc-

rior process, and on the anterior surface on both

sides of the fossa there are well-developed stop

facets. On the distal end, the latéral condyle Ls

more extensive than the médial one.

Calcanéum

The calcmeum Spengtdmtorneryy; wardi has a

symmetrical tuber with a wide but short and

deep postL'riiir fossa. Its hody is straight, the dis¬

tal latéral groove is smooth and tlic distal facct

for the astragalus îs oblique. In the calcanéum ol

P. austtoafricitnus the tuber is asymmetric and

more strongly dcvelopcd, both transversely and

antero-posteriorly. The rtigose postero-medial

area is also more dcvelopcd.

Metatarsal

rhe only spccimcn in the collection is a distal

juvénile fragment (EF 25'94) that shows an open

anterior groove. Its attribution to Sperrge-

bietorneryx wardi is hased on its size.

Subfamily Ci.IMAcXjCHRaTINAF. Hamilton, 1978

Diag.MO.SIn. — Glimacoceracidae with frontal protu¬

bérances of apophyseal nature (Buhenik 1990).

Dentition hypsodont. Premolar row shortened. Lower

molars without palaeomctyx lold. Upper molars with

external fusion ol the lingual and buccal lobes.

Orangerneryx n. gen.

Type srntaES. — Orangerneryx hendeyi n. sp.

Diac.NOSLS. — As for the type species.

Orangerneryx hendeyi n. sp.

Clhnacoceras sp. — i Icndey 1978.

Holotype. — AD 595’94, left frontal fragment with

GEODIVERSITAS • 1999 • 21 (2)

241

Morales J., Soria D. & Pickford M,

Fig. 7. — Orangemeryx hendeyi n. gen.. n. sp., frontal apophyses from Arrisdrift, Southern Sperrgebiet, Namibia, base of the middie

Miocene; A, apophyseal point (AD 1798); B, apophyseal point (AD 1177); C-F, apophysis with trifurcate tip (AD 594’94);

C*E, various views; F, dorsal view; G, H, bifurcate apophyseal tip, latéral views (AD 649 + 763). Scale bars: 20 mm.

GEODIVERSITAS • 1999 • 21 (2)

Miocene giraffoids from Namibia

apophysis, housed ar the Geological Survey of

Namibia, Windhoek (Fig. 6A-C').

LocALITV and AGE. — Arrisdrift (southern

Sperrgebiet, Namibia), early middle Miocene,

approTcimately équivalent co rnainmal zone MN4 of

ibc Fairopean scnle (De Bniiin et ul. 19^2). fickford

( 1994) csrimaies ihe age of tlie siic lo be ca 17A Ma.

EtyMv>EOGY. — For the Orange River vvhicli is rhe

border berween Namibia and Soinh Africa and nunyx

rhe Oreek word for deci. The ppccies i.s dcdicaied to

palaeontologist Dr Q. B. Uendey.

DlAG.Mfisis. — Climacoceratinae wrth elongated

slightly compressed rruncate conical supraorbital apo¬

physes, ornaniented at the base svith rounded

tubcrcles with bifurcated or trifurcaied upper Lermina-

rion (cwo or rhree poinrs).

Du FERENTIAI. DIAGNOSJS. - diffcrs

markedly From rhe ocher two généra of cÜmacocera-

tines by the niorpKology of tts apophyses, which are

short with u widc base which diniinishe.s towards the

apex, giving the apophysi.s an elongated. slightly conv

pressed truncated conical aspect, different from the

cylindrical form thaï occurs in Nyanz(tmetyx Thom^LS.

1984 and Clim/tcoceras Médîmes, 1936.

Description

Holotype; The frontal bone is very thick and

strongly vasculariscd but not enougli to be called

pneumatised. The supraorbical loramen is well-

defined and externally continues to the apophy-

sis following a smooih canal, d'he postcornual

fo.ssa is deep and continues postcriorly as a wîde

but not very deep, wcll-demarcatcd groove. The

apophysis is in the forn) of an elongated com¬

pressed conc, with rhe base wide and dlmin-

ishing in section towards rhe apex, while the base

is compressed transversally (anteroposterujr dia-

merer = 44.5 mm, transversal diameter =

29.0 mm), whereas towards the apex the section

is almost circulât. There is a smooth anrerior

keel, accompanied by a small protubérance, rhe

posterior margin is rounded with a weaker but

more extensive proiuberance. The apex is bro-

ken, so char it is possible lo observe chat ihe wall

of dic apophysis is thick and the central part vas-

cularised (Fig. 8H).

Orher specimens such as AD 130 (Fig. 6D-E),

AD 250, AD 483, AD 596 94 (Fig. 8A-E) and

AD 132 (Fig. 8F'C), are similar to the holotype,

even though thcrc is a certain amouni of variabi-

liry in che shape and size of ihe protubérances uf

the apophyses. At the base of the aptïphysi.s in

AD 596 94 (Eig. 6A-E), there are shallow

siniious canals indicating die courses followed by

blood Vcssels. which suggest diat the apophyses

were covered in skin. AD 131 (Fig. 6F) is the

apical and mediul pan of an ajiopliy.sis which

possesses a latéral protubérance similar lo those

menrioncd above. l'his spccimen indicates rbai

the apophysis bifurcates towards the apex, also

evideneed by the surface ornamentation in the

form of a Y. Other specimens indicaie a more

complcx apex than this fossil, one of which

AD 648+763 (Tig. 7G), illusirated by Hendey

(1978) shows a rip wuh two different si/.ed

points, while atiother AD 594 94 {Fig. 7C-F) is

crifurcâie with ihree approximately .similar si/.ed

points. Fragments uf apophyses such as AD 129

(figured by Hendev 1978) or AD 1798 (Fig. 7A),

AD 1177 (Fig. ■"B), AD 658'94 and AD 659'94,

are probably best interprétée! as poinrs at rhe apex

of rhe apophysis.

A specimen of skull (AD 652'94, Fig. 9B-D)

comprises the famcal with the base of the apo¬

physis and part of the pariétal back to the union

of the temporal lincs, wliere they begin to form

the sagittal crest. T hcrc is a scrong, abrupt change

in slope bccwccn the pariétal and the frontal,

whiie berween and anterior to ihc apophyses

chcre is a deep, rounded dépression. In rhe base

of the right apophysis thcrc is a deep. strong

postcornual fossa similar to that obsci*vcd in the

holotype, which continues disrally as a canal ro

connecl with the temporal line. The temporal

lincs arc wcll-markcd, swclling towards the hase

of the apophyses, and postcriorly the)' unité to

form a visible thickcriitig, which fn the broken

surface has a subrriangular section. There was

probably a strong development of rhe nuclial

cresr. T he Iromals arc very thick atid the bone is

wcll A^ascularised, in particular the large supraor-

bitaJ foramen which e\pand.s into the roof ol tlic

orbit. The bases of the apophyses are not pneu-

matized, and in rhe right onc the long widc volu-

minous base can be observed, while in chc Icft

one, which is more broken, the subrriangular

transversal section can be seen to be similar

to that in AD 596’94. This cranial fragment is

GEODIVERSITAS • 1999 • 21 (2)

243

Morales J., Soria D. & Pickford M.

Fig. 8. — Orangemeryx hendeyi n. gen., n. sp., frontal apophyses from Arrisdrift, Southern Sperrgebiet, Namibia, base of the middie

Miocene: A-E. left apophysis (AD 596’94); A, médial view; B, posterior view; C, latéral view; D, anterior view; E, transversal cross

section: F, G. base of apophysis, two views (AD 132). Scale bars: 20 mm.

GEODIVERSITAS • 1999 • 21 (2)

Miocene giraffoids from Namibia

Fig. 9. — Orangemeryx hendeyi n. gen.. n. sp.. Arrisdrift. Southern Sperrgebiet, Namibia, base of the middie Miocene; A, left D4-M1

in occlusal view (AD 624); B-D. cranial fragment (AD 652'94); B, dorsal view; C, ventral vtew; D, right latéral view; E, F, left P4-M3

{AD-273); E. occlusal view; F, buccal view. Scale bars: 20 mm.

GEODIVERSITAS • 1999

Morales J., Soria D. & Pickford M.

Fig. 10. — A D. Orangemeryx hendeyi r) gen., n. sp.. Arrisdnft, Southern Sperrgebiet. Namibia. base of the middie Miocene;

A. B. axis (AD 1549): cervical vertebra V3 {AD 1447); A, latéral view B dorsal view; C, letî radius, anierior view (AD 00 95): D, left

meîatarsal. anterior view (AD 5'93)‘, E. Propalaeoryx austroafrica^us. Slromer, Elisabethfeid red silîs. northern Sperrgebiet, Namibia,

early Miocene. left distal humérus, antehor view (EF 22'g3)i F-H. Spengebietomeryx wardio. gen., n. sp., Elisabethfeid green sands,

northern Sperrgebiet, Namibia. early Miocene; F, left juvénile distal radius, carpus. and proximal metacarpal (In analomical connec¬

tion), antehor view (EF 23'94); G, right radius, antehor view (EF 21'94); H, left metacarpal, posiehor view (EF 35’93). Scale bars:

20 mm.

I 246

GEODIVERSITAS • 1999 • 21 ( 2 )

Miocene girafFoids from Namibia

usefui in providing cvidence as co rhc correct

orientation of rhc apophyses in OrangemeryXy

especially rhe holoiype AD 595'94, in which it

was inclined slightiy laterally and torwards.

Dentition

rhe cheek lecth are relatively hypsodont, being

similar in many rcsj^cccs lo thc dentition of other

climacoceratids. Lower inolars (Table 2, Fig. 3C)

possess strong metascylids which project laterally.

The protoconid and hypoconid are flattened. The

hypoconid and protoconid are separated, only

unking when wear is adranced. The hypoconulid

of m3 is simple ;Uîd unicuspidate. The prcniolars

are short. The |>4 is variable, alvvays with rhe ante-

rior wing bifurcaie and a sirong labial groove, the

metaconid varies from being isolaccd to forming a

Wall which unités wiih the metastylid. Incisiform

tecth are wcll represented in the collection» none

of which are bilobcd. Upper molars (Table I.

Fig. 9A, E. F) ha\Te internai lobes separated from

cach other. ’Fhc styles are strong, particnlarly the

parastyle and metasryle. The entosryle is weak.

Upper preniolars are short and wide.

Vertébral column

Numerous vcriebrae of Ürangerneryx hendeyi are

preserved, somc of them in articulation, Ail in ail

they possess morphological fearures typical of

modem ruminants. Aniong rhe cer\'ical vcrcebrae

there is an axis (AD 1549, Fig. lOA-R) which is

relatively complété. It is notahly elongated and

thc spinous process is distinerly high and well-

dcvclopcd, whtch suggests thc existence of strong

musculature relaied to mcwenieius of ihe head. A

furrher wcl!-prescrved cervical veriebia

(AD 1447. Fig. lOA. B), like thc previous spéci¬

men, is elongated.

Limh bones

The sample of limb bones assigned to Orange-

meryx hendeyi is very compréhensive, bones of

this spccics being rhc most common mammal

rcmains at the site (over 220 specimens). In the

sample are complété examples of most limb élé¬

ments, but some of them bave been deformed by

compaction and several hâve bêen damaged by

gypsum crystal growth (Table 4, Fig. lOC, D).

The morphology of the limbs is quite generalized,

excepr for their élongation, indicating that

O. hendeyi was not greatly speciaÜzed, retaining

much of rhc aspect of what we consider to be the

basal girartdid condition. Many of the postcranial

éléments of thèse ruminants possess a rarher

constant morphological pattern, whereas others

présent major variabilicy and appcar to be more

closely implicated in the processes of adaptarion

and évolution, even though it is difflcult to déter¬

mine rheir evolutionary signifîcjnce. The exis¬

tence of great wiriability in rhe hone size is aiso

noriceable, wliich should probably be interpreted

as a dtmorphic pattern. The aim ol ihi.s section is

not to provide a detailed study of each postcranial

élément, but to provide remarks on rhe salient

fearures of thc limb skeleion of O. hendeyi.

Humérus

In rhe proximal epiphysis (Fig. IlA), the greater

and lesser tuberosities arc low and there is no

intermediaie ruberclc équivalent to the condition

in more modem giraffes. The distal epiphysis is

comparable to that of Sperrgebietorneryx mardi

described ahove, but ihe radial fossa is more

reduced duc to the grcai proximo-distnl develop¬

ment of the capitulum and the mcdial condylc of

the trochlea, a condition which also differentiates

il from CUmacoceras gentryi and Palaeotragus prT

maevtiSy the humeri of which possess a more

'*giraffid”-likc morphology with rhe médial

condyle low proximally.

liadwulna

The proximal epiphysis of the radius accords

Vv'ith rhe morpholog)' of the distal humérus, and

this bone thus differs in the samc way from its

counterpart in P. primü€vus\ the lacerai facet is

higher and the mcdial one mote inclined in

O. hendeyi. The distal radial epiphysis is similar

to that described in 5. mardis the diaphysis is

quite straight and is antcro-posteriorly compres-

sed. The uina (Fig. 1IB-C’) is not fused to the

radius. les tuber olecrani possesses high latéral

and médial crests, between which is a deep valley,

similar to a specimen of Climavoceras gentryi

Hamilron, 1978, from Ngorora. Kenya.

Carpus

The scaphoid retains the distal posterior facet

GEODIVERSITAS • 1999 • 21 (2)

247

Morales J., Soria D. & Pickford M,

Fig. 11. — Orangemeryx hendeyi n. gen.. n. sp.. Arrisdrift, Southern Sperrgebiet, Namibia, base of the middie Miocene; A. left proxi¬

mal humérus, proximal view (AD 3380); B-C. righl proximal uina (AD 00’95); B. anterior view; C. latéral view: D.right distal humérus,

anterior view (AD 1915) ; E. right calcanéum, anterior view (AO 747); F. right talus, anterior view (AD 613'94); G, right navicular

cuboid, posterior view (AD 317’95); H, I Phalanx, anterior view (AD 501); I, Il Phalanx, anterior view (AD 469); J. III Phalanx, latéral

view (AD 896). Scale bars; 20 mm.

GEODIVERSITAS • 1999 • 21 (2)

Miocene girafFoids from Namibia

Table 4. — Measurements (in mm) of the postcranial skeleton of Orangemeryx hendeyi n. gen., n. sp. from Arrisdrift. Abbreviations:

APD, antero-posterior diameter; TD. transversal diameter; c.c., corpus calcanei; t.c., tuber calcanei: m., maleolus: s., sustenta-

culum.

Humérus

t.c. TD

20.1-23.2

21.6

Length

209.5

m. APD

32.0-39.5

36.2

Proximal APD

64.0-71.2

68.0

s. TD

26.6-30.2

28.2

Proximal TD

54.8

Distal APD

41.8-48.6

45.2

Navicuiar-cuboid

Distal TD

44.6-51.8

47.5

Anterior Height

19.4-25.5

21.7

Posterior Height

25.1-29.0

27.2

Radius

Maximal APD

34.2-40.0

36.2

Lenglh

254.0-287.0

268.0

Maximal TD

31.4-36.6

33.6

Proximal APD

20.4-26.8

23.8

Proximal TD

37 0-46-8

42.1

Metatarsai

Midshafl APD

14.1-22.3

18.1

Lenglh

252.0-280.0

266.0

Midshaft TD

23.0-33.4

28.2

Proximal APD

29.0-40.0

32.1

Distal APD

23.7-32,9

28.6

Proximal TD

26.2-33.0

28.7

Distal TD

34.0-46.5

39.5

Distal APD

19.2-25.3

22.0

^jlng

Distal TD

29.1-37.5

33.2

Olecranon Length

49.7-61.0

54.6

Tuber o. APD

30.1-37.4

33.6

Length

46.5-57.4

51.6

Tuber o. TD

15.2

Proximal APD

18.9-22.6

21.0

Pro. anc. APD

32.6-45.6

39.0

Proximal TD

15.1-19.3

17.3

Proc. anc. TD

11.3-14.4

12.7

Distal APD

10.5-15.0

13.3

Distal TD

12.5-15.8

14.5

Scaphoid

Anterior Height

17.4-21.6

19.5

1 Phfll;4nv

APD

24.3-28.5

26.3

Length

26.5-30.5

28.4

Anterior TD

12.3-14.6

13.8

Proximal APD

17.1-21.0

19.5

Proximal TD

13.0-16.0

14.5

Semilunar

Distal APD

14.3-17.6

16.4

Maximal Height

16.1-20.3

18.6

Distal TD

10.2-14.4

12.0

APD

24.0-27.9

25.7

Proximal TD

14.4-18.6

16.1

ni Phalanx

Plantar Length

30.1-36.6

33.6

Unciform

Dorsocaudal D

22.7-28.0

24.8

Maximal Height

12.2-15.2

14.2

Maximal TD

11.7-14.8

12.9

APD

20.0-24.0

22.3

Proximal TD

13.0-17.2

14.9

Metacarpal

Length

242.0-275.0

263.0

Magnotrapezoid

AD501’95 PQAD2574

AD609’94

Proximal APD

21.0-27.4

23.2

Proximal TD

29.4-38.0

32.5

Anterior Height

14.8

14.6

Distal APD

21.6-26.4

24.1

APD

23.8

21.8

19.3

Distal TD

32.0-39.3

35.8

Anterior TD

19.2

16.8

15.0

Posterior TD

18.5

15.3

16.3

Talus

Latéral Length

38.5-46.3

42.3

Médial Length

36.2-44.0

40.7

Latéral APD

19.7-27.5

23.0

Médial APD

21.5-25.0

24.1

Tibia PQAD2292

PQAD1100

PQAD42

Distal TD

22.5-28.8

25.8

Lenglh

360.0

350.0

317.0

Calcanéum

Proximal APD

71.1

Length

94.0-106.0

100.2

Proximal TD

64.5

59.2

c.c. Length

61.0-75.4

67.9

Distal APD

32.1

29.4

27.2

t.c. APD

23.1-27.7

25.9

Distal TD

39.1

35.6

34.8

GEODIVERSITAS • 1999 • 21 (2)

249

Morales J., Soria D. & Pickford M.

which is losr in somc fossil giralTids such a.s

Palaeotragui mlcrodon (Koken, 188S). Sarno-

therium s'inerisis (Schlosser. 1903)i Deccenn-

therium püchecai Crusakme-Pairo. 1952, and

Sivatherium hvndeyi Harris, I976, but which is

still présent in rhe tvvo extant giraM'ids {Gmijfn

camelopanialu Linneaus, I738 and Okiipht jolm-

stoni). In the scmilunar, the distal latéral faccl is

somewhat wider than the rnedial one. The

magnotrapezoid, which is similar to thac ol

Palaeoîragus primaevns Churchcr. 1970, has a

concave posterior margin and begins to develop a

posrerior keel berween the facets for the semiluiur

and the scaphoid. Iii the unciform. the sernilunar

facet Is parcicularly clongated in a posterior sense,

characiers which it shares with uihet glrafïîds.

Tarsus (V\^. IIE-G)

The astragalus is picsiomorphic. It is similar to

that of P pwnaeina but has a weakly expressed

posterior process in the medial proximal condylc.

There is no strong development of the proximal

latéral condyle as occurs in (2imacfH'erm gentiyis

and the distal trochlea has more or less symmetri-

cal condylcs, In posterior view, the navicular

cuboid shows a very srrong medial cresi disrally

which diminishes tu nothing at the proximal zone.

It has a fossa in a very lacerai position, unlike in

Pprimaevusy (Praffa and ükapia.

Metatamd

The proximal epiphysis of rhe metatarsal has

similar morphology lo that cP Andeganmyx ande-

gaviensis Ginsburg et ///., 1994, and the anterior

groove is open distal ly.

The meiacarpal, pelvis, tibia and calcanéum ail

hâve a generalized morphology similar to that of

P primaevus. Ilowcvcr. the scapula of O. hendeyi

has a IcsS'developcd supraglcnoid tubcrclc ihan

that of PdlûeomtgtLs primanms whcrcas its cora-

coid apophysis' Ls more reduccd than it is in the

latter species.

Discussion

Orangemeryx shares wirh other climacoccradnes

the same morphological pattern of rhe dentition

and the possession of coniplex supraorbiral apo¬

physes, In Nyanziitneryxpkkfbrdi 1984,

and Orangemeryx the frontal apophyses are orna-

meneed ac ihcir bases wnth rounded protubé¬

rances w'hich do not projcct far cnough to hirm

latéral points. In contrast, in Climacocerds ajiica-

n/zt Maclnncs, 1936, and cspcdally in C. gentryi

Hamilton, 1978, rhe points are considerably

elongared, imparting a deer-like morphology to

the apophyses. The morphology of the apo¬

physes permits us to separate the various spccies

intü twü gfoups : oiie comjrrising the two species

of C/imacoieras plus Nyanzatneryx ’l’homas,

1984, the other consisting of Orangemeryx.

PH\TOGENETIC RELAddONS

The relationships betsveen these two groups, now

con.sidcred to be subfamilies of the Family

Cliinacoceraiidae (- Cli macoceridae of

Hamilton, 1978), is based on the faet that they

share tlie same morphtjlogical pattern of the den¬

tition and the incipient élongation of the ncck

and the limbs. The presence of elongared cervical

vertebrae, including the atlas, ^^’as aiready nored

in Olbnacacernî gentryi \>y Hamihon (1978), the

atlas of W'hich is pa'>pornonally longer th,in it is

in Palaeotragus prtmaevits and Saniorhvrlum dfrP

canm Churcherr 1970. Even though no atlas of

Orangemeryx hendeyi has been found, compari-

son between rhe axes of rhjs species and that des-

cribed by Chiircher (1970) for Palaeotragus

priniuevHS is suggestive in this respect, and ir»di-

care.s to us that tins new' genus als{) possesscd a

long ncck.

TTic new' fossil material dc.scribcd herc tends to

confirm this rclationship bccausc Sperrgebieto-

meryx^ and Cltmacoceras (C. gentryi)

share elongated cervical vertebrae, including the

atlas, a chariicter which indicates a notable spe-

ciali/ation towards feeding froni high Food

sources. In ihc forms ol which the postcranial

skcleton is known, it is possible to confirm that

the cxcrcjnitics arc clcaiiy elongated.. 7‘hc clado-

gnini (Eig. 12) summarizes chc phylogencuc rcla-

tionships between the dcscribcd forms,

considering Andegamer^fx Ginsburg et al., 1994,

to be the oucgroup.

The Family Climacoceratidac has previously been

placcd close lo the giralfe^ on ihe hasts ol rhe

presence of a bilobed lower canine in Climaco-

250

GEODIVERSITAS • 1999 • 21 (2)

Miocene giraffoids from Namibia

Fig. 12 . — Phytogenelic relationships of Spcrrgebietomeryx

n. gen. Propalaeoryy Sfrom 0 r and Orangemeryx n. gen (o other

giraffoids. Nodes: 1, (primitive characters) dentition moderately

hypsodont: upper and lower premolar senes long: premolars

elongated and thus gracile; lower molars with strong stylids;

third lobe of m3 simple; hypoconid isolated. a conséquence oî

its dear séparation trom the entoconid and me anterior lobe; the

p4 has a simple metaconid. directeo posleriorly. anterior wing

simple, Its mpfpholtDgy differing Utile trom thaï ol the p3: lhe pi

fe présent wilh Iwo slrong roots: upper piemolars long. P2 and

P3 with well-defined anterior lobes, upper molars with strong

slyles and laie union of the iniernal lobes to The ectoloph:

appendicular skeleton gracile, with a lendency to élongation of

the etemenîs; 2. cervical vnnebrae. including the atlas^ elonga-

led; 3. loss o( p1 ; 4 hypBodoi'l denlitioo, The p4 wiili metaconid

disposée! more transvemally, with a tendency to turn anteriorly,

anterior wmg blIurcMtfi (and thus very different trom p3). tenden-

cy towards flatltning of lhe iniernal wall of Lhe lower molars;

5, presence of supraorbitnl apophyse:; wrth protubérant orna-

mentaiiüns ât Ihn hasA. with a hilufcate lerminatlon loss of pi,

6, apophyses wilh elongated slighlly <;o»npreiiised itionical mor-

phology below the bifurcation; 7, apophyses elongated and

cylindrical; 8, apophyses with well-defined points perpendloular

to the long axis of the apophysis (C. afticanus): apophyses com-

plex, ramified as in some cervids {C. gentryi).

ceras gentryi from Fort Ternan (Hamilton 1^)78),

but the présence of this character is not certain,

as was noced by Churcher (1990), At Arrisdrift,

there are at least sixreen lower incisiform teeth

attributed to Orangemeryx, none of which is bîlo-

bed. 11^ fossil canines and Indsors are preserved in

the saint* ratio thaï occurred in life, chen at least

four canines should be présent in the Arrisdrift

sample, in which case the lower canine in

Orangemeryx consisted of a single lobe. Other

authors hâve advocated ihis relaiionship

employing for the most part dental charactêrs

(Gentry 1994; Gentry & Hooker 1988; Janis &

Scott 1987) or hâve inerely considered ir as

incertae sedts within rhe Giraffoidea (Geraads

1986).

Apparently, the precucious spécialisation of the

postcranial skcicton séparâtes the Climacocerati'

dae from the GirafFidac. Neverrheless, primitive

giraffes, such as Zaraja zeheni Hamilton, 1973,

of the early middle Miocene ot Gebel ZeltcUt

Libya. ai.so po.ssessed an elongated atlas.

However, a deeper study of the postcranial skele-

ton of the forms iitvoivcd in this radiation of the

Giraffoidea is rcquired hefore we can confirm

that this speciaJization is a character rhat permits

the inclusion of the Girafiidae and the

Climacoceratidae in a monophyletic group.

These new ruminant fossils support rhe idea

expre.ssed by Ginsburg et ai (1994) rhat prior ro

rhe appearance of pecorans with frontal protubé¬

rances rhe group would hâve been srrongly diver-

sified, à diversification which afïccted not only

lhe dentition, but aiso the postcranial skeleton.

We can now détermine with somc précision the

relationships of most of the pccoran fornis lack-

ing cranial protubérances which are related at the

level of sister gtoups to pecoran familles which

do pü.ssess such protubérances. Morales et al.

(1995) described a horniess bovid from

F.lisabethfeld, a cliscovery which corroborâtes this

view of ruminant évolution. Fhcsc findings

strongly .support the hypothesis that chc appea-

rancc of cranial protubérances -w'as a biological

phenomenon which occurred virtually synchro-

nously {ca. lS-17.5 Ma) but indepcndcnily in

varioas lineages of pecorans (comprising at least

lhe Cervidae, Falacomciycidae, GiraBoidca and

Rovidae). Ir corroborate.s the hypothesis of

Morales et al. (1993) in which the appearance of

cranial appendagt's is considered to liavc been an

organic response ro global scale environmental

cliangcs which occurred towards the end of the

early Miocene.

GEODIVERSITAS • 1999 • 21 (2)

251

Morales J., Soria D. & Pickford M.

Acknowledgenieiils

We thank NAMDEB (M. Lain, J. D. Ward,

B. Biirrell) tor acccss to thc Sperrgebiet,

Namibia, and h)r funding and logistical support.

Support was also providcd by rhe Mission for

Coopération and Cultural Alfairs, Windboek

(Y. Maire, N. Weil), ihc Collège de France

(Y. Coppens), the Muséum national d’Hisioire

naturelle, Paris (P. Taquet), the Museo Nacional

de Ciencias Naturales, Madrid (proyectos

PB/95/0n3 and PB/95/01 14)> the Geological

Survey of Namibia (B. Hoal) and the Spanish

Agcncy for International Coopération (AECI).

Wc thank Drs B. Senut, P Mein,. D. Gommer}',

A.-M. Bacon and Ward l'or help in dtc ficld.

Rescarcli Permission was approved by the

National Monuments Council, Namibia

(G. Hoveka, A. Vi>gi). Wc thank Dr D. Cicraads

and an anonymous reviewer for their comments

on the manuscript.

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GEODIVERSITAS • 1999 • 21 (2)

253

Snout proportions in some Eurasian hipparions

(Mammalia, Equidae): taxonomie and functional

implications

Ann FORSTEN

Finnish Muséum of Natural History,

PB 17, FIN-00014 Flelsinki (Finland)

ann.forsten@helsinki.fi

Forsîen A. 1999. — Snout proportions in some Eurasian hipparions (Mammalia, Equidae):

taxonomie and functional implications. Geodiversitas 21 (2) : 255-278.

KEY WORDS

hipparions,

snour proportions,

phylogeny,

Kinction,

ABSTRACT

In the hipparions, length and breadth of the snout and depth of the nasal

opening are noc well correlated with one another, nor with the shape and

placement of the preorbital fossa, recently considered taxonomically impor¬

tant in these equids. These characters are distribuied in a mosaic-like fashion

among specics, making species différentiation and phylogenedc reconstruc¬

tion difRcult.

MOTS CLÉS

hipparions,

proportions du museau,

phylogénie,

implications fonctionnelles.

RÉSUMÉ

Les proportions du museau chez quelques hipparions (Mammalia, Equidae)

dEurasie : implications taxonomiques et jonctionnelles.

Chez les hipparions, la longueur et la largeur du museau ainsi que la profon¬

deur de la fosse préorbitairc ne sont pas clairement corrclées entre elles, ni

meme avec la forme et la position de cette fosse. Pourtant la valeur de ces

mesures a été récemmeitt considérée comme ayant une signification taxono¬

mique importante chez ces équidés. Ces caractères suivent une distribution

en mosaïque, au sein des différentes espèces considérées, et leur utilité tant

pour la différenciation spécifque des hipparions que pour leur analyse phylo¬

génétique est restreinte.

GEODIVERSITAS • 1999 • 21 (2)

255

Forsrcn A.

INTRODUCnON

While tlic preorbital fossa of the cheek has

recently becn given great weight in hipparion

raxonomy> even used for delimiring “supraspeci-

fie grüii[)s”- rhe proportions and structure of tbe

snout and rbe nasal opening bave largely been

ignored. Complété skulls of Hipparion de

Christol, 1832, are rare. Mosc often che snout

and/or dic cranium aie broken oft^ leaving rhe

middie part of rhe skull wich the cheeks, tooth

rows and orbics prescivcd. f'hus ihere îs much

Icss data on rhe snour of Hipparion than on the

preorbital fossa of the chcck.

l'he function of the fossa in Hipparion is still

unsolved, airhougb chere are several alrernarive

hypothèses (see discussion in Setve 1V27: 67-78).

In recent equids the shallow fossae serve as

attachmcni areas (or Icvator muscles of the upper

lip (Ellenberger &: Baum 1943; Zhcgallo 1978);

the very nitibik lips foncriofi together with the

incisors in g.uhering fotïd. "l’he extension of ihe

nasal opening in Hipparion may refiect the deve¬

lopment of the nasal diverticulae, in receni

equids siiuated within the nasal opening

(Ellenberger Bauni 1943; but sce Gregory

1920).

The shapc of the snout in fossil and extant herbi¬

vores has rccendy been debated (e.g. Bunnell &

Gillingham 1985; Solounias &: Moellekcn 1993;

Dompierre & Churchcr 1996; Eisenmann 1998)

and there hâve been atrempts to interprète it eco-

logically. The shape o( rhe snout is believed to be

dictated by chc mode ol fccdlng. A broad, antc-

riorly flatrcncd snour is interpreted as indicaring

grazing or chc unselectivc gathering of fotjd near

ihc ground. A narrow snout, with tlie incisors in

a sharp arc, is interpreted as indicating hrowsing

or the sélective gathering of tood at different

heights (for the illustration of equid snours of

different breadth, see Eisenmann 1998, fig. 5: I-

10 ).

In an carlier paper, l (Forsten 1983) discusscd

the variation in the placeinent and shape ot the

preorbital fossa in somc Old World Hipparion,

In the présent paper I will desetibe, compare and

discLiss the snout and the nasal opening. Old

World hipparions group into broad- and narrow-

snouted taxa, but with many intermediates.

rhere is vvcak coircspondence with groups dell-

miccd on the basis of the preorbital fossa. AIso

snout proportions and nasal opening extension

are only weakiy correlated. How sbould rhese

characters be evaluaced and weighted taxonomi-

cally and pltylogenctically?

METHODS

In the upper jaw>. snout Icngrh was measured as

the distance from the prosihion (Il-II) to rhe

middie of a line uniring the anterior tips of P2

(Gromova 1952, table I, sktill measure 18);

snout widrit wavS measured as rhe ourer distance

beliind 13-13 (GromovaV measure 40). l’Ite nasal

opening is tbe distance anicriorly Irom berw'cen

tbe premaxillac (not including tbe upper sym-

physis) postcrioi’ly to whcrc the nasals and pre-

maxillac/maxilhtc mect (Gromova's measure 28).

Skull Icngth, since seldoin measuieable, was here

substiiuLcd hy the distance P2-anterior rim of

orbii (Gromovas measure 11), which is roiighly

correlated with basal lengtli.

lu the lower jaw snout breadth was measured as

tlie oiiter distance behind i3'i3 (Gromova 1952:

table IV, measure 14) and syinphysial length

anteriorly from betwéen il-il posteriorly to ihe

symphysial noicb (Gromovas measure 11).

Relative .snout width is expressed in scarter dia-

grarns, ploiting maxillar snout width ro snout

Icngth and mandibular snout width to symphy-

sial lêngih. rcsptxtivcly. Snout Icngth îs rckted to

skull length by plotting ir lo tbe disiancc 1*2-

orbit. Nasal opening Icngth is cxpte.ssed boch as

an absolutc mevisure and in relation to the tooth

row. l’he preorbieal fo.ssa is discussed in terms as

in l'orsien (1983). 1 calculatcd 95% cquiprobabi-

lity ellipses on the data of H. schlosseri-rlietrichi

from .Sainns Q1 and Samos without more exact

locâIit^■ data and of H. nioldaincmn Cjromova,

1952 from Taraldia and Novoelizavetovka, using

metrical dara, rben used rhe ellipses as modeis in

che diagrams. I did not plot the spécimens ased

for calculacing the ellipses, but type specimens

are ploiied in bold leitcr abbreviations.

The maierials used and the institures in which

are kept the macerials seen are listed in the

Appendix.

256

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some F.urasian hipparions

Fig. 1. — Upper snout breadth plolted to snout length in European hipparion skulls: measurements In mm; 95% equiprobabiTity

ellipses calculated and drawn on the data of Hipparion schlosseri-dietrichi (Samos Q1 -Andriano and Samos withoul exact locality)

and H. mofdavicum (Tarakliya and Novoelizavetovka). Ptotted are specimens from the localities: Cer. Cherevichnoe; Ch. Chimislia;

K. Karaslari; M. Maragheh; ML. Mt. Luberon; MoL Molayan: Sq. Salonikî (coll. Arambourg & Puyhaubert); T. Tchobruchi; Tu.

Tudorovo; UD, Umen Dol. Bold type abbreviatlons in figure depict type specimens: Ch, H. pregiganteum {Chisnau 4040/84); Mc.

H. campbefli iUn'w, California Riverside No. 13/1342); Mg. H. gettyi (Wien No. 840); Sd, H. dietrichi (Münsier S 1/7); Ssch, H. schlos-

seri (Wien 1911 V 114}: Ta. H. moldavicum (Moscow PIN 1256-3639); Tu, H. tudorovense (OGUM 1780). Observations in paren¬

thèses approximafive.

DESCRIPTION

Europe anu ti ie Near East

Hippariom loith a short andJor broad snout

Hipparion prostylum Gervais, 1849 [localities:

Mt. l.uberon, France; Saloniki, Grcccc; Karaslari

and Umen Dol, Macedonia (former Yugoslavia);

possibly Maragheh, Iran], H. schlosseri'dietrichi

Antonius, 1919'(Wehrli, 1941) fSamos without

exact localitv (Sondaar 1971, pl. Il: a); Samos

Ql, Q4. Q6 (Sondaar 1971, pl. Il: b);

Vathylakkos, Prochoma-1, Ravin des Zouaves

(Koufos 1987. fig. 3), Greece; Basiboz (Forsten

& Garevski 1989, photos 3. 4), Macedonia; and

Maragheh] hâve a short and broad snout (Fig. 1 :

upper ellipse). The snout is short also in relation

GEODIVERSITAS • 1999 • 21 (2)

257

Forsten A.

130 140 150 160 170

Fjg. 2. — Upper snout length plotted to P2‘Orbit distance in Européen hipparion skulls; measurements in mm; 95% equiprobability

ellipses calcuiated and drawn on the data of Hippanon schlosseri-diêtriahi (Samos Ql-Andriano and Samos wtthoul exact locatity)

and H. moWsWcümtTarakliyaand Novoelizavelovka). PIoHed are specimens from: Cer, Cerevichnoe: Ch. Cbimislia: M, Maragheh;

ML, Mt. Luberon: Mol, Molayan; Sq, Saloniki; T, Tchobrucht. Bold type abbreviations in figure depict type specimens: Ch. H pregi-

ganteum: Mc, H. campbelfi: Mg, H. gettyr, Sd, H. dietrichi, Ta. H moldavicum-.Tu. H fudofovense, Observations in paren(he5es are

approximative.

to skull lengtfi (Fig. 2: lowcr ellipse) and in thc

lower jaw ihe snout is broad in relation to the

length of rhe symphysis (Fig. 3: upper ellipse).

The single preorbital fossa of the skull is placed

rather far in front of the orbit. It may be faintly

delimited and shallow or well-defined, in citber

case more or less pocketed. There is no clcar dif¬

férence in skull morphology berween rhe smalter

specimens (e.g. H. prostylum and the holotype of

H. dietrichi^ Münster SI/7) and thc larger ones

258

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

50 60 70 80

Fig. 3. — Lower snout breadth plotted to symphysial length in European hipparion jaws; measurements in mm; 95% equiprobability

ellipses calculated and drawn on lhe data of Hipparion schlosseri^dietrichi {Samos Q1-Andriano and Samos without exact locality)

and H. moldavicum (Tarakilya and Novoelizavetovka). Plotted are specimens from: Ch. Chimislia; M. Maragheh; ML, Mt. Luberon;

RZO, Ravin des Zouaves; T, Tchobruchi; Tu, Tudorovo.

(c.g. Wicn 1911 V 114, thc holotypc of

H. schlosseri, and Budapest No. 274) (Figs 1, 2:

ML, and in bold; Sd, Ssch). The schlossen

and dietrichî were given the same species;

although Antonius’ name “jrA/owr/” is older,

thcrc is some uneertainry as to its validicy (inadé¬

quate description and illustration of the type).

Until the question as to the correct name is sol-

ved, I use both united by a hyphen: H. schlosseri-

dietrichi.

In this group aiso belongs H. molayanense

Zouhri, 1992 described from Molayan,

Afghanistan (Zouhri 1992). In thc two skulls

seen, the snout is short, but not as broad as in

the former group (Zouhri 1996, pl. 59) (Figs 1,

2: Mol). The preorbital fossa resembles that of

the former group in being shallow, oval-egg-

shaped, and situated far in front of the orbit.

d'he above mentioned short- and broad-snouted

hipparions, H. prostylum^ H. schlosseri-dietrichi^

and H. moLiydnense^ also hâve a short nasal open-

ing, anteriorly cither blunt or softiy pointed,

ending at a level wcll in Iront of P2. The nasal

openiug is short also compared with snout and

slcull length, excepe in Paris Mol. 040 from

Molayan.

Among the skulls with a short-broad snout fall

the type and the referred specimen of H. garedzi-

cum Gabuniya, 1959 from LJdabno, Georgia

(Fig. 4: U, in bold). The nasal opening is short,

ending at a Icvel in front of P2 buuthe preorbital

fossa is well-defined and deep (see Gabuniya

1959, pl VI: 1). On the other hand, among

skulls with a shallow preorbital fossa situated far

from the orbit can be mentioned the specimen

B-50 from Bazaleti, Georgia, referred to as

GEODIVERSITAS • 1999 • 21 (2)

259

Forsten A.

Fig. 4. — Upper snout breadth plotted to snout lenglh in European hipparion skulls; measurements in mm; 95% equiprobability

ellipses calcutated and drawn on the data ot Htpparion schbss&n-dietrichi (Samos Ql-Andriano and Samos wilbout exact locaJity)

and H. moldavicum (Tarakliya and Novoelizavetovka). Plotted are specimens of H. matthewt (scatter to ttie left) from. B. Boluska;

Dyt, Dytiko (data Koutos 1983. table 1), Q5. Samos Q5: S, Samos withoui exact focaliiy. Ploiied also liippanons în the H primige-

nium group (scaMer lo rtie righl) trom; B. fieluska: VC. Vila de Caballs; Gr. Greboniki: N. Nesebr: OH. Oued el Hammam;

Pi. Pikermi: 04. Samos 04: S. Samos without exact tocality: U. Udabno: W. Inzersdort Dold type abbreviations In figure depict type

specimens- Gr, H. giganteum (OGUM lOtS), Hos. H catalstunicum (BMNH 16397): Sm, H. màtfhewi (Bud&pesi Ob/5S7);

U. H. ga^ecI^icum (Tb*lisi No 156/13)-

H. garedzicum by Melad/e (1967, table VIII) and

as H. moLiyanense by Zouhri {1992^ 1996), but

which dificrs iront both in the snout being long

and narrow (Fig. 5; Bz to the far right in dia-

gram). The type skull of H. tudorovense

Gabuniya, 1959 (OGUM I7H0; Gabuniya

1959, pl. V: 2) irom Tudorovo, Moldova, aiso

bas a shallow prcorbiral fo.ssa siruaied rclatively

far in front of tbc orbii, but a narrow snout

(Figs 1, 2: Fu, in bold). The nasal opening in B-

50 from Bazaleti and OGUM 1780 from

Tudorovo ends Icvel with F2, A very Iragmentary

skull (Inst, üf Paleobiology, d'bilisi No. 148/I91t

Gabuniya 1959, pl. VI: 3) from Kiourteviouf

50 km north of bake Urmia, Iran, the holorype

ol H. urmiense Gabuniya, 1959 bas a shallow

and faim (ossa and a nasal opening exrending

level with P2'r3. Since the snout is lacking and

the orbits are not visible, the placement of the

preorbital fbssa in relation to the orbic and tbc

snout proportions are unknown.

A rclatively broad snout is characterisric of the

skull ot H. mattheiv! Abel, 1926 [localitics:

Samos Q5 (Sondaar 197T Fl. Ia*b) and Samos

without exact locality data {H. nicosi Bernor

Tobien, 1989), Ravin de la Pluie, Vathylakkos,

and Prochoma-l {H. mncednnicum Koulos,

1984; Kouios 1987, fig. 8), Dytiko (//. matthe-

un and /•/. peridffivtiuii}u Villalta 6c Ctusafont,

1957; Koiiios 1987), Salonîki, Grcccc (coll.

Arambüurg 6i Puyhauberr); ?Umcn Dol and

Beluska, Maccdonia; and Ploski Blagoevradsko

{H. murvtaton Nikulov, 1971), Bulgaria] but the

relative breadth is les.s rluiii in rhe //. prostylmnl-

scblosscri-dietrichi group (Figs 4, 6: scaticr to the

lefi in diagrani). In the lower jaw the snout is

medium broad relative ro symphysial lengch

(Fig. 7: scacrer ro the Icft in diagram). The pre-

utbitul fossa in mauhewt varies from well-defined

to almosi absent; în somc specimens front Q5 it

is double, consisting of a posterior fossa proper

and an anterior, smaller subnasal fossa (Forsten

260

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

Fig. 5. — Upper snoul breadttt plotted to snout lenglh in European hipparions; measuremenls in mm; 95% equiprobability ellipses

calculated and drawn on the data of Hippanon schlosseri-dietrichi (Samos Qi -Andriano and Samos wiîhout exact locality data) and

H. moldavicum (Tarakliya and Novoelizavetov/ka) Plotted are specimens ot: Hipparion verae (Gr. Grebeniki: K, Karaslarl ); H. medi-

terraneum [Bz. Bazaleti (specimen B-51 to lhe left in diagram); Dyt, Dytiko (data Koufos 1988. table 1); Pi. Pikermi (data partiy

Koufos 1987, table i)); Hipparion sp. with a double fossa (Q1 and ?Q5. Samos quarries); Hipparion sp. (P. Piera). Plotted are aiso

skulls ûl H. proboscideum (scatter to lhe right in diagram) from; Cer. Cherevichnoe: RZO, Ravin des Zouaves; Spr, SI and Q1.

Samos quarries. The skull B‘50 Irom Bazaleti with a weak fossa falls in among the specimens of H. proboscideum. Bold type abbre-

viation in figure depicts type. Gr, H. \/erae (OGUM 1016).

1983). l he opcnlng in H. marthewu Hke

snout lengtli, varies froni short to long or irom a

levcl 1 cm in front of P2 to Icvel with P3 para-

style. The fragineniary nasal opening in

München 1899 Vil 31h (H. nicost holotypc)

from Samos exleruls level with P2 and is vhus

comparable lo the holoiype ol manhewi

(Budape.st OK/357), aIso from Samos. In the

rather long-snoutcd specimens with a double

fossa from Samos Q5, the nasal opetiing is long.

The skulls of IL matthewi irom QS re.semhie

some larger skulls with a double fossa tound

from Samos QI-Andriano (see Porsten 1983,

fig. 3), pos.sibly aiso from Samos Q5 and

Gülpinar, Turkey (Higs 5, 8: Ql, QSj; they arc

hcre referred co as H. sp. (with a double fossa).

Thcy bave a relatively short-broad snout and the

nasal opening ends level with P2, but both the

snout and the nasaJ opening arc longer in rela¬

tion to .skuli and snout Icngth, rc.spcctively, than

in H. schlosseri-dietnchi of similar size and from

the same localiries. 1 lii.s untiamed trtxon scems to

be rclaccd to the hipparions in rhe H. jnediterm-

Tinmr {Koxh îk Wagner, 1853) group. in which

the fossa is situated close to the orhii, but is rare-

ly double. The proportions of the snout and chc

depeh of chc nasal opening in thèse specimens

from Samos are like in H. m{'dite}ri7}n'un/ from

pikermi, Grcece, but relatively bruader and deep-

er, respectively, than in the varions local forms of

H. moldavicum Gromova, 1952 from Moldova

and Ukraine, belonging in the same species

group.

Hipparions ivith a long snout andlor nasal opening

Extrême for ics long and narrow snout among

GEODIVERSITAS • 1999 • 21 (2)

261

Forsten A.

Fig. 6. — Upper snout length plotted to P2-orbit distance in European hipparion skulls; measurements in mm; 95% equiprobability

ellipses calculated and drawn on the data of Hipparion schlos$en<iietrichi (Samos QVAndnano and Samos wiihout exact localfty)

and H. moldavicum (Tarakliya and Novoelizavetovka), Plotted are specimens ot H. matthewi (scatter to the leti in diagram) Irom:

B. Belushka, Q5 and S. Samos quarries. Plotted are aiso spécimens of the H. primigenium group (scatter to Ihe right) from;

B. Beluska: Gr. Grebeniki: Hos. Hostalels; Hw HÔwenegg; N. Nesebr OH, Oued el Hammam: Pi. Pikermi; Q4 and S, Samos quar-

ries; U, Udabrio: VC. Vila de Caballs: W, Inzersdorf. Bold type abbreviations In figure depict type specimens: Gr H. giganîeunr,

Hos. H. catalaunicunr. Sm H maithewi: U. H garedzicum.

the European hipparions is the specimen B-50

from Ba/aleri with a shallow fossa (Meladze

1967, table VllI) (Figs 5, 8: B/, ro the far right

and top centre in diagram). Specimen B-51 from

the same locality, but with a maximaliy dcvelop-

ed, double preorbital fossa, has a medium long

snout similar in ics breadth: length proportions

to that in H. moldavicum (Meladze 1967,

table Vil) (Figs 5, 8: Bz in centre of diagram). (n

borh specimens the snout is longer than in

H. mediterraneum^ and in the lowcr jaw (Meladze

1967, table IX) the symphysis is longer and the

262

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

Fig. 7. — Lower snout bnsadih plotted lû symphysiat langth în European hippaflon jaws; measufemef>îs In mm; 95% equiprobability

ellipses calculated and drawn on lhe data ol Hipparion schlossen^d/etrichi (Samos OvAndnano and Samos wiihou! enacl locality)

and R moldavtcum (Tarakiiya and Novoalizavotovha)- Plotted are specirrrens of H /Tî(?ffnevvriscaner to the lefi) Irom; B, BelusKa:

Dyl. DytîKo (data Koutos T988. table 2): Q5 and S. Samos quarries; RPI, Ravtn de fa Pluie; Sq Saloniki. Plotted are also (scattor lo

the nghi in diagramj not separable specimens of hipparions ot ihe H primigemum group and R probosctdeum from- Ad S, Ol and

Q5. Samos quarries; B. Befüska; CL, Can Llobatores; E. Eppe‘sne*m: Gr. Grebeniki: K, Kafaslari: Kf, Kalta; M Maraghch:

N. Nesebr. OH. Oued el Hammam. Pi. Pikermi; RPf, Ravin de la Pluie V, Vozarii W Inzersdorf and Wienerberg Bold type abbre-

viations in figure depicî type specimens: Sm, R matthewi. Observations in parentneses are approximative.

snout narrowcr (Fig. 9: Bz). Compared for their

nasal opcning dcpch, that in B-50 is absolutely

shorter rhan in B-SI, but in boch .specimens ihc

nasal opcning rcachcs Icvel with V2.

Hîppnrion sp. Front Fiera, Spain, has a wcll-

dcfincd, pockctcd, pcar-shapcd to roundcd rrian-

gulaj* prcorbital lossa and a long snout in relation

to skull lengdt (Forsten 1968, pl. 2) (Figs 5, 8;

P). For its brcadrh, the snout reseniblcs iltai in

H. mediterraneurn and //. venie Cjabuniya, 1959

(Fig. 5: P. Pi. Gr). The nasal opcning is short,

ending front a level 2 ont in front of P2 to Icvel

with P2 paraconc; il is short also in relation to

snout and skull lengch.

Hippiirion proboscidtum Sixxdttu 1911 llocalities:

Samos Ql and SaniOsS without exact locality data

(Sruder 1911, figs 1, 4a; Sondaar 1971»

pl. Illa-d), Ravin des Zouaves (Koufos 1987,

fig. 3), Grccce; Vozarzi and Karaslarl, Maccdonia

(Forsten & Garevski 1989, photos 1, 2); Valea

Sarii, Roinania (Forsten 1980); Kavakdere,

Turkey; and Chcrevichnoc, Ukraine, {H. sp. cf

proboscidcitm Forsten Krakhmalnaya, 1997,

fig. 7)1, with an arrangement of the double

prcorbital fbssa similar to that jn specimen B-5l>

has a narrow snoui similar lo rhat in B-50, but

the snout is not long in relation to skull Icngth

(Figs S, 8: 8pr, SI, Ql, RZO, Ccr). The nasal

opening is long and deep. extending from level

with P2 parastyde to P3 mosostylç. In its snout

proportions H, proboscideun) rçsembles the

second large species from Samos Ql-Andriano

and Q4, called II cf. probosckkuni and IL pri~

migcniiiui (v. Meyer, 1829) (Sondaar 1971,

pl. 111: c; Forsten 1983, in Appendix referred to

as H. aft. brachypus Hcnscl), in which the single

prcorbital fossa and nasal opcning arc as deep as

in the former, but in which the snout is longer in

relation to skull lengch and broader than in

H. proboscideum (Figs 4, 6: S, Q4). This taxon,

GEODIVERSITAS • 1999 • 21 (2)

263

Forsten A.

Fig. 8. — Upper snout length plotted to P2-orbit distance in European hipparion skulls; measurements in mm; 95% equiprobability

ellipses calculafed and drawn on îhe data of Hipparion schfosseri-dieirichi (Samos Ql-Andriano and Samos wilhout exact locality)

and H. moldavicum (Tarakiiya and Novoelizavetovka). Plotted are specimens of H. rnediterraneum [Bz, Bazaleti (specimen B-51 in

centre of diagram); Pi. Pikermil. H. varae fGr. Grebenikl); Hipparion sp with a double tossa (Q1, Samos 01): and Hipparion sp.

(P. Piera). Plotted are also spécimens ol H proboscideum (scalter to ihe right) from: Bz. Bazaleti (specimen B-50 in top-centre of

diagram); Cer, Cherevichnoe; 01. SI and Spr. Samos quarries, RZO, Ravin de Zouaves. Bold type abbreviation in figure depicts

type; Gr. H. varae.

similar to ihc large one front Pikermi (H. bmehy-

pus Henscl, 1862, .scc Koufos 1987: pl. Ill)

(Figs 4, 6: Pi), leads over to liipparions wirh a

mostly long snout, comprLsing the H. mediterra-

neiim and the H. pr 'tmigenium species groups.

Hipparioiis with a ynedium long and narrow snout

A medium long and narrow snout characterizes

the local samples of H. moldavicum [locaUtics:

'larakiiya, Novoelizaverovka, Tudorovo,

Chimishiia, Cherevichnoe, and Chiobruchi,

Ukraine and Moldova, possibly Maragheh, Iran)

{Gromova 1952, tables l-III; Gabuniya 1959,

pis III: 2, 3> IV: 1, 2; Watabe ^ Nakaya 1991;

Forsten & Krakhmalnaya 1997, fig. 6) (Figs 1, 2:

lower and upper ellipse, respectively)]. The nasal

opening extends from a level 1 cm in front of P2

ro P2 mesosryle; ir is short in relarion ro snout

and skull length. în the lower jaw the snout is

narrow in relation to symphysial length (Fig. 3:

lower ellip.se). Hipparion moldavicum lias a well-

defined, single preorhital fo.ssa placed close to rhe

orbit, a character which it shares with H. medi-

lerranenm from Pikermi, but has a narrower

snout and shorter nasal opening even taking into

account îts smaller skull size. Similar narrow

snout proportions are found in skulls from

Maragheh. dcscribed as //. getiyi Bernor, 1985

and H. campbelli Bernor, 1985 (Bernor 1985), in

OGIJM 1780 (iy[)e o( H. tudnrovense) from

Fudorovo, and in the type skull (Kishinev

4040/84) of H, praegiganteurn Tarabukin, 1967

from Chimishiia, Moldova (Figs 1, 2: "lit, Mg,

Mc, Ch, in bold). The single preorbital fossa of

264

GEODIVERSiTAS • 1999 • 21 (2)

Snout proportions in somc Eurasian hipparions

50 60 70 80 90

Fig. 9. — Lower snout breadth plotted to symphysial length In European hipparion jaws; measurements in mm; 95% equiprobability

ellipses calculated and drawn for Hipparion mediterraneum from Pikermi and H. verae from Grebeniki. Axes aiso drawn on the data

of H. schlosseri-dietrichi (Samos Ql-Andriano and Samos without exact locality) and H. moldavicum (Tarakiiya and

Novoelizavetovka)- Plotted is single specimen B-54 jaw from Bz, Bazaleti.

these skulls is roiinded rriangular tü oval, shallow

or witli a posterior pocket, and situated niodcra-

tely far from the orhil. I hc nasal opening ends

level with P2; it is particularly long in relation to

snout and skull lenglh in the type skulls (Wien,

No. 8401 and BMNH 44574/cast of Univ.

California Riverside No. 13/1.342) of H. gettyi

and H. campbelli from Maraghch. Hipparion get¬

tyi may l^e a yuunger synonym of//. tudorovettse^

with which it shares similar snout proportions

and proportions between the orbil-fossa: P2-

orbit distances, and similar extension of the nasal

opening. Hipparion campbelli may be a younger

synonym of H. urmiense^ with which it shares a

similar long tooth row and the extension of the

nasal opening (see Watabe &: Nakaya 1991 ).

Hipparions with a medium long and broad snout

Althüugh similar in the shapc and placement of

the preorbital fossa, H. mediterraneum (Pikermi

and Dytiko, Grcecc, Koufos 1987, pis 1, 11;

Kûufos 1988, pis 1, 2; possibly Maraghch, Iran,

and Bazaicii, Georgia) has a broader snout than

H. moldavictitm although ovcriapping the lattcr

in snout proportions (Pigs 5, 8: Pi, l)yt). l'hc

nasal opening is deeper, rcaching Icvcl with P2,

even with P2-P3.

Hipparion verae from Grebeniki, Moldova, has

GEODIVERSITAS • 1999 • 21 (2)

265

Forsten A.

Fig. 10. — Upper snout breadth plotted to snout length in Asiatic hipparions; measurements in mm. For comparison the axes are

drawn on the data of Hipparion schlosseri-dietrichi (Samos Ql-Andriano and Samos without exact (ocality) and H. moldavicum

(Tarakiiya and Novoeli^avetOv/Ka). Abbreviatfons. CGC Chang Chia Chuang and 2 miles W of Chang Chia Cliuang CMK Ch'eng

Chia Mao Kou. Hd. Htpparior} dermatorhinum: HKP. Hsiao Ku Po; HK$. Hstao Kou Shan: Hl. H licenti: HL. Hsi Liang; HLK, Huang

Lu Kou; Hr, Pfobosdcfippancn roanantis {vaxious locations). KN Kfiirgis Nuf Kp Kalmakpai tant. Lanlian. LKL Lu Kao üng;

LT, Ling Tou: LWK. bao Wang Kou NH Nan Ho' NHH. Nan Hao Hsia; PNP. Po Niu Po; Pp. P 'pater. Ps. P stnense PTT. Pai

Tao Tsun: Sor, Sof. Sw Siwaliks; YM. Young Mu Kou: 30. Loc. 30; 43, Loc. 43: 44 Loc 44; 49. Loc. 49: 70. Loc. 70:73, Loc- 73;

109. Loc. 109, Bold type abbrevtations in ligure depict the lyprs. 49, H. plocodus Sofve *927 (Uppsala No 3824), 30 H tossatom

Sefve, 1927 (Uppsala No. 3821). Lant. H. weihoanse llu Ü èt Zhai, 1978 (IVPP V 3113 1); Ps, Probosadipparion smense Sefs^e

(Uppsala No. 3925 & 3926).

snout proportions rcscmbling those of H. medi-

terraneum (Gabuniva 1959, pis II: 1, 3, III: 1,

VIII: 2) (Figs 5, 8:'c:r. Pi). In thu lower jaw t)f

both forms the snout is broad in rclaiion lo chc

length of the symphysis (Fig. 9: Gr, Pi), evcn

rcsernbling the jaws ol (he H. prostyluni/schlosseri-

dietrichi coaiplcx. Ifippctruw vertlv has a diffe-

rcnrly shaped and placcd preorbital (ossa and tlie

nasal opening is shallower than in //. meditena-

neum^ reaching (rom a level 2-1 cm in Iront of

P2 to P2 nicsosryle. Like in the lutter, the preor-

bital fossa is occasionally double,

in skulls frem Basibo/, ümen Dol and Karaslari

(Macedonia, former Yugoslavia), referred to as

H. verae in Forsten Garevski (1989, photos 5-

7), the prcorbital fossa spills into the fossa bucci-

nacor. in sonie cases forining an isolaied, anterior

subnasal fossa Icvcl widi P2. In thèse spccimeus

chc placement, shapc and depth of the prcorbital

fossa varies, as does tlie depth ol the nasal ope¬

ning (ending from 1 cm in Iront ol P2 to level

wirh P2 paracone). The snout is medium broad

in t\vo measureable spccimens (PMM Kar 86/73

and IGir 203/73), belonging cither to this species

or to //. prostyhindschlüsst'rî-dietrkhi (Fig. 5: K).

Hipparimis lacking a pmjrhital jossii

The earliest known Hipparion in Europe lacking

266

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

Fig. 11. — Upper snout lengtti plotted to P2'Orbit distance in Asiatic hipparions; measurements in mm. For comparison are drawn

the axes calculated on the data of Hipparion schlosseri-dietrichi {Samos Ql-Andriano and Samos without exact locality) and H. mol-

davicum (Tarakiiya and NovoeÜzaveïovka)- For abbrevialions ses Figure 10 Bold type abbreviaiions in figure depict types:

Hd, H. dermatorhinum {Uppsaïa No. 3872): Lant, Hipparion weihoensis: Ps. Proboscidipparion sinense: 30. H. fossatum.

a preorbital fossa is H, p/atygertys Gmmovay, 1952

from the Meotian of Tarakiiya (Gromova 1952,

table IV). Its upper snout is unknovvii, but in

three lower snouts the symphysis is lonj; and the

snout medium broad in the mcasincablc spéci¬

mens. Gromova (1952) reterred lhe .spccimen

PIN 1256-.3634 to H. sp. or H. plaiygtnys.

HipparionpliZtygmys may bc a junior synonym ot

H. hippidiodus Sefve, 1927 trom China (see

below).

From the Pliocène is known Probtneidippurion

rocinanth (Hernandcz-Pacheco) from Vlllaroya,

Spain, and Kvabcbi, Geur^ia, lacking or with a

faintly defincd preorbltal l^ossa. This specie.s

represents the dentally advaneed caballoid hippa¬

rions, refcrabic to a genus of their own:

Proboscidipparion SefVc, 1927 (Forsten 1997).

The snout of ihe skull and jaws from Villaroya i.s

long and medium broad; thaï of the skull from

Kvabebi (Vckua 1972, Ris. 27-28. tables XVllI,

XIX) i.s short and broad. The nasal opening is

short, ending in front of or Icvcl with P2> rcspcc-

tively. Proboscidipparion rocinautis is aiso found

m China and Mongolia, where spécimens tend

to hâve a narrow snout (see below).

Phe Hipparion primigenium

Among the early (Vallesian) forms of rhis group

represented by skulls with siiouts are the hippa¬

rions from Vila de CabalLs and Hostalet.s, Spain

[H. catalannicum Pirlot, 1956; Pirlot 1956,

pl. V); Hôwcncgg, Gcrmany; Jnzcrsdorf, Ausiria;

Ne.scbr, Bulgaria {H. prestdaitnm Nikolov, 1971

and H. nesebricum Bakalov & Nikolov, 1962;

Nikolov 1971, table I: 1, la); Kalfa and Braila,

Moldova (//. sarmalicum Lungu, 1973; Lungu

1973, Ris. 1, cable 1); and Oued cl Hammam,

Algcria {H. africanurn Arambourg, 1959); latci

(Turolian) forms are found at Lklabno, Georgia

(H. gartdzicîirn Gabuniyu, 1959, pl. VI: 1);

vSamos Ql-Andriano and Q4 (H. cf. proboscu

Sondaar 1971, pl. lllc; hcrc H, aff. brachy-

pus), Pikermi (H. brachypas Koufos, 1987,

pl. 111), and Grcbeniki (H. giganieum Gromova,

1952, table XII), cvidenily aIso ai Rcluska,

Karaslari, and Vozar/i, Macedonia. While the

early forms hâve a iriangular or oval fossa, in the

lacer fbrnis the fossa is pear-shaped. The snout is

long and broad (Figs 4, 6), but the Icngth of the

nasal opening varies considerably between the

local samples. It is very shallow in the specimens

GEODIVERSITAS • 1999 • 21 (2)

267

Forsten A.

Fig. 12. — Lower snout breadth plotted to symphysial length in Asiatic hipparion jaws; measurements in mm. Abbreviations as in

Figure 10, in addition to Htc, Hipparion tchicoicunr, Ts. Tan Tsun: 12, Loc. 12: 52, Loc. 52; 71, Loc. 71; 110, Loc. 110. Bold type

abbreviations in figure depict types: Ps. Proboscidipparion sinense: 30. Hipparion fossatum.

from Hostalets (ending 3 cm in front of P2),

Inzersdort (3-2 cm). Kalia (2-1 cm), Howenegg

(3-1 cm), Utlahno and Grebcniki (from 1-2 cm

in front ol P2 to P2 anterior lip); medium deep

from Vila de Caballs (level widi P2 anterior lip),

Oued cl Hammam (ju.si in front of P2 to P2

paracone), and Nesebr (J^2 mcso.style); and deep

in ihe skulls from Pikermi (P2 mc.so:>iyIc-P3

paraconc) and Samos Ql-Andriano (lovcl with

P3). Hipparion garedzicum from Udabno (see

above) differs from rhe rest of tbe group in

having a short snoui relative to skiill length

(Figs 4, 6: U> in hold).

In rhe lowcr jaw ihe snout is broad in two

(MNHN 8^>, 143)» narrow in a third (MNHN

98) of the jaws from Oued el Hammam; il is also

broad in ihc jaw (Univ. Thessaloniki, RPl-1)

from Ravin de la Pluie, Grecce, referred to as

H. prirnigcnium (Koufos 1986), and from Can

Llobateres (Fig. 7: OH, RPI, CL). In die other

local saniples in ihis group rhe lowcr snout tends

to be narrow and tbe .symphysis long, particular-

ly in che large forms from Pikermi, Samos, and

Grebeniki.

Asia

Hippariims with et short and broad snout

1 here are few hipparions in Asia with as pro-

nouncedly short and broad a snout as rhe mem-

liers of the H. prost)4um/st'hlosseri~dietrichi group.

Most A.siatic hipparions, e.g. ail the Chine.se spe-

cies kepi in Uppsala described by Sefve (1927)

and ihc majoriiy of rhe specimens in the

AtVINH, hâve a narrow snout (Figs 10, 12). As

for snout length in relation to rhe distance P2-

orbit, ihere are borh long and short-snoiiicd

forms (Fig. 11). The conformation of rhe nasal

opening and the nasals show great variation

among the finds, as dues the preorbital fo.ssa.

A skull (New York. AMNH I976I) from

0.5 mile souib-wcst of Dhok Paüian, Paki.stan,

callcd H., antelopimtm Falconer ik Cauilcy, 1847

(Matthew 1929), has a medium long but broad

snout (Figs 10, 11: Svv) and a small, although

posteriorly dcep> preorbital fo.ssa placed far in

front of ihe orbit. i’hc nasal opening cnd.s at a

level 1 cm in front of P2 and is short also in rela¬

tion lo snout and skull length. ’Fhe species repre-

sented by this skull seems to be a vicar of the

268

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

Huropcan-Ncar Kastcin H, prouylumhcblosseri-

dietrichî groiip mencioncd abovc.

From Chang Chia Chuang, vShanxi, China, thcre

arc rwo skulls (AMNII 44-B 421 juv. and 44--B

427) in which the snout is quire broad (Fig. 10:

CGC). The prcorbical fossa, situared tar from the

orbit, is small and oval-pcarshaped; the nasal

opening appears .short. From 2 miles west of

Chang Chia Chuang, there are iwo skulls

(AMNH 21-B 38 and 25-B 49), the one with a

broad, the odicr with a narruw snout (Fig. 10:

CGC); ihe snoui is aiso short in relation to skull

length (F'ig. I I; CGC). Fhe nasal opening is

anteriorly pointed and deep, cxtcndiiig levcl with

P2 metaconc-F3 mcsostylc. 1 hc deep iossa is

situared close lo the orbit. Of these skulls,

AiMNH 21-B 38 with a nasal opening extending

levcl with V3 mcsostylc, rcscmbles the pcculiar

Chinese //. liechti Qiu, Huang & Guo, 1987

(Qiu et al. 1987, pis IX. XII) (Bcijing, IVPP

4 HP 20764. 2076", 20769 juv. and London,

BMNH 44377/casi of F-.AM 125708), charactc-

rized by its scrongly rciractcd nasal opening,

rcaching Icvel with Ml mesostyle, but cvidcntly

unrcduccd iiasals. l'he snout in H. Ikenti is

known only in BMNH 44577 and is lathcr long

and medium broad (Figs 10, 1 1: Hl). ’l hc pre-

orbiral fossa is double, the posterior one situared

close to the orbit (Qiu et al. 1987, fig. 20).

A skull from Nan Hao Hsia (AMNH 35*B 255)

has a medium long and broad upper snout, but

the snout of the jaw of the samc spécimen is nar-

row (Figs 10-12: NHH). The nasal opening is

very short, ending 2 cm in front of P2; the

preorbital fossa is pear-shaped. A skull from

Hsiao Kii Po (AMNH 53-B 641) resembles the

previüus one it; havmg a broad snout (Fig. 10:

HKP), very short nasal opening, and a pear-

shaped preurhital fossa.

Hippariori ùlegUfis Csxoxnovx^ 1952 from Pavlodar,

Kazakhstan, resembles some //. maithewl in

having a relaüvely broad upper snout (Fig. 10:

Pv), as well as in rhe placement and shapc of the

single preorbital fossa (Cromova 1952, cable V;

Forsten 1983, Rg. 4). The nasal opening scems

short, extending from 1 cm in front, to ihc antc-

rior tip, of P2. The fivc* mcasurcabic lowcr iavvs

correspond ro those of some H. matthewi in the

snout being narrow (Fig. 12: Pv). This species is

Rg. 13. — A, upper snout of H. dermatorhinum, showing deep

nasal opemng and long, low snoul: holotype, üppsala No. 3872,

subadull (aller Sotve 1927, table 1:1); B. upper snout of H. mot

davtcum. showing medium deep nasal opening and snout; hoio-

type, PIN 1256.^3639 (after Gromova 1952, table I 3). C. upper

snout of H. giganteum. showing very short nasal opening: holo¬

type. OGU 1015 (after Gromova 1952. taole 12: 2). Drawings

not to scale

an Asiatic vicar of rhe dwarf H. mattlmvi, from

which it differs in the morphology of its teerh

and in the proportions of its limb bones.

Hipparions with a narrow snout

Extrême among the Asiatic hipparions with a

GEODIVERSITAS • 1999 • 21 (2)

269

Forsten A.

long and narrow snour is H. tchicokum Ivaniev,

1966 (Qiu étal 1987, pL XIII); the snout is nar¬

row also in the lower jaw and thc symphysis long

(Fig. 12: Htc). The nasal opening is vcry short,

ending at a Icvcl almosç 2 cm in front of P2; it is

short also in relation to snout lengch. The large

prcorbital fosvsa is evidently situated close to the

orbit (orbit not visible).

Hipparion dermaiorhlnum Sefve, 1927 from

China has a long snout in relation to skull length,

and the snout is low and narrow (Sefve 1927,

table 1: 1-3; Qiu et ai 1987, pis XLl, XLli)

(Figs 10, 11: Hd). In thc lower jaw thc snout is

similarly narrow and ihe symphy.si.s long

(Fig. 12: Hd). The nasal opening is retracted,

rcaching Icvel with P3, it is long also in relation

to snout and skull length; the nasals are unreduc-

ed. The prcorbital fossa is well-defined but shah

low.

An upper snout (rom Sor, I*ad/hikistan, shows

the pre.sencc ol a double preorbiiaJ fossa

(Zhegallo 1978, Ris. 73). The narrow snout

(Fig. 10: Sor). thc nasal opening extension, and

the double fossa closely resemble those in the

specimen R-5I from Bazaleti.

Hipparions lavking a preorhitu! fossa

A skull (PIN 2433-360) from Kalmakpai,

Kazakhstan, referred lo as H. Inppidiodus because

it lacks a preorbital fos.sa (Zhegallo 1978,

Ris. 61), lus a long, narrow snout but may he

laterally cruslicd, since a second specimen

(PIN 2433-460) from thc saine loculiry has a

medium broad .snout (Figs 10, II: Kp). In boch

specimens the nasal opening ends Icvcl with the

meracone oF P2. Originally //. hippidiodas was

described from Loc. 115, Kingyang, China

(Sefve 1927; f'orsten 1968, pl. .3; Qiu et ai

1987, pl. XXXVIIl); irs snout is unknown.

Skulls from Khirgis Nur-2. Mongolia, called

//. mogoicani Zhegallo (Zhegallo I97S)> also lack

a fossa. The snout in rhe mcasureable specimen

is medium broad like in PIN 2433-460 from

Kalmakpai, but sliorcer (Fig. lOi KN). The nasal

opening rcaches level with P2 mesostyle.

Chinese hipparions belonging in rhe species or

specics groLip Prolwscidipparioa rocinantis-houfe-

nense (HernarideZ'Paclieco)-(’Icilhard & Young,

1931) with advanced, caballoid cheek teeth and

lackinga preorbital fossa (sec above), bave a long

snout in relation to skull length and the snout is

narrow (Qiu et al. 1987, pis VI. VU; Forsten

1997, fig. 7) (Figs 10> 11: Hr); also the snout of

the lower jaw i.s narrow (Fig. 12: Hr). The nasal

opening is short, extending From 0.5 cm in Front

oFP2, to level wirh P2 paracone.

There are four skulls oF rhe rwo (?) species oF

cypical Proboscidipparkaiy P, sinense Sefve (Sefve

H)27, table VI: 22-24, V[l) and

Marsumoto, 1927 (Qiu e( al 1987, pis l-IV).

l'he snout is rather shori in relation to skull

lengch and narrow (Figs H). Il: Pp, Ps and Ps in

bold). Also thc symphysis is short, taking imo

account the size of rhe jaw, and thc snout is nar¬

row (Fig. 12; Ps and Ps in bold). The nasal ope¬

ning is enormou.sly retracted, extending from

level with Ml rnctacone to M2 mesostvlc. and

the nasals arc rcdiiced and foreshortened. There

i.s no fossa.

DISCUSSION

Phylogenetic and functional

c:ONSrDERATIONS

l'here arc différences berween local forms (spe-

cie.s, suKspecie.s, local populations) of hipparions

for the proporiions of thc snout and chc exten-

.sion of the nasal opening, but vviihin a local

form chese characters arc quitc stable. Characters

of rhe snout and nasal opening occur in combi¬

nation with different shapes and placement, even

absence, of chc preorbital fossa. Snout breadih:

length proportions are nor corrdated with rhe

extension of rhe nasal opening. Flow to deliniit

species and .supraspeciffe groiips when characters

of the snout, nasal opening and nasals, and

preorbital fossa arc di.stributed mosaic-likc, a.s are

characters of thc teeth ,and linib-bone size and

proportions? Whai do rhese characters and the

différences berween forms signifÿ^ Do they bave

phylogenetic and functional significance? Which

characters should be given more, which less

weight when phylogénies are discu.ssed and

reconstructed? How lo déterminé which charac-

ters are primitive, which derived, i.e. rnorpho-

clinc polaritics?

The hipparions {Hipparion, Prohoscidipparion

270

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

Sefve, 1927, Neohipparion Gidlcy, 1903,

Narmippus Mauhcw, 1926, and Ihcudhipparion

Ameghino, 1904) can be disdnguished froni the

ancestral merychippincs on tlic basis ol the chcck

tecth in the lower jaw, particulaily ihc lowcr

molars. 1 he lowcr molars oï the mcrychippines

retain a primitive, low double knot with litllc

differentiated and rounded mecaconid and

metastylid loops, while in the hipparions rhe

loops of the molar double knor are well difleren-

tiated, rhus advanced, Ail Fiirasian and African

hipparions are truc hipparions, with welhdiffe-

rcnciatcd loops ol the double knot of ihe lower

molars, indicating dispersai to the Old World at

a stage when ihe dental pattern already had be-

comc moderni/cd.

In the hipparions, a short snout and shorr nasal

opening could be cotisidered primitive, since

rhey characrcrize the merychippine ancestors,

c.g. ^Neohipparion coloradense' (originally of

Osborn, 1918) froni the early Barstovian ol

Boulder (^uarry, Nebraska, and Lue Barstovian ol

NE Colorado (MacEadden 1984, figs 52, 54, 58.

59). and Cnrnwhippiirwn' goorisi MacEadden &

Skinner, 1981 (rom the laie Barstovian of IVinity

River Fit 1, Texas (MacEadtlcn 1984, figs 121,

123). On the other hand, if a shon-broad snout

in a hipparion is an adaptation to gra/dng, it

sbould be considered a late, advanced character,

since gra/ing evolved wiih rhe spread ol c4

grasses about 7-5 Ma ago (C-erling et ni 1993).

Sonic merychippincs, alihough dentally primi¬

tive, already had a rachcr long snout, e,g.

"'Eoequits" wiboni Quinn. 1955 (Quinn 1955,

pis 10: 1-3, 11: l'3, 12, 13 ), which although

large!' is prohably closely related to '’dïippnrion'

shirleyi MacFadtlen, 1984 (boih are froin Texas

and Barstovian in age; MacEadden 1984, figs 28,

29). On the other hand, sonie North American

hipparions or near-hipparions, although dentally

advanced, retained a short snout and nasal open-

ing, c.g. Ncohippntion nfjinc" (originally of

l.eidy, 1869) froni che late Clarendonian of

Upper Miller Quarry, Nebraska (Machadden

1984, fig. 63). ^'Gnnwhippnrhnr sphenodus (ori¬

ginally of C^ope, 1885) from the Valentinian of

Railway/llailroad Quarry, Nebraska (MacFaddcn

1984, figs 124, 126), and ^Cornujhipparion"

occideritale (originally of Leidy, 1856) from the

late Clarendonian of Gidiey Horse Quarry, Texas

(MacEadden 1984, figs 133, 136). The breadth

of the snout in relation to snout length is not

given in thèse hotscs.

'Elle nasal opening *m che North American hippa-

nons does not extend posteriorly beyond rhe

Icvel of P2, but otren ends ar a level 1-4 cm in

front of P2 (MacEadden 1984, table 1).

Ahbougb in the Eurasian hipparions the nasal

opening may bc cqually short, c.g. In early

H, prirnigeniurn^ the snout is gencraJly longer in

relation to skull length. In the H. prostyluni/

schlosscri-dietrichi groiip bot h rhe nasal opening

and snout are short. In the Eurasian hipparions a

deep nasal opening, posteriorly extending to a

level beyond P2, bas evolved several rimes and in

combination with diftercnrly shaped, even

absent, prcorbiral fo.ss.ie, e.g. in the large

Hipparion from Samos Ql-Andriano and

Pikermi, in il. prohoscidcurrn H. licenti, H. der-

rnatorbiniim, and Probosddippnrion smeme. The

deep nasal opening in somc hipparions has given

risc io hypothèses that thèse horses had a tupir-

like pruboscis (Sefve 1927; Solounias &

Dawson-Saunders 1988). In extant mammals

with a lengthencd nosc, e.g. tapirs, dépliants,

saiga and dik diks, rhe nasal bones arc forcshortc-

ned, thus allowing mobility of the nosc

(Macdonald 1984). In most of rhe known hippa-

rions vvith an cxccptionally deep nasal opening,

the nasals arc nornially long. Only in some

Proboscidipparion (sintnse and ^^pater') arc the

nasals reduccd in length, the nasal opening bet'

ween che premaxillas is narrowly slit-like, and the

upper symphysis long. Sole among the hippa-

rion.s tfie .snout in these Proboscidipparion ducs

resemble thaï in a tapir (Sefve 1927). However,

the deep nas'al opening and foreshortened nasal

bones in these Proboscidipparion also resemble

those in a nioose, Àlccs alces (Linnaeus, 1*^58),

wliich has no probu.scis but a large overlianging

upper lip. Extant horses, which bave a modera-

rcly deep nasal opening and long nasals, hâve

very mobile bps. 'l'he hipparions may, to a

varying dcgrce, also bave had mobile bps usefui

for gathering grass or browse.

In tlie extant horses rhe diverciculae nasi form

hliiid sacks in cotinectioti with the nostrils and

occupy the nasal opening (Ellenberger & Baum

GEODIVERSITAS • 1999 • 21 (2)

271

Forsten A.

1943; bue see Gregorv 1920). The longer and

wider the nasal üpening> the more room for che

diverriculae. According to Gregory (1920) the

diverticulae in exranc Equiis (Linnacus. 1758)

partly occupy the shallow subnasal {= preorbiral)

fossa. In rhe liipparions nasal opening extension

and dcvelopnicni oF che fossa are not correlated.

Well-develoj'îcd and posteriorlv deep in the large

hipparions from Samos Ql-Adriano and

Pikermi, rhe fossa is small and situared far from

rhe nosrrils in // hcefitty and is absent in

Probosàdippnrhm sittense and 'épater'. The diver-

ticulae comprise rhe vomeronasal organ wirh sen-

sory pachways ro rhe hypothalamus. The

vomeronasal organ is believed to he involved

wirh rhe "f1ehmen’\ Fr olfacrorial tïistmg of fero-

mones (Estes 1972), and wiih cerrain low vocali-

zations.

The gencraLly narrow snoui in Old Woild hippa¬

rions may hav^e to do wirh thèse horscs having

been chiefly browsers or mixcd-fccdcrs (Hayek ei

al. 1991). Candidates for che gr.r/.ing niche

woLild bc the broad-snoLited founs in the

H. prostyluw/schlosseri-dietrichi cornplcx and

H. antelopinum. Eiscnmajin (1998) rccently

plotted Icasr diastemal breadrh to symphysial

lengrh in josvs of Old World hipparions. She dif-

ferentiated specimens wirh a broad and those

wirh a narrow jaw (F.isenmann 1998, figs 3, 4)

and discusscd jaw (>roportions and incisor mor-

phology in relation to mode of feeding (whether

grazing or brovvsing). Wirh a few exceptions hcr

diagrams correspond ro mine» plorting lowcr

snour widih tu symphysial length. The excep-

rions are specinieris probably wrongly identified,

e.g. in her ligure 3 Ntn 7, 8 are evidenriy

H. verae not gip^aiitcuyn , and No. 9 is not

H. medhmaneuni.

TAXONOMU' CONSlDhR/VnONS

In hipparion taxonomy and phylogeny» the

preorbital fossa is ofeen given more weighr than

orher characters» sometimes even ac che near

exclusion of orher characters (si;e Eisenmann et

ai 1987). .Solely on the alleged re^iemblance of

the fossa» the narrow-snouted IL getty} ( =

H. tudoroveine^.) and H. campbtdlï {-

H. timnensd) witb a long nasal opening arc said

to be the ancestor and descendant, respectively,

of the short and broad-snoured H. prostyltim

with a short nasal opening (Hernor et ai 1996).

The member.s in ihe H. prostylum/scldosseri-

dietnchi group arc helieved to hâve given i ise to

hipparions lacking a preorbiral fossa, although

tliere are no signs rhac the fossa did disappear in

rliis group. Hipparioa hippidiodudplatygenys from

Tarakiiya, ihe carliesr known hipparion lacking a

fossa, has a medium broad .snout and is of the

samc âge as rhe members in che former group,

while rhe hipparions from Khiigis Nur-2 aiid

Kalmakpai, also lacking a fossa, may bc younger.

Thcir nasal opening is' deep and long. On the

supposcd similariry of che fossa the long-snou-

red, lai^c hipparion from Samos Ql-Andnano,

wirh a deep and long nasal opening, is teferred to

as H. gignyitemn (Bernor et ai 1996), the type

form of which has a very short nasal opening.

On the oihcr hand, H. macedoakuni is helieved

to represent a .spccies different Irom H. rnntthewi

(Koufo.s 1986; Bernor et ai 1996), although

Identical to ihat species in skull and jaw rnorpho-

logy, meiapodial si/.e and proportions, and in

protoconal length and pUcaiioti coiint of its

lecth. Sonic of thc.se conilicting idcnliflcaitons

seem lo hâve been dictated by the presumed

stratigraphie position ol che fmd, rather chan by

les morphology. However, stratigraphie position

is nor a taxonomie character.

HK-I-AKKiN Mt'i.niM.t niSr-ERSAL?

Old World Hipparion has been thought ro repre-

seni multiple dispersais from North America,

rfits idea was also based on preorbital fossa mor-

pholog)'*. ignuring the morpholog}' of the cheek

reeth and snout, and the faunal evidence against

multiple equid dispersai. In rhe Miocène the

Notvh American equid fauna was diverse, com-

prising anclhtheriines, para-merythippines, plio-

hippines, and hipparions, often sevcral généra of

each group in the same faunas (Forsten 1989).

Of the anchilheriines only Anehitherium

y. Meyer, 1834 dispersed to the Old World, iiojic

of the parwmerychippines or pliohippines made

ir, and there is no evidence thar more than one

hipparion successfuily crossed the Bering intcr-

coiuincnial connection. None of the lypical

North American hipparionid (,sub)gcncra

Pseudhipparion^ Neohipparion, or Nannippiis has

272

GEODIVERSITAS ■ 1999 • 21 (2)

Snouc proportions in some Eurasian hipparions

ever been found in the Old World. Ncithcr bas

any of the scveral Eurasian hipparions with a

deep nasal opening ever been found in Norrh

America.

CONCLUSIONS

It is dangerous to base hipparion taxonomy, sys-

tematics and phylogeny, and hypothèses about

intercontiiieniul dispersai, on single charactcrs,

e.g. on the preorbital fossa alone’ (F.iseninann et

ai 1987). Skulls are rare, mosi fos.sil samples

consist of isolated teerh and lîmb bones, which

in mulrispecies samples may be difficült/impos-

sible to übjettivel) couple with the skulls or

idencify to spccies. The more character.s thaï arc

taken into account, the more düficult is the déli¬

mitation of speties and lineages, l)ecausc ol ihe

mosaic combination of charactcrs (sec also

Hiscnmann 1998). I bclicvc that gcncra of equids

should bc characterizcd on their check teeth. par-

ticularly on the cheek tecth of the lowcr jaw.

Wirhin a genu.s occur subgroup.s (.spccies groups,

in some cases worthy of subgcncric .séparation),

which sometimes can be circumscribcd on .seve-

ral charactcrs m common, e.g. plicaiton count of

the upper check tecth, proportions of the limbs,

preorbital fossa nu)^pholog)^ and snout propor¬

tions. Many tlnd.s cannot bc placed wiihin a

single subgroup, as ihcy .share charactcrs with

-several. 'l'here is no reason why the preorbital

fossa should hâve priority in determining sub-

groiip membership, but neither has any other

character Référencé to spccies or species group

should be based> îf possible, on the holomorph

of the find.

Stratigraphie provenance is nor a taxonomie cha-

racter. Stratigraphie schemes, using species of

fossil horses as markers, stiould be regarded with

suspicion. More often than not they arc idcalized

and do not truly reflect the time range of the fos-

sil spccies, whïch olten occur at an earlicr date

and/or survive for a longer timc than presuppo-

sed by the schemes. Subjecrive ^‘morphologieal

trends’’, often eonsciously or uneonsciousiy

constructed to fil and/or to support allegcd strati¬

graphie successions, are cqually suspicîous.

Fossil cquids should not bc forced to fit a strati-

graphy, but should bc objectivcly interpreted

against the background of stratigraphy.

Acknowledgements

My sincère ihanks arc duc the curators and carc-

takers of the fossil collections studied for this

paper (sec Appendix). I also thank Dr. V.

Eisenmann and an anonymous reviewer for usc-

ful comments on the manuscripi.

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Submitted for publication on 2 April 1998;

accepted on 31 Jnly 1998.

274

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in some Eurasian hipparions

APPENDIX

List of hipparion specimens used in this paper.

Listed are only specimens, which hâve yielded

measurements on the upper and/or lowcr snout.

Listed are not specimens lacking or vvith frag-

mentary snout, although represcnting the same

Hipparion sp.

(wiili double preorbiiai fossa), not named.

LocAiJTY. — Sanios Ql (and Q5?) and Samos

wiihouc exact localiiy data.

Ace. — Turolian.

Matekiais. — Samos; (AMNH 20628. probably

205V9, 22908, 94905 & no No., ail Ql) iWc skulLs,

(AMNH 94906 Ôf no No., Ql) skull and jaw, possi-

hly aiso (AMNH 207S6 22912. Q5): (SMI- no

No. and possibly no No.)- two jaws; Geologieal

Muséum, Lausanne (No. 15 K 61 (J, 843, Andriano)

thiec juws; Garncgic Muséum, Pittsburgh (No. 2 I 26)

a jaw.

//. maubvîvi or H. sp (with a double lossa): Sainus

(AMNH 22928, Q5) jaw; (SMI* no No.) two jaws;

(MNilN 1955-1 7.^)ajaw.

H. sp. (wiil> double lossa) or H, ichlosseri-dietrichr.

Samos (AMNH 20651, 20721 & no No.. Ql; No.

10733) Tour jaws, Geol. Mus., Lausanne (Nos. 78 &:

1031, Andriano) two jaws; Nat. Hist. Mus., Vienna

(1911 V 135)ajaw'.

Hipparion sp.

not named, not H. mediterraneum.

L(x:alit\'. — Piera, Spain.

Ac.E. — Turolian.

Mai'HRIALS. — Institut Paléontologie Dr.

M. Crusafont, Sabaddl (IPS.P 6359, 6360, 6365,

6372, and two specimens withouc a readablc number)

skulls.

Hipparion afiicanum Arambourg, 1959

Type LOCALITY. — Oued el Hammam, Algeria.

Ac;e. — Vallesian.

Materials. — (MNHN 141 & no visible No.) two

skulls, (MNHN 89, 98, 143) three jaws.

Hipparion antelopinum Falconer (^utley, 1847

Type EOCALI TY. — Porwar Plateau, Siwaliks, Pakistan.

Ace. — Turolian.

Material. — (AMNH 19761) a skull.

spccies, from the same localities, and présent in

the same collections. When studied in the

1960-ties, most collections were still inadequate-

ly filed, as reflected by the frequent lack of col¬

lection numbers.

Hipparioyi brachypus Hensel, 1862

Type I OCAUTA'. — pikermi, Greece.

Age. — Turolian.

Materials, — Pikermi: British Muséum of Natural

History, London (BMNH unreadable No.) skull,

(BMNH 11211, 1 1213 & 11217) three jaws;

(MNHN no No.) skull, (MNHN 31 & rhiee .speci¬

mens withoiit a number) four jaws; Geologieal

Insncucc, Gorungen (no No.) skull; Swedish Muséum

of Naiural Hi.siory, Stockholm (no No.) jaw;

Humholdt Muséum, Berlin (no No.) skull; Yale

IVabody Muscutti ol Natural History (No. 11768)

skull with jaw,

In -addition; Natural Hisior)' Muséum, Bascl (PK 89)

a jaw of ciiber H. hnnhypus or H mrdUrrtancum.

H, alf hrttchypHi' Samos; (AMNH 22838, Q4) a

skull, pos.sih!y (AMNH 22912 & 22922, Q5); (SMF

4707) a skull, (SMF no No.) a jaw; Geol. Inst.,

Budapest (ÜK 425) a jaw; Nat. Hisi. Mus., Vienna

(No. 1911 V 1 18) a skull: (jCoI. In.st., Lausanne (No.

73 Ôc 148, Andriano) skull jaw, (No. |75,

Andriano) a jaw; Hessisches Landesmuseum.

Darmstadt (Ss 45) a jaw. — Beluska, Macedunia (lor-

mer Yugoslavia): Prirodnjacki Mu/cj, Beograd (PM

2791) jaw. karaslari. Macedonia: (PMM 193/73)

jaw. — Maraghch, Iran: (MNHN 561) a jaw.

H. aft. hniibypns or H. prnlmcideimr. Samos: (AMNH

20640, Ql) a jaw; Inst, of’CJcoL &c PaleoncoL,

MünsitT (Sl/no No.) a jaw.

Hipparion cataiaunîcum Pirlot, 1956

Type \ vX:AIJTY. — Hostalets de Pierola, Spain.

Age. — Vallesian.

Materials. — (BMNH 16397, type //. catalan-

nicum) a skull.

Hipparion dennatorhinum Sefve, 1927

Twf. LOCAlTTY'. — Loc. 30, Bao De, Sbanxi, China.

Age. — Turolian.

Matekiai5. — Loc. 30: Paleontol, Inst., Uppsala (M

3872, type) skull. — Huang Lu Kou, China (AMNH

41-L 310) skull and jaw. — Bao De, Sbanxi, China:

(IVPP no No.) skull and jaw, (IVPP 8243?) skull and

jaw, (IVPP no No.) jaw.

GEODIVERSITAS • 1999 • 21 (2)

275

Forsten A.

HippaHon elegans Gromova, 1952

Typk LOCAI.ri Y. — Pavlodar, Kazakhstan.

Age. — l’umlian.

Mat erials. — (PIN no visible No.) a skull, (PIN

2346-2479. -2516, -4883, 2413-2847, -2864) five

jaws.

Hipparioft garedzicum Gabuniya, 1959

IVpe UîCAl.l'i'Y. — Udabno, Georgia.

Age. — I.ate Vallesian-Early Turolian.

Materials. — Musei Grusii, Tbilisi (156/13 type

H. garedztmm 270/34} c\vo skulls.

Hippainon pgnntettm Gromova, 1952

Ty]‘E LOCALITV. — Grcbeniki. Moldova.

Age. — Early Meotian/Late Vallesian/liarlv l urolian.

Materiajs. — (OGU 1012/2, 1015 type & 1017)

rhree skiilU, (GGll 10 18) skull and ja\v, (OGU

unreadable No.) a jaw.

In addition OGU 908, a skull, is intermédiare bet-

ween H. verae and H. giganteum.

Hipparion bippidiodiis 1927

Type lo(L\Lity. — Loc. 115, King Yang Hsien,

Gansu, China.

Age. — ?'l urolian.

Materials. — Kalmakpai, Kazakhstan: (PIN 2433-

340, -360.-340) ihrcc skulls.

Hipparion licentiQjw^ Huang & Guo, 1987

I’YPK Lt)c:ALITY. — Sianiiotsun, JYliaiigkou, Yushe,

Shanxi, China.

Age. — Ruscinian.

Material. — (BMNH 44577, cast F:AM 125708)

skull.

Hipparion matthewî Abel. 1926

Type locaI ri V. — Sarnos w'irhour exact locality data.

Age. — Laie Vallesinn to and including l urolian.

Materials. — Samos. Greece; (AMNll 22907 &

22936, Q5) two skulls. (/VJVINH no No., Q5) a jaw;

(SMF 4/10 & no No.J two skulls. (SMF no No.) u

jaw; Nat. 1 list. Mus., Vienna (no No.) a skull: British

Muséum oF Naiiiral Hisiory, l.ondon (BMNH

14071, casi of type) skull tfc jaw; Geol. Inst.,

Budapest (t'ïK 557, type oF //. manl^rwi) skull and

jaw. — Itaviii de Pluiéi Circece; Univer.slty of

Thessaloniki {RPi-21 type H. macedonicum Koufos &

RPl'36) two jaws. — Beluska, Macedonia (former

Yugoslavia): Prirodnjacki Muzej, Beograde (PM

2659/197 & 2751 ) two jaws.

Hippariot! inediten'aneinn (Roth & Wagner, 1855)

Type i [»C-Ai rn. — Pikermi, Greece.

Age. — Turolian.

Materiai.S. — Pikermi (BMNH 11215) jaw;

(VINHN no No.) skull and jaw. (MNIIN 514-31 &

no No.) two jaws; Nat. Hisi. Mus., Vienna (no No.) a

skull, (no No.) a subaduir jaw; Universiry of

Caldornia Muséum, Berkeley (LIf'M 63422) a jaw;

US National Museutn, Washington, D.C. (No. 267)

a jaw; Swtdish MuseioTi ul Natural I^istory,

Stocldinlni (ncr Nod two j.i\v.s; Cie<ilogit:al lustitule of

rhe Universiry, Gôtringen (no No.) a jaw; Naruial

FTsrory Muséum, Stuttgart (no No.) jaw to skull.

H, cf. medhenvncunn Bazalcii, Georgia: Insiiiute of

Paleobiology, I blJisi (B-51) skull, (B-54) a jaw. —

Manigheh, Iran: (MNIIN no No.) a jaw; Nat. Hist.

MuseLKiL Vienna (no No., Ketchawa) jaw; National

Muséum, Praha (no No.) jaw.

Hipparion ntogoia/m /hrgallo, 1978

Type, I.üCALTTA^. — Khirgis Nur, Mongolia.

Agi*. — Turolian.

Materials. — Khirgis Nur: (PIN 3222-193 type &

no No.) two skulls.

Hipparion 7nolayanefise7^o\x\\x\, 1992

Type LOCAILIT. — Molayan, Afghanistan.

Age. — Turolian.

Matkriaia. — (MNHN Mo 040 & 1758) two skulls.

Hipparion moldavicum Gromova, 1952

Type IOCAI I T Y. — Parakliva, Moldova.

Age. — Meorian/Turolian.

Materiai.S. — Tarakliya: Palcomological Institute &

Muséum, Vloscow (PIN 1256-2922, -3639 type,

-3647, -3648 Sc no No.) ihe skulls, (PIN 1256-3619,

-3620, -3638. -.3641, -3642, -3643- -3'’00, -4189,

-4191, -6605,.-6944, -7027 A; four withour a mun-

hcr) jaws, — Novocli/jvetovka, Ukraine: (OCî-U

1233, 1.306, 1307, 1313, 1314, 1459,. 3369. 3371)

.skulk (OGU 1394^95, 1401, 1403, 1404) j-aws. —

Cdieicvichnoc, Ukraine: Dcpi. Paleonrol. &

Pnleomoi. Mus.. Kiev: (No. 45-2(i65, -3849. -3925)

rhree .skulls. - Tchobrudii. Moldova: (OGU 3081)

skull. (OGU 30781 two jaw.s; Mus CdrdvonikitÎAe,

Moscow (No. 2080 & no No.) two jaws, (No. 2026

ik 2081) ,1 .s-kull and ja'v m.iy be Liiher from

Tchobruchi or Grcbeniki.

H. aff moldavicum: Novaya Emefovka-2, Ukraine:

276

GEODIVERSITAS • 1999 • 21 (2)

Snout proportions in somc Eurasian hipparions

Depr. Paleontoi. Paleonrol. Mus., Kiev (No. 25-

2439, '2923, -3005, *3200, -3310) skull.s. -

Chimishlia, Moldova; In.sricute of Geology àc

Paleontology, Uiiiv. Bucharest (No. 66/(378), 378a

&: no Nu.) thrce skulls, (no Nu.) jaw: Kegianal

Muséum, Cliisiiiui (No. 44040/79 & -/8I) jaws. ^

Maragheh, Iran: (RMNH 3924) skiill; (MNHN thrce

specimens vviihoLii No3 chree jaws-, Nai. Hi-tc. Mus.

Vienna (no Nt)., Kopran Si Kerchiiwa) two faws.

Hipparion platygcnys 1952

(?younger synonym ol H. hippïdiodii'^ .SefÀ'e 1927).

Tvpl- LOCAUTY. — Tarakiiya, Moldova.

Age. — Meotian/Turolian.

Materials. — (PIN 1256-2942, -3634 & no No.)

ihrcc jaws.

Hipparion praegigantctim farabukin, 1967

TvT'I', — Chimishlia, Moldova.

Age. — Mcotian/l'urolian.

Mai ekials. — Kcgionul Muséum, Chisnau (No.

4040/84, type) skull, (No. 4040/83) jaw.

Hippario7t pHmigetiium (v, Meyer, 1829)

Type l-OCAi.rrY. — Eppelsheim, Germany.

Age. — Vallesian.

Materials. - F.ppelsheimî Geol. Inst., Budape.sr

(OK 22) a jiiw. -- 1 lowenegg, CJermany: llumboldt

Muséum, Berlin (no No., casr) skull and jaw;

Hcssisehcs 1 .nndc.sinnscutn, Darrnsiadi (Ho 58/VI)

skull and jaw, (Ho. 54B) a faw. — V'Üa de Caballs,

Spain: (IPvS no No.) skull and jaw. — (]an Uobateres,

vSpaln: (IPS no No.) jaw. — In/.ersdoi4, Austria:

Natural Htstory Muséum, Vienna (SK 1346 Sc no

No.) tw<3 skulls, (No. 1875 V^I 5) a jinv. —

Wienerberg, Austria: Nar. Hî.sc. Muséum, Vienna

(1842 LVIl 11) a jaw. — J’rottes, Austriar Gcologital

Instiiutc, Univ. oT Vienna (No. 958) a jaw. —

Balravar, Hungary: Geol. In.st., Budapesr (No. 90) a

jaw; Nacural Hiscor>' Muséum, Budapest (No. 319b,

390, 391) chree jaws. — Nc.sebr, Bulgaria: Geological

In.sritute, Univ. Sofia (No. 130) skull, (No. 40, 133 &

no No.) three jaws.

Hipparion proboscideum Siudcr, 1911

'rwv, l.oCAi.n V. — Samos vviihoiit exact localirv data.

Age. — 'f'urolïan.

Materials. — Samos: (AAINH 20771, QI. AMNH

20772, QIs) two skulls; (SMP 4706, 4708, 4709)

three skulls; (SI/4) a skull, (possibly SI/208) a jaw;

(possibly BMNH 4359) a jaw. — Ravin des Zouaves:

Univ. Thessaloniki (RZO-60) skull.

H. sp. (cf. proboscideum):. Chcrevichnoe, Ukraine:

Depr. Paleonrol. Paleonrological Muséum, Kiev

(No. 45-2664, -4282) two skulls.

II. proboscideum or II. sp. (widi double fossa), Samos:

(Sl/277) a juw.

Hipparion prostylum Gervais, 1849

d’VPE LGGAI.M Y. — Mr. J.ébéron, France.

Age. — d'urolian.

Materials. — Faculté des Sciences, Lyon (no No.) a

skull with jaw; l’acuité des Sciences, Montpellier (no

No.) a lower snoui.

Hipparion prostylum!H scblosseri-dietidchi

an entily nut well defined, resembliiig both species.

Type LOC AII IY. — None, but présent in Southern

Macedoniu (lormer Yugoslavia) and nurthern Greece,

possibl) also in Maragbeh, Iran,

Age. — Turoiian.

M.ATERTALS. - Salonikk Greece: (Coll. Arambourg Si

Puvhaubcrt) Miisétim National d’Hisroire Nacurdle.

Paris (MNHN 1919-8) rwn skulk (MNHN 1911-

23) a jaw. Ravin des Zouave.s, Greece: Univer.siry

of Thessaloniki ïRZO-76, -105, -145. -154) Four

jaws. — Karaslari, Maccdonia (fonner Yugoslavia):

Prirodonaueen Muzej na Makedonija, Skopje (PMM

86/73, 93/73 subadulr Si 203/73) skulls, (l’MM

410/73) a jaw.

in addition: Llnien Doi, Macedoni. 1 , (PMM 99/60) a

.skull of elther H. mauhcwi or H. prostylum!H.

scblosscri-dietricbi. — Maragheh. Iran: (MNHN three

specimcivs without No.) three skulls.

Hipparum saMnatiewn L ungu, 1973

Type LOCAI ri y. — Kalfii, Moldova.

Age. — Middie Sannarian/Vallesian.

MaJERIAIS. — KalFa: Tiraspol Pedinstitut, formerly

Tiraspok now Cdiisnau (no No.) fivc jaws. — Brada,

Moldova: (PIN 646-12) a skull.

Hipparion schlosseri'dietrichi

AnconiiLs, l919'(Wehrli, 1941)

Both trames were given evidenrly the same species;

Anrontus’ nanir is oldcr. but possibly nor valid (des¬

cription Incomplète, type not figured). Wchrli used

the genus namc Hemihipparion.

4'ype I.OGAl.rn . — Samos without exact localicy' data.

Age. — Turoiian.

Materials. — Samos, Greece: American Muséum of

Natural History, New York (AMNH 20596, 20598,

20608, 20692, 20997 & two without number, ail

Samos Ql) seven skulls, (AMNH 20603, 20650,

GEODIVERSITAS • 1999 • 21 (2)

277

Forsten A.

20655, 20667, 22787, ail Ql) five jaws, (AMNH

22860, Q4 6c no No., ?Q4) cwo skulls, {AMNH

22990, Q6) skull; Senckcnber^ Muscum, l'rankfuit

(SMF no No,) thret jaws; Nutiiral I üstoty Miiscum,

Vienna (No. 191 ! V 114, type H. schlossert); possibly

Geological Jnstirute, Budapest (No. 274) skull;

ïnstitiite of Geology & I^aleontology, Milnster (Sl/7,

type HernUnpparum dietrichi 6c Sl/28) iwo skulls, pos-

sibly (Sl/236) a jaw; Teylers Muscuni, Haarlem (No.

15470 & no No.) nvo skulls; Geological Muséum,

Lausanne (No. 132) skulls (No. 6c 195 Irom

Samos, Andriano) rvvo jaw.s; University of Texas-

Ausiin, Bur. Econ. Geol. (No. 40275 cast of CMNH

P. 12868) skull. — Maragheh, Iran: Nat, Hist. Mus.

Vienna (no No., Kopran) jaw.

Hipparion tchicoiewn Ivanjev, 1966

Type LOCAI I TY. — Beregovaya, Russia.

Age. — Vilkifranthian.

Materials. — Shamar, Mongolia: (PIN 3381-53)

jaw. — Peihaitsun, Maesegou, Yushe, China: (IVPP

THP 19009 & 19013) skull and jaw. —Yid-

jouantsun, Yushe, China (IVPP THP 10302) jaw.

Hipparion hulorovense Gàhnn\yd.s 1959

Type LOCALi n'. —Tudorovo, Moldova.

Age. — Meorian/Turoban.

Materials. — (OGU 1780 type //. tudorovensé) a

skull, (OGU 906, 1783 & no No.) three jaws.

Hipparion t>^#ï^Gabuniya, 1979

(originally H. ^romovae Gab. 1959, preoccupied by

H. ViTlalta & Crusafont, 1957)

Type i üCAI h y. — Grebcniki, Moldova.

Age. — Early Meotian/Late Valle.dan/Larly '1 urolïan.

Materials. — Grebcniki; University of Odessa

Muséum, Odessa (OGU 916, 917, 1012/1 & 1016

type) four skulls, {OGU 889, 897, 898, 905, 1016,

1462 & no No.) seven jaws; Department of

Paleontolog)' & Palcontological \luseum, Kiev (No.

408-47 6c 408-114) two jaws; Muséum Ordjo¬

nikidze, Mosçow (Nos. 2027, 2051, 2079 6c no No.)

lour jaws, (No. 2061) a skull, said to bc from

Tchobruchi, resembics H. vetetc.

H. aff. vèrae\ Bekuskà, Macedonia (former Yugoslavia):

fPM 2660/196) skull, (PM 2743/195) jaw.

Proboscidipparion Matsumoto, 1927

Type locali iy, — unknown.

Age, — Ruscinian-Villafranchian.

Materials. — Nihersun, Yushe, China: (IVPP THP

20763) skull, (IVPP THP 30756) jaw. — Loc. 26,

Pcihait.sun, V'ushe, China (IVPP THP 14312 lecto-

rype) skull and jaw.

Proboscidipparion rocinantis

(Hcmandcz-Pacbeco, 1921)

Type IOCAI ITy. — Puebla de Almoradier, Spain.

Age. — Villafranchian.

Ma ITRIALS. — Villaroya, Spain: (IPS 2085) a skull,

(IPS V 196, unreadabie No. & no No.) three jaws. —

Kvabebi, Georgia: Inst. Paleobiol., Tbilisi (K-48) a

skull. — ühamar. Mongolia: (PIN 970/2086) a

jaw. — Peihaitsun, Matsekou, Yushe, Shanxt, Cliina:

Institutc of Verrebrate Paleontology & Paleo-

anihropology, Beijing (IVPP THP 10331 6c 10508)

skull, skull and jaw, (IVPP THP ]0()97) a jaw. —

Yinkiangtsun, Yushe, China (IV'PP l'ill* 10733)

skull and jaw. — Hsi Chwang/Hsiao Chuang, Shanxi,

China (AA4NH 96-B 1031) skull and jaw, (AMNH

64-B 815) skull.

Proboscidipparion sinense Sefve, 1927

Tvpl. l.OCAl I LY. — Mien Chili Hsien, Lankou,

Henan, China.

Age. — Villafnuichian.

Materials. — Mien Chili Hsien: Palcontological

Institute, Univ. Uppsala (M 3925 & 3926 type) skull

and jaw. — Yushe, Shanxi, China: Beijing Natural

History Muséum (no No.) skull and jaw.

278

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Références bibliographiques

Denison R. H. 197H. — Placodermi, in .Schulr/,e

H. P. (cd.), Handbook of PaUfiichthyology^

Volunte 2, Gustav Fischer, Stutigarr, 128 p.

Marshall C. R. 1987. — Lungfish: phylogeny and

parsimony, in Bemis W. H., Burggren W. W.

&: Kemp N. E. (eds), The Biology and

Evolution of Lungfishes, of Morphology

1: 151-162.

Schultze H. P. & Arsenauk M. 1985. — The pan-

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Schultze H. P. 1977a. — l'he origin of the rctra-

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Evolution. Plénum Press, New York and

London.

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GEODIVERSITAS • 1999 - 21 (2)

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Refcrences

Denison R. H. 1978. — Placodermi, in Schultze

H. P. (ed.), Handbook of Paleoichthyology,

Volume 2. Gustav Fischer, Stuttgart, 128 p.

Marshall C. R. 1987. — Lungfish: phylogeny and

parsimony, in Bemis W. F., Burggren W. W.

& Kemp N. F. (eds), The Biology and

Evolution of Lungfishes,/o//r;W of Morphology

1: 151-162.

Schultze H. P. ^ Arsenault M. 1985. — The pan-

derichihyid fish Elpistostege: a close relative to

tetrapods? Paleontology 28: 293-309.

Schultze H. P. 1977a. — The origin of the retra-

pod limb within the rhipidistian fishes:

541-544, in Hecht M. K., Coody P. C. &

Hecht B. C. (eds). Major Patterns in Vertebrate

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geodiversitas

1999 • 21 ( 2 )

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137

147

157

167

215

229

255

Indexed in

CeoRef (Online Database for Bibliography and Index of

Ceology), Blological Abstracts, ASFA (Aquatic Sciences and

Fisheries Abstract), Pascal, Zoological Record

Conception Graphique : Isabel Gautray

Morales J., Soria D. & Pickford M.

New stem giraffoid ruminants from the early and middie Miocene of Namibia

Forsten A.

Snout proportions in some Eurasian hipparions (Mammalia, Equidae): taxonomie and functional

implications

Cifelli R. L. & Madsen S. K.

Spalacotheriid symmetrodonts (Mammalia) from the médial Cretaceous (upper Albian or lower

Cenomanian) Mussentuchit local fauna, Cedar Mountain Formation, Utah, USA

Freneix $.

Bivalves du Trias tariquide de Los Pastores (Algésiras), Espagne. Leur signification

Poplin C.

Un paléoniscoïde (Pisces, Actinopterygii) de Buxières-les-Mines, témoin des affinités fauniques entre

Massif central et Bohême au passage Carbonifère-Permien

Pereda Suberbiola X. & Taquet P.

Restes de Rhabdodon (dinosaure omithopode) de Transylvanie donnés par Nopesa au Muséum

national d'Histoire naturelle de Paris

David H., Dauphin Y., Gautret P., Pickford M. & Senut B.

Composition en acides aminés d'os de mammifères fossiles de deux sites du Plio-Pléistocène

d'Angola. Comparaison avec la conservation de la phase minérale