A Small Collection of Fosils MIDDLE EOCENE

8/11/2019 A Small Collection of Fosils MIDDLE EOCENE

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CONTRIBUTIONS FROM THE MUSEUM OF PALEONTOLOGY

THE UNIVERSITY O MICHIG N

VOL. 24, NO. 18, p . 190-203 2

pls.,

text-figs.

M RCH 1,1977

A SMALL COLLECTION O F FOSSIL VERTEBRATE S

FROM THE MIDDLE EOCENE KULDAN A A ND KOHAT

FORMATIONS O F PUNJAB PAKISTAN)

PHILIP

D.

GINGERICH

MUSEUM OF PALEONTOLOGY

T H E U N IV E RSIT Y O F M ICH IG A N

ANN ARBOR


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CONTRIBUTIONS FROM T HE MUSEUM OF PALEONTOLOGY

Gerald R . Smith Director

Robert V. Kesling Editor

The series of contributions from the Museum of Paleontology is a medium for the

publication of papers based chiefly upon the collection in the Museum. When the number

of pages issued is sufficient t o make a volume a title page and a table of conte nts will be

sent t o libraries on the m ailing list and t o individuals upo n request. A list of the separate

papers may also be obta ine d. Correspondence should be directed to the Museum of

Paleo ntology The University of Michigan Ann Arbor Michigan

8109.

VOLS.

11-XXIV.

Parts of volumes may be o btained if available. Price lists available

upon inquiry.


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A SMALL COLLECTION OF FO SSIL VERTEBRATE S

FROM THE MIDDLE EOCENE KULDA NA AN D KO HAT

FORMATIONS OF PUNJAB PAKISTAN)

BY

Philip D . Gingerich

Abstract.-In 19 75 the au tho r made a new collection of fossil verte brates from the

Middle Eocene Kuldana and Kohat Form ations in the Kala Chitta mountains of the

Punjab Province in Pakistan. Ve rtebrate s were f oun d a t twelve localities in the vicinity

of Lamm idhan and Ch harat. All of the identifiable mamm als came fro m Lam midh an.

Two additional mammal specimens from Lammidhan in the British Museum (Natural

History) are also described and illustrated fo r the first time .

It is suggested tha t Gandakasia may b e a primitive cetacean ra ther than a meso-

nychid co ndy larth. The new artiodactyl genus Lamm idhania is proposed for the

anthra cothe re species Anthracobu ne wardi Pilgrim, 19 40 .

Discovery of a small concentration of mammal specimens at one locality suggests

that there is very good potential for the discovery of additional middle Eocene mam-

mals in the K uldana Form ation and its lateral equivalents.

INTRODUCTION

The Eocene vertebrate faunas of Europe and North America are now relatively well known, and

our knowledge of the Eocene faunas of central Asia is increasing.

It appears that a series of broad

holarctic migrations t oo k place at the beginning of th e Eoc ene, establishing a relatively hom ogen eous

fauna in No rth America, Eur op e, and central Asia just be fore these three con tinenta l areas became

isolated from each oth er. This unified faun a diversified rapid ly, with distinctive mammalian groups

evolving t o d omin ate each region. In the late Eocene and early Oligocene anoth er series of migra-

tions mixed elements of each of these faunas. The whole Tertiary history of mammals in the north-

ern hemisphere can be seen as a series of five or so cycles of pangeographic migration, endemic

deve lopm ent, followed by pangeographic mig ration, et c. Climate seems to have played an im po rtan t

role in the origin and dispersal of new mammalian groups, at the beginning of the Eocene at least

(see Gingerich,

1976 ,

and it is important, whenever possible, that mammalian faunas in equatorial

regions be sampled, since these regions appear plausibly to be centers of origination for a majority

of new higher taxa. Thus the faun as in the southern parts of each of the northern con tinents are of

special interest. Th e Eocene mamm alian fauna of the Indo-Pakistan subco ntinen t is one such ex-

ample.

The Eocene faun a of Indo-Pakistan is of particular interest for an oth er reason. It is generally

agreed that the Indo-Pakistan subcontinent moved northward during the late Cretaceous and early

Tertiary, joining the central Asian continent sometime during the Eocene. Better knowledge of the

early Tertiary land mammal faunas of Indo-Pakistan should help to date the time of first substantial

190


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VERTEBRATES FROM THE EOC ENE O F PUNJAB (PAKISTAN)

Anthracobune ?)wardi (Lammidhan, ?Jhalar)

Anthracotheridae ?) nde t. (Safed T ob ah , Jhalar)

HELOHYIDAE

?)

Khirtharia dayi

(Lamm idhan, ?Jhalar)

CREODONTA

MESONYCHIDAE

Genus indet., sp. n. (Safed Toba h)

Additional fossil vertebrates were collected from the general vicinity of Lammidhan and Jhalar

by a German expedition from Munich, all being described as coming from Ganda Kas. Unf or-

tu na te ly , no m ore precise locality or stratigraphic in form ation has ever been published for any of

these specimens. This collection was described by Dehm and Oettingen-Spielberg (1958), who

expanded the fauna from the area to include the following species:

TAENIODONTA

STYLINODONTINAE ?)

Basalina basalensis

gen. e t sp. nov.

Taeniodont indet.

CREODONTA

MESONYCHIDAE

Gandakasia potens

gen. et sp. nov.

Ichthyolestes pinfoldi

gen. et sp. nov.

Creodont indet.

(Ichthyolestes?)

Creodont indet.

CONDYLARTHRA

HYOPSODONTIDAE

Promioclaenus? gandaensis sp. nov.

PERISSODACTYLA

BRONTOTHERIIDAE

Eotitanops? da yi sp. nov.

HELALETIDAE

Teleolophus? daviesi

sp. nov.

Helaletid indet.

ARTIODACTYLA

DICHOBUNIDAE

Haqueina haquei gen. et sp. nov.

Khirtharia d ayi Pilgrim

Pilgrimella pilgrimi

gen. et sp. nov.

Pilgrimella wardi (Pilgrim)

Anthraco bune pin foldi Pilgrim

Anthracobune? daviesi

Pilgrim

Van Valen (1965) subsequently transferred the above species 'Fromioclaenus? gandaensis to

a new genus

Dulcidon,

which he placed in the insectivore or condylarth family Paroxyclaenidae.


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VERTEBRATES RO M T H E E O C E N E O F PU N J A B PA K ISTA N )

94

J

Kilometers

1 2 Miles

Streams

Footpaths

and above 15 00 ft

Paved Road

. Milepost

ossil Locality

TEXT-FIG. 2- Map showing middle Eocene foss i l loca l i t ies a t Lammidhan on the road be tween Basa l and Camp-

bellpore see text and tes t- f ig. 4 .


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P. D.

GINGERICH

dilian d enta ry, and coprolites. Also, this locality yielded to oth fragments, an edentulous mandible

fragm ent, a com plete proximal en d of a fem ur, and pieces of long bones of mammals. The collec-

tion from this locality is catalogued as UM 65858.

Locality

6: Kuldan a F orm atio n. This locality is in variegated red and violet beds in the valley west

of milepost 60.4 on the Basal-Campbellpore road, and it has yielded

Planorbis

a broken shark s

to ot h, broken bone of fish, turtle s, and crocodilians, coprolites, and a to oth fragme nt, vertebrae,

and bro ken phalanges of mammals (all catalogued as UM 658 59) .

Locality 7:

Red, yellow, and green shales of the Kuldana Formation near the village of Chharat,

typ e locality of th e Chhara t S eries. Beds in this area are nearly vertical, and platy limestones of the

Koh at Form ation are exposed just to the nort h of this locality. Th e locality yielded fragme nts of

crabs, turtle bone, crocodilian teeth and bone, and a fragment of a mammal tooth (all catalogued

as UM 65860).

Locality :

Kuldana Formation, about one-tenth of a mile west of Locality 7 and on the south side

of the tra ck . The stratigraphy in this area is comp licated by faultin g, and the relationship of this

locality to Locality 7 is no t clear. This locality yielded a large ?cranial bone fragment of a fish,

reptilian bone fragm ents, coprolites, and a weathere d mamm alian phalanx (UM 6 586 1).

Locality

9: Goo d exposures in the Kuldana F ormation six-tenths of a mile west of L ocality 8 ,

on b oth sides of the track along a small dry strea mb ed. Exposure s farther west of this loca lity

were prospected and some bo ne fragments were fo und . Locality 9 yielded only a crab claw, and

bone fragments of fish and reptiles (UM 65862).

Locality

10:

Kuldan a F or m atio n. This locality is in a bed of fine-to-coarse grained clastic carb ona te

that is abo ut three feet (1 m) thic k. The locality is on the sout h side of the hill straight west of

milepost 60 .4 on the Basal-Campbellpore road. broken shark s too th , fish and reptile bo ne , and

coprolites (UM 65863) were found here, and a mammalian mandible with a deciduous premolar in

the British Museum (BMNH M-15806) is preserved in a block of matrix of the same lithology.

Locality

:

Kuldana Form atio n. This locality is in

a

westward continuation of the same beds

exposed at L ocality 5 , in an area straight west of Locality 2 and milepost 59.7 on the Basal-Camp-

bellpore roa d. Locality 11 was the richest locality discovered, yielding

Planorbis

a shark s too th,

reptilian bone, and a vertebra and carpal bones of mammals (all catalogued as UM 65865). In addi-

tio n, associated fragments of a b ony fish are catalogued as UM 6 58 64 , and the following mamm al

specimens were collected in a small area of sharply eroding red badlands near the west end of the

exposure (see text-fig.

3).

All were fou nd within a square 15 feet (5 m) on a side, near the very

highest level of the red bed s:


Dehm and Oe ttingen, left P, (UM 65869 )

Haqueina haqtiei

Dehm and O ettingen, left M2 (UM 65 867 )

Khirtharia day i

Pilgrim, lef t mandible with P4 (UM 65870)

Lammidhania wardi

(Pilgrim), left astragalus (UM 6 58 72 )

A

small artiodactyl astragalus of

?Haqueina haquei

(UM 65 871 ) was found in the clastic carbonate

bed forming a low ridge in the eastern part of Locality

11

about 50

(1

5 m) below the m ain mamm al

locality stratigraphically.

Locality

12: Kuldana For ma tion. Abun dant bone was fo un d coming ou t of green marls in a small

exposure across the road to the west of Locality 2 , near milepost 59.8 on the Basal-Campbellpore

road.

Planorbis

and a high-spired gastropod, crocodilian and turtle bon e, coprolites, and mam malian

bone were foun d here (UM 658 66) .

Localities 2-6 and 10-1 2 are in the Lamm idhan area o n the road between Basal and Campbellpore


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TEXT-FIG. 3

Locality

11.

Upper figure shows a general view lookin 3 north east across the valley of Locality

1 1

towa rd the village of Gan da Kas in the distance. Area yieldins a conce ntratio n of mammal speci-

mens is in red beds in the lower lef t corner of th e photopraph. Lower f igure a lso looking north -

east shows a closer view o f the area yielding mammalian tee th and bone . Specimens were fou nd

on the s lope in the lef t center of the pho tograph.


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197 P

D.

GINGERICH

Koha t Form ation Kuldana Form ation

fianorbis

L e v e l

,LOCALITY 2

Level of LOCALITY 12

Bone Level

of

LOCALITIES 5 I

/ I

A 97

A60

A

NUMMULITE PLA TY Green

Red

SHALE LIMESTONE VAR IEGA TED BE DS

TEXT-FIG.

4

Stratigraphic relationships of localities at Lammidhan.

Diagrammatic section taken along the

BasalCampbellpore road between mileposts 59.7 and 60.1.

2

miles

1 4miles

9

-

i.3miles

To

CAMPBELLPORE

TEXT-FIG.

5

Map showing middle Eocene fossil localities along the track leading to Chharat, west of the Fateh-

jangCampbellpore road. All are in the Kuldana Formation see text).


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VERTEBRATES FROM THE EOCENE O F PUNJAB (PAKISTAN)

198

(see text-fig . 2). Th e stratigraphic positions of the localities, where kn ow n, are shown in text-figure

4 . Localities 7 -9 are in the Chharat area and can be reached by driving no rth from Fatehjang on the

road t o Campbellpore for

3

miles, then driving west on a dirt track past the prom inent white shrine

fo r 1.3 miles (see tex t-fig . 5).

NOTES ON T HE FOSS IL MAMMALS FROM LAMM IDHAN

The new collection fro m Lamm idhan described here includes representatives of fo ur genera:

Gandakasia Ichthyolestes Haqueina and Khirtharia. In addition, undescribed specimens of two

additional genera,

Anthracobune

and

Lammidhania

g. nov. were discovered in the British Museum

(Natu ral Histo-ry) collection fro m Lam mid han . Each of these is discussed briefly below .

Gandakasia potens

Dehm an d Oettingen-Spielberg, 19 58

Pl. 1 figs. l 2

Gandakasia

has very large, distinctive molars with a characteristic notch in the anterior margin.

Th e specimen foun d at Locality 2 (UM 658 68) preserves most of the crow n, bu t the talonid has been

broken off. Th e dentition of this genus is to o poorly known t o permit determination of which toot h

in the m olar series is represented. Because of breakage the length of the to ot h cannot be estimated.

Its maximum width is 8.6 mm .

Dehm and Oettingen-Spielberg (19 58 , p. 1 5) suggested tha t the maxilla of a mes ony chid described

by Pilgrim (19 40, see pl. 1 , fig. 1 0 here) from Safed Tobah m ight belong to

Gandakasia

bu t, having

studied the specimens in question, it is difficult to see how the lower teeth of

Gandakasia

would

occlude with te eth o f the confo rmation suggested by t he Safed Tobah specimen.

It is also wo rthy of note th at this specimen of

Gandakasia

is the only mamm al found in the Kohat

Form ation , which represents a distinctly more marine facies than the underlying Kuldana beds. The

possibility exists that Gandakasia is a primitive archaeocete rather than a mesonychid c ond ylarth .

Ichthyolestes pinfoldi Dehm and Oettingen-Spielberg, 19 58

P1. 1 , fig. 3

UM 6 586 9 is a partially erup ted left lower premo lar, probably P z . Th e length of the too th is

approximately 11 mm , and the width is 4.6 mm. Judging from its size and morphology, the to oth

probably represents

Ichthyolestes pinfoldi.

Haqueina haquei Dehm and Oettingen-Spielberg, 1958

P1. 1 , fig. 4

UM 65867 is a broken left upper molar, probably

M

The size of the tooth cannot be measured

accurately, bu t it is of the correct size and m orphology to occlude with the lower teeth of Haqueina

haquei.

An astragalus, UM 6 587 1 measuring approxim ately 8.2 mm in bread th across the proximal troch-

lea, is slightly narrower than the astragalus of Khirtharia dayi in the British Museum (BMNH 15 80 0,

measuring approximately 21 mm

in

leng th, 1 0 mm in breadth across the distal trochlea, and 10.5 mm

across the proximal trochlea). The specimen was found several meters below the to ot h of Haqueina

haquei

stratigraphically, bu t it probab ly belongs to this species.

Khirtharia dayi

Pilgrim, 1940

PI. 1 , figs. 5-7

A newly discovered left mandible fragment

UM

65870) is the first specimen of

Khirtharia

pre-

serving a premolar. Th e premolar, measuring 6.9

r r

in length and 4.6 mm

in

wi dt h, was still in th e

process of erupting.

An

x-radiograph of the mandible revealed no erupting crown under the roots


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199 P. D. GINGERICH

preserved behind this too th , confirming th at the t oo th in question is P4 and the preserved ro ots

were for M I . Comparison with molars of Khirtharia day i described by Pilgrim (1940), and Dehm

and Oettingen-Spielberg (19.58) shows that P4 was very small relative to molar size in this genus, as

in the related Bunodentus from the ?middle Eocene locality of Kalakot in Jammu-Kashmir (see

Ranga Rao, 197 2).

Anthracobune pinfoldi Pilgrim, 1940

P1.2, figs. 1-4

The type specimen of

Anthracobune pinfoldi

includes left and right mandible fragments each pre-

serving p arts of M Z- 3. While studying the type specimen in the British Museum (BMNH 157 92 ), it

was discovered t ha t an isolated molar trigonid (BMNH 157 94) fits on to the left M2 of the typ e

specimen (see pl. 2, figs. 1-2). n isolated MI (BMNH 32 169 ) that undoubtedly belongs to this

species was discovered in the collection from Lammidhan (see pl. 2, figs. 3 4 ) . This MI measures

25.5 mm in length and 16.3 mm in width . Furthermo re, comparison of the type specimen of

Anthra-

cob une f?) daviesi

Pilgrim, 19 40 (BMNH 15 79 5, a left maxilla w ith P ~ - ~r P2-3) indicates tha t it is

surely conspecific with

Anthracobune pinfoldi,

and

it

may possibly be part of the same individual

animal as the type specimen of

A. pinfoldi.

Lammidhania

new genus

Lammidhania wardi

(Pilgrim, 1940)

P1. 2, figs. 5-6

Pilgrim (1940) referred a third Lammidhan species, Anthracobune(?) wardi, questionably to

the genus

Anthracobune.

Dehm and Oettingen-Spielberg (1958) recognized that this species cer-

tainly does no t belong to Anthracobune, and placed it in their new genus Pilgrimella. An undescribed

M3 fro m Lamm idhan in t he British Museum almost certain ly belongs t o this species. The species was

previously know n only from the typ e specimen, th e broken talonid of a lower molar, and thus the

new M3 ad ds considerably to o ur knowledge of the species. The M3 (BMNH 32 168 ) measures

32.7 mm in length and 15.6 mm in width. The ty pe specimen (BMNH 1579 9), probably the talonid

of M I , measures 12.7 mm in width. Considering the specimens now know n, it is clear tha t this

species does not b elong in th e genus

Pilgrimella.

Dehnl and Oettingen-Spielberg (1958) identified the type specimen of

Pilgrimella pilgrimi,

an

isolated u pper mo lar, as an

M

From the arrangement o f the cusps on the specimen, it appears more

likely th at it is an M ~ .n either case, the new M3 of

A.

wardi

mentioned above indicates that the

species wardi had tee th significantly larger than tho se of Pilgrimella pilgrimi. Furthermo re, the lower

molars of wardi are to o selenodont t o occlude with upper molars of the morphology of Pilgrimella.

Th us Pilgrim s species

wardi

is here placed in th e new genus

Lammidhania.

It differs from

Pilgrimella

in being more selenodont. Lammidhania appears to be a primitive anthracotherid. Additional more

comp lete specimens are required to permit mo re detailed comparisons.

An isolated astragalus fou nd a t Locality 11 (UM 6 58 72 , pl. 1, figs. 8 , 9) probably belongs to

Lammidhania wardi,

or possibly to

Pilgrimella pilgrimi.

The astragalus measures 35.7 mm in total

leng th, 15.0 m m in width across the distal troch lea, and 14.6 mm in width across the proximal

trochlea.

PLATE 1.- (Figs. 1-7 x2 , 8-10 natur al size) FIGS.

1 2-

Gandakasia potens from Lammidhan,

UM

65868 ,

trigonid of lower molar; 1 stereo photo grap h of occlusal view; 2, lateral view. 3- Ichthyolestes pinfoldi f rom

Lammidhan,

UM

65869, left lower premolar, stereophotograph of lateral view.

4- Haqueina haquei

from Lammi-

dhan , UM 658 67, left upper molar, stereophotograph of occlusal view.

5, 6- Khirtharia dayi f rom Lammidhan,

UM

65870, left mandible fragment with P4 and roots of MI; 5 , stereophotograph of occlusal view; 6 , stereo-

pho tograp h of lateral view. 7-

Khirtharia dayi TYPE

f rom Lammidhan,

BMNH

157 96, left m andible fragment

with M2, lateral view. 8 , 9- Larnmidhania wardi o r Pilgrimella pilgrimi from Lamm idhan, UM 65 87 2, left astraga-

lus; 8 , stereopho tograp h of dorsal view; 9, stereo pho tograp h of ventral view.

10-

Mesonychid from Safed Tobah,

BMNH

158 05, edentulous m axilla, stereopho tograph of occlusal view.


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201 P.

D.

GINGERICH

CONCLUSIONS

The following fauna of middle Eocene mammals is now known from Lammidhan.

U

except

Gandakasia come from the Kuldana beds:

?CONDYLARTHRA

?MESONYCHIDAE


?CETACEA

?PROTOCETIDAE

Gandakasia potens

Kohat Formation)

ARTIODACTYLA

DICHOBUNIDAE

Khirtharia da yi

Haqueina haquei

ANTHRACOTHERIDAE

Anthraco bun e pin foldi

Lamm idhania wardi

?Pilgrimella pilgrimi

The most imp ortan t conclusion, based on discovery of the small concentration of fossil mammals

at Locality 11 , is that there is very good pote ntial for the discovery of additional middle Eocene

mamm als in the Kuldana Form ation and its lateral equivalents.

ACKNOWLEDGEMENTS

It is a pleasure to acknowledge the generous assistance and encouragement of Dr. Ibrahim Shah,

Director, and Mr. Mahmood Raza, Assistant Director, Paleontology and Stratigraphy Branch, Geo-

logical Survey of Pakistan. Similarly, I tha nk Professor David Pilbeam, Yale University, for his

assistance and encouragem ent. Fund s for field work were provided through th e Smithsonian Foreign

Currency Program.

Museum research was conducted during tenure of a NATO Postdoctoral Fellowship at the Uni-

versitk de Montpellier. I am grateful t o Drs. A. G entry , P. Andrews, and Mr.

J.

Hooker, British

Museum Natural History), Lon don , and Professor R . Deh m and Dr . N. Schm idt-Kittler, Munich,

for assistance in studyi ng collections in their care. Dr. D. E . Russell, Paris, provided casts of some

of the type specimens described by Dehm and Oettingen-Spielberg.

In Ann Arbor, Mr. T. Abdelnour made the x-radiograph of the mandible of

Khirtharia

R. G.

Habetler and

G.

Gunnel1 cleaned a nd catalogued th e new collection, and Mrs. Gladys Newton typed

the manuscript.

PLA TE 2.- All figures natural size) FIGS. 1 , 2 -

nthracobune pinfoldi

TYP from Lammidhan, BMNH 157 92

and 15 79 4, left mandible with M2-3; 1, stereophotog raph of occlusal view; 2, lateral view. 3 ,4 -

nthracobune

pinfoldi

from L amm idhan, BMNH 32169 , right mandible fragment with MI ; 3 stereophotograph of occlusal view;

4, lateral view.

5 6-

Lammidhania wardi

from Lammidhan, BMNH 32168, left mandible fragment with MJ;

5

stereo pho togra ph of occlusal view; 6 , lateral view.


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2 3

P

D. GINGERICH

LITERATURE CITED

COLBERT, E. H.

1938.

Fossil mammals from Burma

in

the American Museum of Natural History: Bull. Am er.

Mus. Nat. Hist., v. 74 p. 255-436.

DEHM, R., and OETTINGEN-SPIELBERG, T.

1958.

Palaontologische und geologische Untersuchungen

im

Tertiar

von Pakistan.

2 . Die mitteleocanen Saugetiere von Ganda Kas bei Basal

in

Nordwest-Pakistan: Ab h. Bayer.

Akad. Wiss., Math-Nat. Kl., v.

91

p.

1-54.

FATMI, A. N.

1974.

Lithostratigraphic units of the Kohat-Potwar province, Indu s Basin, Pakistan: Mem. Geol.

SUN. Pakistan, v. 10 p. 1-80.

GINGERICH, P. D.

1976. Cranial ana tom y and evolution of early Tertiary Plesiadapidae Mamm alia, Primates):

Univ. Mich. Pap. Paleon t., v. 15 p. 1-140.

LYDEKKER, R.

1887.

Th e fossil Verteb rata of India: Records Geol. Surv. India, v.

20

p.

1-79.

PILGRIM, G. E.

1940.

Middle Eocene mam mals from north-west Pakistan: Proc. Zool. Soc. Lon don , ser. B,

v. 110 p. 127-152.

PILGRIM, G. E., and C OTTER , G. d e P.

1915.

Som e newly discovered Eocene mammals from Burma. Rec. Geol.

Surv. India, v. 47 p. 42-77.

RADINSKY,

L.

1965. Early Tertiary Tapiroid ea of Asia: Bull. Am er. Mus. Nat. Hist., v. 129 p. 181-264.

RANGA RAO, A.

1972. New ma mm alian genera and species from K alakot zon e of Himalayan fo ot hills near

Ka lako t, Jammu-Kashmir St ate , India: Special Pap. Oil Nat. Gas Com m., Dera D un, v. 1, p. 1-22.

SAHNI, A., and KUMAR,

V.

1974. Palaeogene palaeobiogeography of the Indian subcontinent: Palaeogeography,

Palaeoclimatology, Palaeoecology, v. 15 p. 209-226.

SZALAY, F.

S.,

and GOULD,

S

J .

1966.

Asiatic Mesonychidae Mamm alia, Cond ylarthra): Bull. Amer. Mus. Nat.

Hist., v.

132

p.

127-174.

VAN VALEN, L.

1965.

Paroxyclae nidae, an exti nct family of Eurasian mamm als: Jour. Mammalogy, v.

46

p. 388-397.

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