Agriotherium intermedium n. sp. from the Pliocene bone breccia of Weze

-' C T A

\'01. II

I ' A L A E O N T O l . O G I C A , 9 5 7

JAN STACH

P O L O " ' ; ' C A No . I

AGR I OTHERIUM INTERMEDIUM

N. SP . FROM THE PLIOCENE BONE BRECCIA OF WEiZE

Stu dy on th e T er tiar y bone br ec cia F auna from WE: ze near Dzialoszyn in P oland

PART VII I*

.4. bst r act . - The writer describes fragme nts of den tition of Ag riot he rium Wagn.

(= Hya ena rc t os Fa lc. & Cau tl.) fr om the Pli ocen e bone breccia recovere d at Wp,ze near Dziat oszy n, Poland. He refers these remains to an in divi dual of a new spec ies from genus Agriot he riu m in te rme dium, which occupies an in termediate sy- st emati c position between A. insigne (Gerv.) fro m Montp ellier in Fr ance and A. si- valense (Fa lc. & Cau tl.) fr om Siwalik in India, in what con ce rns the shape of the

cr own of molars, that of M2 in partic ular.

DESCRIPTION OF MATER IAL

The bone breccia discovered at WE:ze n ear Dzialoszyn f r om whic h th e writer has already described ArctomeLes pliocaenicus, Ursus wenzen- sis and Nyctereutes s p. ', all belonging t o the group of carnivores, has also yielded the remains of Agriotherium

(=

Hyaenarcto s), another carnivore whose size greatly exceeds that of an y other animal forms as so ciated with the s ai d breccia.

The remains here cons idered are small , mostly detached fragments, m ainly referable to one individual.

1) The larg est of these remains

is,

a fragmen t of the right upper jaw , sh owing remnants of th e fourth premolar and t he first and second mo- la rs in a damaged condition.

The other r em a ins consis t of:

2) .lingual edge of cro wn of left upper se con d molar

(M2);

" Parts I-V - see Act a Geo!. Po!., vol. II-V/1952-55 ; parts VI-VII - Acta Pa-

!aeon t. Pol., vol. 1/1956.

1 Arctomeles pliocaenicus n. gen. & sp , from the Melinae subfam ily. Acta Geo!.

Po!., vol. II/195I. Ursus wen zensis, new speci e s of a small Pliocen e bear. Ibid., vol.

III/ I 953. Nyct er eu t es (Can idae ) in the Pliocen e of Poland. Ibid., vol. IV/1954.

2 ^{JAN STACH}

3)

br ok en off ex ternal cus p of righ t up per firs t molar

(M');

4)

posterior portion of the right upper fourt h pr emola r (P4);

5) fragment of left m a ndibl e with re mnants of lowe r fourth p remo- lar

(p.);

6) a detached, well preserved lower third in cisor

(13);

7) a likewdse be au tifu ll y preserved complet e left lower first mola r

(M1);

8) ri ght lower f irst molar (M

1) ,

sli ghtly damag ed.

T he fragment of right maxilla , men tioned a t the h ead of th e her e sp e cifi e d remains , is, by far, the most diagnostic specimen:

The upper fourth premolar of this fragment of maxilla is ba dly da- maged , th e crown and t he anter ior p art bein g altogether m is sing. T he ou tlin e of the base o f the cr own in t he po sterior end of t oot h, however, as w ell as the ou tlin e of it s r oot are di scernib le. No t much inference can be drawn from this to ot h as regar ds its sha pe an d si ze . The only con- cl usi on of some pr ob ab ili ty is tha t th e len gth of t he to oth had b een 29 mm.

In this it agrees wi th th e size asce rtaine d for that sa me tooth in Agrio- t h eri u m ins igne (Gerv.) b y Ch. Frick (1926-30) . The figure of the 29 mm length h as been ob tain ed by taking in to a ccount the 12 mm length of the broken off pos terior cusp (metacone) of t he tooth u nder item 4, an d b y com paring it wi t h the sa me ousp in A. in sign e (Gerv.) fr om Montpelher whic h is lik ewi se 12 mm long, as fig u re d by Frick (1926-1930 , f ig. 20) in a na tural size drawin g of t he comp lete crown of the upper fou rth premolar.

Th e upper firs t molar

(M')

in this fr agmen t is likewise in a very poo r state of preservation, as all t he four cusps are missing, having been partly broken off after the an im al had perished and p artly worn do w n, namely in the lingual side of the crown. On the preserved ba se of crown, how eve r , it is poss ib le to rec ognize its outline and m easure its size. The base shows to be of sub-quadrate shape with corners somew h at r ounded . Ex te rn all y the t ooth is 26 mm l ong, 25 mm m edi ally and 22.2 mm in ter- nally. T h e crown of the tooth is thus narr ower lingually an d with i nn er border m ore rounded. Th e cr own, as measu r ed at mi d-to oth len gth is 25 mm wi de. Th e size of the external po st erio r cu sp (m etacon e) in this toot h may be fi gured out 'by m easu r in g the br ok en off cusp of a n ot her M' , under item 3. T h is cu sp , as measured externally,is 15 mm high incl u di ng t he cingul u m, 10 mm with out the Clin&u lu m, an d 13 mm w ide. The summit of t he c us p is pointed, the edges consp icu ou s , t he s urfa ce wrinkl ed , this b ein g better recognizable on t he external si de of the cusp.

Th e up p er secon d m ol ar

(M2)

is in a fa r be tt er state

of

pr eservation,

th e paracone and t h e external border only b ein g m is sing. Perfectly pre-

s erved are its s econ d ex ternal cusp (metacone) an d the whole ldngual

AGRIOTHERIUM INT ERMEDIUM N. SP. FR OM THE PLI O CEN E 3

part of the crown , including the cingulum, and th e ridg e running length- wi se through the centre of the tooth. The crown of this tooth is L ike wise sub-quadrat e in shape, with rounded a ng les , Th e length here is 23.5 mm ex ter n al ly, 24.5 mm at mid-len gth a n d 25.2 mm on the lingual side ; the wi d th

is

26 mm an te r iorly, 25 mm m edially an d 24 ' m m po s te riorl y . These three dim ens ions are onl y ap prox im ate si n ce t he extern al border has not been preser ved an d on ly the trans it ion of the ro ot in to the crown is anteriorly we ll d isce rnible. Th e height of the m e ta con e as m easured ex tern a ll y, w ithou t t he cingulum, as this is missing ,

ds

6.8 mm, t he length of t h e ed ge descending lingu all y is 8 mm, t hat of the po sterior edge, mea- sured to th e inner border of the he r e pr es en t ci ngulum, is 9 m m. Th e cus p is i n the s hap e of an almost regul-ar pyrami d,

wdth

well sh owing anterior, lingu al an d po ster ior edges , an d fi ne ly wrinkle d surface . Th is cusp was separated f rom t he anterior cus p (pa r ac on e) b y a rat her p ro - minent depressio n . A gently cu r vin g ri b, i n the shape of a diagonal ex te n ding lingu ally to t he poste rior angle of t he squ a r e, an d well p r e- se r ved over its compl et e l ength, branches off from the po sterio r bord er of the missin g paracone. H

is

difficult to in dica te the exact position of t h e lingual cusps on this r ib. The grea te st h ei ght , 3.8 mm, is at ta in ed by t h e ri b medially , opp osit e to the depre ssion sepa r ating the parac one fr om t h e m eta con e.

It

is als o h er e that the r ib attains its maximum width of 6.5 mm, w hile a t it s in itial and termi nal part it is 3.8 and 3 rom res- pective ly . Auelong a ted, more or less tri ang ular area se para te s the r ib a lo ng its entire lengt h f r om th e sharply outh ne d bo rder of crown , for me d by the cin gul um. Th is fl a t area is covere d by den sely arrange d bu t s ha ll ow w ri nkles . Ther e is no li ng ual exten sion w hateve r of th e pos te r ior en d of the crown,

i.

e. no t alon . The circumference of th e to oth consis ts of a si ng le cin gulum, part icularl y high (6.8 mm) on th e lingual side of the crown .

A detached fr ag m ent of t h e lingual edge of th e left upper secon d ' molar (item 2)

is

probably r ef er ab le to th e s a m e indiv idual even though i ts length (26.5 mm) exceeds some w ha t that of the lingual borde r of the just described M2. In

.this

fragmen t als o is the dense transverse w rinkling readtly recognizable over the su rfa ce s eparatdng th e rib from the cin- gulum.

A fragment of t he Left mandibular branch has been recovered con-

si s tin g of a part ex ten din g from the posterior end of the canine to the

be ginnin g of t he first molar (M

1 ) .

The l ength of this fragment is 48 mm,

the height - 51 mm. Of the dentition a trace of the lower third premo-

lar

(P3)

has been preserved, with fragment of a single root emb edd ed in

a rather small alveolus, 5 mm in diameter, al so of the lower fourth pre-

molar (P

4)

directly adjacent.

It

is this remnant of the fourth premolar

4 ^{JAN STACH}

w h ich made i t possible t o asc erta in t he e x a ct length of the to ot h to be 18 mm an d i ts m aximum median br eadth as 10 mm. The area between t he 'canin e too th w hic h is miss in g he r e , ' an d t he a lveolus of th e t hi r d premolar does not s how any t r ace of teeth .

Very well pre served is the crown 0.£ a de t a ch ed left l ow er th ird inci- s or (1

3),

In shape i t mark edly resem bles th e crown of this in cis or in bears. A sm all e r acce ssory cusp b r an ch es off laterally to wards t he canine fro m t he m ain shaft of th e crown.

It is

externally sep arated from the cen tral sh aft b y a sh allow groove, wh ile ling ually three nibs des cend from the s um m it of the crown to it s base, na mely along the inner edg e of the shaft, mediall y, an d fro m t h e external edge ,. uniting a t the b as e of the crown . Th e heigh t of the cen tral s haft measu redexternally f r om the b ase of th e crown is 17.5 m m , that of t he sl ide branc h 13 mm; t h e su m m it of the shaft overtops by 4.5 mm the s u m mit of the s ide cu sp.

The wi d t h of the crown measured at the base is 11 m m, w h ile t he th ick- n es s of t he to oth at

d

ts bas e is 12 mm.

In a very sa tisf a ct or y state of p r eservation ar e . also two' de tached s pecimens of the rig ht an d left mandibul ar fi rs t m ol a rs . Th ey both, pro- bab ly , belonge d to t he same individual as th ey differ neithe r in shape no r in di mensions . Their s ize is r at h er large, the basal l en gth of th e cr ow n : bein g 37 mm, frontal wi dt h 17.5 mm, that

in

t he mi ddle al ong t h e Iine o f the central

^CUSIP

(prot oconid) 18.7 m m, while at the e n d, w ith in th e t alonid, i t is 20.5 mm. Th e pr in cip a l cu sp (p r otoconid), w hich is stro ngly d ev el ope d, attains 21 mm w he n meas ured fro m the lower border of the cingulum on t he outside . Its basal w idth is 16 mm , th e le ng th 14 mm.

A distin ct na r r ow r idge, 10 m m long, run s from the su m m it anter iorly , uniting directly w ith the posterior r id ge of the pre ced ing ant erior cusp . T he sh or t er (4 m m) posterior ri dge bifurca t es only j usta little be- l ow the s um m it t o for m two s harply ou tline d ri dges, 8 mm in length, wh ich enci r cle th e fla t triang u lar postenior wall of the 'cus p , s t eep ly d escen din g t o the dep r ess ion of the talo ni d. The lateral external ri dge is p oorly disc ernib le , whereas numer ous fi ne wrin k les are well shown at this s ide , arran ge d a lmost par allel t o ea ch other , from th e s um mit to the cin gul u m. The anterio r cu sp, the pa r aconid, measured at the ba se , a s 12 m m thick an d 11.5 mm lon g . Its heigh t m ea sured from t he

Uower

margin of the cin g ul um is 15 mm.

It

sh ows on ly t w o distinct ridg es, the anterior one descen di ng fr om th e su m mit to th e b as e where

<it

unit es w it h th e cin gulum, and the posterior one , 6 mm in length . Th e

ex ternal surface of that cu sp is a ls o fin el y w ri n kl e d. On th e ling ual si d e

the summit of the m et a con id r ises beyond the main cusp. Its h eigh t mea-

s u r ed inte rnally f ro m the l ow er ma rgi n of the cingu lum is 13 .5 mm,

w hil e meas ure d from the side of the talon id it is 6.5 mm ; t he meta-

AGRIOTHER IUM INTERMEDIU M N. SP. FROM TH E PLIO CE NE 5

conid is 7.75 mm lon g. Ant eriorl y the me taconid to uc h es the base of the large protoco nid, form ing with it an d with the posterio r cusp th e trigo- n id ,

28

mm long. Re ally it fo rms th e beginning of t h e intern al delimi ta- tion of t he talonid . Beyond it, on th e lingual side is the conspicu ousl y lo- wer e nt ocon id, with h ei gh t of 10 mm m easu red from the lo w er m argin of cingulum, followed b y the still lower entoconuli d , which pa r tially closes up po steriorly the taloni d by its ar cuate low summit ri dge . The hypoconid rises externally , opposit e th e two la st small lingual cus ps . Its , heigh t measur ed ex te rnally from the lower m a rgin of th e c ing u lum is

14.5

rnm, th a t m easur ed fro m the talon idal dep r ession bein g 6 rnm ; it s thickn ess is

12

mm , the posterior su mm it r idge is 5.8 mm long, w hil e the anterior, reaching to the base of the cardina l cusp is 8 mm lon g. No cusp or th ick ening is disce rnible on this r idge.

T h e talonid, being thus lingually delimited by three cusps - the m e- tac on id in clude d - an d labially by t he hypo con id an d its anterior ridg e, is re lative ly b road and rather long . T he ci ng u lum surrounding th e crow n at t he b ase is high (7 mm) on the outer s ide, b elo w the h yp ocon id, tha t is to s ay posteriorly. It is , h owever , relative ly weak ly marked, near the s u mm it

lit

ascends and becom es almost entirely obli ter a te d, to be again, though very indistinctly discernib le, near the anterior cusp. Lingually the c in gu lu m is almost completely obliterated.

GENERIC AND SPECIFIC DETERMINATION

Th e s hap e and size of the here described teeth lead w ith perfec t s u- rety to the inference t hat they formed part of the den ti t ion of an indi- vid ual belonging to a group of species or igin all y known under t h e common generic name of

Hsjaenar ctos

(Falconer

&

C au tl ey).

Remains of an individual from th is genus were first recovered from Pl iocene deposits i n the Siwalik Hills of India. Th ey consisted of a s t r ong ly damag ed right mandible wit h P

4 ,

M

1

and M

2 ,

also with alveoli referable t o two premolars and M

_a,

which are all missin g. The nex t find w as th at of part of a sku ll w ith preser ved canin es, P4_M2 in a damaged condition , and alveoli of P 2_P 3. Th ese r em ains were in

1836

descr ib ed by H. F alcon er an d Cauiley an d ref err e d by t he m t o the s kele ton of a b e ar which they called Ursu s siv alem is F alc.

&

Cau tl .

T he autho rs themselves, an d la ter in

1837

also A. Wagn er , la id e m-

phasis on the dentition of the d es crib ed specimen not agree in g with that

typical of th e genus Ursu s, so muc h s o that th e ge ne ric n ame of Agrio-

therium was suggested by A. Wagner fo r the in dividu al from the S iwali k

H ill s. So m e ti me af ter th e name of Huaenarct os Falc .

&

Cau tl . was in-

trodu ced for that genus by

R.

Owen who ass erte d t hat the auth ors h ad

6 ^{JAN STACH}

o r ig inally intended to call their specimen by this name gui de d b y the rese m blance in shape of P ! to that tooth in the hyena. T his name , a cce pted by H . Falconer (1848) as su bgeneric for this type of foss il m ammal , h as b ee n since used by palaeontologdsts as generic.

G en us Hu aemarctos (Falc, & Cautl.) has been subse quently s plit up in t o g enus Agriotherium Wagn. with Agriotherium sivalense (F alc. &

Cautl.) a s genotype, genus I n darctos P ilg r im with I n dar cto s salmantanus P ilg r im as ge n oty p e, an d s ubge n us Hyaenarctos (Lydekkerion ) F r ick which has not b een generally accepted, with H y aen a1 " ctos paiaeisuiicu s L ydekker as genotype.

T he main differences between Agriotherium W agn. and In darctos Pilgr. lie in th e s h a p e of dentition , of the upper jaw part icularly. T hey have been summed

UiP

by Ch . Frick , W . D. Ma tthew, G. E. Pilgrim and others . Th e chi ef on es are as follows :

1. In Indarctos Pilgr. the anterio r extern al Hceesso ry cus p, the pa- rastyl e, on P\ is poorly developed while it is promin en t in Agriotherium W agn .; in Agriotherium this tooth is large an d longer than the next molar, MI.

2. In In dar ct os Pilg r. t he basal se ct ion of M! is sub-quadrate, while i n Agriotherium W agn. the lingual border of the crown is s horter than t he labial an d with angles d istinctly r ou nded .

3. In Indarctos Pilgr. th e labia l poster ior bo rd er of M2 extends int o a m ore or less promine n t ta lon, whil e in Agriotherium W agn . t here is n o tal on on

this

toot h.

4. In Indarctos P ilgr. M , is relati vely long, with the talon id s ome- w hat shorter t ha n t he trigonid, an d the h y p ocon id lower than the en- to conid , w hi le in Agrioth eriu m W agn . M, is r el a t iv el y sh or t, with the t alonid m ark edly s h orter than the trigo ni d, a n d the hypoconid higher t h an th e en t oconi d.

On d iffer ences of dentitio n , as h ere spe cifie d, betw een Agriotheri'um Wagn. an d Indar cto s Pilgr. one is w it h perfect sure ty led to the inference t h a t the described remains b elong ed to the skelet on of an inddvidual from gen u s Agriotherium Wa gn. N amel y:

1) MI is wi th the li ng ual bo rd er d istinctly sho rte r t ha n the labial an d with angles r ou n d ed ;

2) th e lingual posterior borde r of M2 d isp lay s no post erior con v exity , which me an s t hat t he crown is witho ut a talo n ;

3) the anterior part of M

l,

the trigo ni d , is cons iderabl y l on ger than the post erior, the talonid; t he crown of t his t ooth

is

l arges t w ithi n the ta lo ni d, a n d t he h ypoconid

is

h igher t han the ento con id .

It

is fa r more diffi cult specifica lly t o i de n t ify t he re mains of Agrio-

t heriu m, fou n d in the Wr::.ze br eccia.

AGRIOTHERIUlvI INTERlvIEDIUlI1 N. SP. FR O M TH E PLIOCENE 7

The paucity of remains thus far recovered from diverse sites, in mo st cases consisting of the detached upper or lower jaw only, has sometimes p r ev en ted t h ei r undoubted id en tification wi th this or an oth er s p eci es of Agriortherium Wagn. or of Indarctos Pilgr.

For this r eason such rem ains were mostly referred to some species wit h in the group of Hy,aenarctos w it h out generic differentiation into Agriotherium and Indarctos. In t his way , a score or so of sp ecies have been es tablishe d - part of which are probably id entic al - or th ey were s imply called Hyaenard, tos sp.

The r ecove ry, t h erefor e, from the sa m e depo sits , with in so small an a r ea as that occupied by the Wp.ze brec cia , of f ragments of both the uppe r an d lower dentition of an in di vi dual, undoub tedly re fer able to one species f r om gen us Agriotherium, prov id es impor tant evide n ce for the know- l edge of t he dentition of t h is species.

Since d ental dimensions s er ve as s up plementa ry evide n ce in spe cific di ag n os is and sinc e th e gr oup of forms known under the common name of H yaenarct os has a ve ry wi de r ange of dis tribution within the Ho- l a r cti c ar ea, it see ms mos t ad visable to t abu late da ta concerning th e dim ensi ons of t he here studied t eeth . in s pe cies t hus far ide ntified f rom . the ge neral group of Hy aen ar ct os (see t able on p. 8-9).

A revie w of the sp eci es li st ed in t he he re annexed t ab le shows th e predom in an ce, within the en tir e H ol ar ctic a r ea, of species referred by their au th ors to genus In darctos P ilg r , Th e predominance wo uld , perhaps, be still gr eat er, should the upper se con d molar be always present in t h e r em ai ns of forms belon ging to the H y aen arct os grou p, rec orde d from diffe rent sites. On the pres ence or absen ce of a t alon in this to ot h , the r emains could without doub t be referred to some s pe cies with in gen us In darctos or Agrio theriu m r espe ctive ly . Sp ecie s tentati vely determine d as Hya ena rct os

sp ,

wou ld then also disappear from the list.

The assignmen t of a fo rm to genus Agriot he rium or to genus I ndarct os on the f irs t molar only , does not always seem quit e j ustifie d to t he writer . A comparison for exa mple of this too th ,collec ted f r om the Wp.ze breccia, w ith th e same to oth recovere d f ro m the . isl an d of Sames , t he sh ape of w h ich is exce llently s how n on ph otog r aphs in a p ap e r by H . Helbin g (1932, fig. 3 a-b) , here reproduce d in pl . II , fig . 5-6, shows th eir al m ost iden tical s imilarit y . In both , t he h yp ocon id is h igh er than t he entoconid and a distin ct e ntoconulid is p r esen t in additio n to t he ent ocon id. An d y et , on the a bsen ce of talon in t he upper s econd m ola r in the Wp.ze sp ecim en , i t is ge n us A gri ot herium, while the prese n ce of a pro m inen t t alon in the same t oot h b elonging to t he Sam os s pecim en refers it t o genus Indarc tos.

The fo llo win g are, aft er Ch. Frick (1926-3 0), sp eci es undoubt edly

r ef er ab le to Agriotherium Wa gn .:

8 ^{JAN STAC H}

Table sh owing dimens ion s of P'', M', M' and M, in spe cies with in the grou p of Hyaenarc tos (in rnmj

Spe ci es ^pI M' M, Patria !Ge n us

',

I W~ze,

Pola nd Montpellier'l

Fran ce I

Boutonnet,

i

Fr a n ce

I

^{ViFrancealette,}

IAl COi, Spain Agr.?

41.5 40X22.5 38X21.5 37

X

20.5

24 25.2X 28.5 27.4

v

26.5 24.5

X

25 25 / 25

27.8Y 27.5

30\, 23 insignis

Gerv a is, 1853

sp. Gerv ais, 1853 insignis

Gerv ais, 1853

1

29.1X21

I

inter m edi1!111 n. sp.

sp . He lb ing, 1932

sp. Flow er, 1877

pon t i cum

I

Kormos, 1913 laurillardi Me-\

neghini, 1863

i ar ctoid es

Deperet & I Lleuca, 1895 24.6

30

X

30

24.6X22

I

^28.2X22

41:x19.5

33

25

Felixtow, Agr .?

England

Baltavar, Agr.?

Hung ary

IMon te Ba m- Agr.?

1 boli, Italy

Or ignac, In d. France

29

X

21 ! 27

X

23.5 30X 24.5 1

27)!27.5

I

^{39.5 . 21 Siwalik,}

I

Ind ia Ag r . I

I

^Montredon,,I

France ,

I

^{Pikermi, I'}

Greece

Ind. Agr.

Siwali k , In d ia Siwalik.

India Valles-Pe- lnedes,Spa in

I I

^{Sarn o s,}

Gre ec e I

:G au -Wein h.

Ge r m any 40

38

42.5 ! 21.5

1

I

41. 6

X22.2

I

31X27 28X 23 29X30.5

26 34X26

25.1X 18.3

27X25.5 II 30 . 28 29X 29.9 29X27

32~<22.5

32

><

22

27)< 21

29.1X 24,3

I

silJalen si s

I

Fal coner &

Cautley,183 61 punjabiensi s

Lyd e k ker, 1878 sp. Tob ien,

1952

palaei ndicus Lydek ker, 1878 urct oides

Deperet & , Lleuca. 1895 \ utt i cus Dames, 1

1883

sp . Helbing , 1932 IJire ti Villatt a

& Crusafo nt,

1943

AGRIOT llER lUlYl INTERlYlEDIUlYl N. se. FR OM THE PL I OCENE 9

Ta ble show ing dim en sio n s of pI, MI, M~ and MI in species within the grou p of Hyaenarctos (in mm)

(contin ued)

Species ^{pI MI M~} M,

I

Patr ia Genu s

oregonen si s

Merriam,1916 31.7X22.6 , -

schneideri

I I

Sellards, 1916 . -

I

29.8X30.2

I I

sp , Matthew, 1918 lagrelii

Zdansk Y,1924 27.3X21.4

sp, Freud en- berg, 1910

-~_!~

Agr.?

Agr.?

Ind.

Florida, USA Ore gon,

USA

Ma r agh a, Ind. Pe rsia

Ed en, California Salt Range ,

I

India

I

Pao-Tedis tr.l China

I

Pao-TediSh"1 Chi n a

i

Sou t h Ch ina Ag r .?

SnakeCreek Agr.?

USA

I

Tehuichil a,

I

^Agr·?

MeXIC O

44X23

42.8X21.4 37.5X20.8

,

: 46.5X24

i

26X24 35X 27

41X23.5 1 I43.5X23.5

I

35.:1X 27.2 ' 32.6X 23.4 29X25.5

27.3X25

i

\

, I

- I

I

32.3X21.7

35.4

X

25.8 30.5

X

32.3 , 36.5

gregor i Frick, 1921 sinensis

Zda n s ky ,19241 sp , Lyde k ker,

1844 sa lm on t anus

Pilgrim, 1913 mara ghanu s

Mequenen, 1925

1) Agriatherium insigne (Gervais, 1853)

(=

HyaenaTdt:os insignis Gervais) from Montpellier in F r ance;

2) A . sivaLense (Falconer & Cautl ey, 1836)

(= U7'S US

siv alensis F alc.

&

Cautl.) fr om S iw al ik in India ;

3) A. palaein di cum (L yd ekker, 1878)

(=

Hu aenarctos palae indicum L yd ek.) from Siw a lik in India ;

4) Agrio therium sp. (Helbing, 1932)

(=

Hy aenarctos s p. Hel b.) f ro m Vialet t e in F r anc e.

The fo llowing are a lso rega r ded b y Ch. F r ick

(l.

c.) as b el on gin g t o Agriot heri u m :

Hya enarcto s sp. Ger v ais , 1853, from Alcoy in Spa in (fragment of upper jaw wi t h P ' and a r emnan t of the ante ri or part of MI);

H. " schneid eri" S ellards, from B r ews t er , Fl ori d a , USA

(M'

only) ;

JAN STACH

Hya enarctos sp. Flower, 1877, from Felixstow near W al d r ing field , e as t of Englan d (MI from Red Cr ag only);

H. gregori Frick, 1921 , from Eden in California, U SA (damaged P ' and M I, also P 4-M 2 )'

Thus tru ly but one single species

fir om

Agriotherium Wagn. h as been recor de d from Europe, namely that of Agriotherium insigne (Gerv.). This spe ci es has been established on fra gment of the upper jaw, wi th partly preserved dentition, belonging to one individual from Mo ntpellier, P . Gervais, when de scribing the den tition of this individual, also publish- ed in 1853 and 1859 a natural s ize drawing of th e fourth premolar and of both molars. Th es e are t eeth with s ha pe which offers particular interest ow ing to th eir comparability with th e t ee th of Agriotherium recorded f ro m Wf{z.e. The lower dentition of A . insigne (Gerv.) was , on the other hand , described in 1939 b y J .

Vbret

on ev ide n ce of three mandibular frag- m ents colle cted from Boutonnet, M ontp ellier. Excellent photographs show n in table II , fig.

²

a-b of J . Vir e t 's paper , ena ble a close comparison to b e m ade of the s ha pe of the fi r st mo l ar in A. insigne (Gerv.) with that of t h e. sa m e tooth in the W

~z e

spe ci men .

F or a bett er comparison of these t e eth the following drawings are reproduc ed in pl . II of the present pap e r: upp er dentition (P4_M2) in species A. insigne (Ge rv.), A . siva ben se (F alc.

&

Cautl.) and A. palaeindi- eus (Lyd.) , M2 in Agrioth erium sp . (H elb.) , an d lo wer den t ition (MI) in I ndarcto s sp. H eLb. and M

1

in A. insign e (G e rv.) described by J. Viret.

When comparing th e size and shape of the maxilla r molars in the W G ze. speci men wit h dimens ions in ot he r spe cies of Agriotherium sp eci- fie d in th e an nex ed ta b le , also with the d r awing of tee th by Ch . F'nick i ncl u ded in his desc r ip tion of A . insigne (G erv .), it is observable that the teeth in the

W~ze

specim en are some w hat smaller and the crown in the be t t er preserv e d mola r s omewh a t differen tl y shaped, to say:

1.

The length of the anter ior e x t e r n al C USp2

in

the

W~ze

specimen pr actic all y eq uals th a t of th e next cusp, the metacone

(12.2 :12

mm ), the b oun da ry lin e where t he ir bas es m ee t is at mid-length of the t oot h, w hile in A. ins ign e (G e r v .) th e anterior cu sp is di stinctly long er than the pos- teri or (15 : 11 mm) and t he bases of the cusps meet beyond the mid-length po int of th e too th .

2.

Th e ante ri or bo rd er of crown in M2 of th e

W~ze

sp eci m e n is a lm ost parallel t o the post erior bo rder , wh il e in A . insigne (Gerv.) the borders are inclined to each oth e r at an acu t e a ngl e.

2 Though in the Wr:ze specimen this cusp is broken off, we can see outlined the re the cours e of the cin gulum from the edge of which the measurements wer e made. The ling ua l edge of th is cusp isdiscer nible on the crown of the tooth, as well as the linewher e the base of this cusp meets with tha t ofthenextcus p, the metacone.

AGRIOTHERIUM INTERMEDIUM N. SP. FROM THE PLIOCENE 11

3. The posterior border of the tooth in the

W~ze

specimen is almost peq: En riicul ar ly dis p ose d to the lingual border, with an inconspicuous depression at its mid-length, while in A . insigne (Gerv.) the posterior border is ' b r oadly r oun ded lingually an d displays a distinct cavity up- wa r ds

of

its mid-length.

In m an y of th ese characters M2 of t he

W~ze

specimen approaches the s am e tooth in the s peci me n from V iale tte, by H. Helbing (1932) defined as Hyaenarctos s p. In the Vialette s pe cime n, however, the rib extending obliq u ely over the en tire length of the crown is quite distinctly divided in to s eparate lingual cusps.

Much closer than to

A.

insigne (G erv.) does M2 in th e

W~ze

specimen come , as shap e is concerned, to A. siv alense (Falc. & Cautl.) , a species w hi ch J. Viret is in cli n ed to identif y with A . insirrne (G erv.). In A. siv a- lense (Falc.

&

Cau tl.) , however, t h er e is some slight t en den cy t o develop a t al on , ow in g to w hich th e lingu al bord er of this too th is som ew h at lon- ger t h a n the la bial (27 X 25 mm) , according t o dimensions shown by the dr-awi n g in Ch . Fr ick's p ap er.

The lower fi rst molar is less diagnostic in the matter of comparison an d sp ecifi c id enti fication , In the

W~ze

spe cime n M

I is

37 mm long, while its m aximum width on the taloni d is 20.5 mm ;

1n

A. in signe (G erv.) the se di me ns ions are not much g reater, bein g after J. Vire t (1939) 38-40 mm and 21.2- 22.5 mm respec tive ly. Th e t oot h is, ho wever, shaped s om ew h at di ffe re n tl y, to say :

1. I n the W

~ze

specimen , o n the li ngu al si de of M

I ,

tw o distinct cusps ri se in additio n t o th e m e ta con id , that is th e entoconid an d the e n tocon ulid, as an exte nsi on of this border of t h e crow n . In A. insigne (Ger v .) the ento con ulid is mi ss in g in this tooth, th e entoconid onl y cons- t it u ti n g b esid es t he met aconid the lingual wall of th e cr ow n .

2. The cingulum of t he crown on the labial side of the toot h is less promi ne nt in t he

W~ze

s pe cime n and it , is not discontinuous as in the t ooth of A. insigne (G erv .).

Th e complete oblit er ation of t h e cr own in M

I

belonging to t h e spe- cimen of A. sivalense (F alc.

&

C autl.) im pe des the comparison of its sh ap e with t h at of t he crow n ° i!I1 M

1

of the

W~ze

spe cim en.

On t he ground of diff er en ces of sha pe , observed in teeth just mention- ed, belongin g to th e

W~ze

s pecim en a n d those in

A.

insigne (Gerv.) and

A.

si'l:alense (Fal c. & Cau tl .), an d also in considerati on of t he great geo - gr aph ical dis t an ce separating

W~ze

from the Siwalik H ills in India which have yie l de d t he remains of

A.

siv alens e (Falc. & C au tl.), t he present w r it er b eli ev es it advisab le to assign an independent n ame to th e

W~ze

spe cim en, namel y that 'of A. i.ntermed-ium n. sp . Some lucky discovery

o f m or e ample m a ter ial , at t hese or ot he r s ites, w ill pe rh a ps in the future

12 ^JA N ST AC H

l ea d t o the kn owl edg e of va r ia tion s,

if

any, . an d of t he ex te n t t o wh ich.

they m ay hav e a ffe cte d t h e sh ape of den titi on in in di vid u a ls f r om Agrio- t h erium Wagn .

GENERAL REMARKS

Fo ss il remai ns belon ging t o anima ls fro m t he g r oup he re considered an d origina lly refe rred to und er the common n ame of Hya enarcto s (Falc.

& Ca u tl.) h ave been recor de d fr om Sp a in , Fra nc e, Gr ea t B ri ta in (F elix- stow near Wa ld field r in g), Ge rma ny , Po land , Hu n gary , Italy, Gree ce (P i- ke rmi, island of Samos), P ersi a, Ch in a, India , also Californ ia, Neb r asc a, Or egon , Florid a in t he USA, an d Mexico . The a rea in habite d by this gro up of carnivo res is thus seen t o have been a vast on e, s ince it extende d ove r ne arly th e whole of south ern and cen tral Europe an d stret che d on east- ward to As ia , occupying als o w ide terri to ri es w it hin No rth Amer ic a.

This sugges ts g reat similari t y if not ide n tity of li vi ng conditi ons of th is group of animals, prevailing during a cer t ain p eriod of ti me ov er i m m ense territories under th e same ge ogr ap hica l la ti tude. These e n- vironmental conditions were probably responsible for the developmen

t

of geo gr aph ic and eco logic fo rms showing no important differences.

Should only those forms from the ' com m on group be taken in to account as are now referred to under the g eneric name of Aqriotheruun. Wagn., still, the occurrence of A. insigne (Gerv.) in France, of A. intermedium n. sp. in Poland and of A . sivaLense (Falc. & Cautl.)

in

the North of India (Siwalik) points out to the existence at one time of common links uniting very closely related forms, distributed along a track extending fro m France to India.

Some difficulties are encountered in determining the time of occur- rence of the Hyaenaret cs group. Geologists do no t always agree as t o th e age of deposits from which fossil remain s have been recovered r eferabl e to r epres enta tives of that group , the age ass igned t o them r anging f rom the Upper Miocene t o the Upp er Pliocen e. D epos its which have y ielded remains of Agriot herium are regarde d as o f P li ocen e age a n d it is the Midd le or th e Upper Pl iocene periods whic h appear to h ave witnesse d the greatest specific abunda nce of this genus . Fossil r emains of In dar c- tos, on the other han d, are more copious in the Lower t h an in t h e Middle Pliocene and In dar ct os is by G. E. Pilgrim, H . To bien and others r egard- ed as th e time-marker for t he Hipparion faun a in t he south of Eur ope and Asia.

T h e carnivores from t he

W~z e

bone br eccia, which have thus f ar

been given cl oser con sideratio n, resemb le foss il faunas r ecorded f r om

the Mcn tpellier deposits in France. Since the age of these deposits i s

n ow b eing s hifted from the Lower to the Mid dle P li ocen e it is h en ce to-

AGRIOTHE RJUM IN T ERME DIU M N. SP. FROM TH E PLIO C ENE 13

b e inferred that the remai ns of

A.

inter mediu m are like wi se assig nab le t h e Mi ddle P li oce n e age .

Thou gh th e mo rp ho logy of species of Agriotherium Wagn. com es qui t e close to that of Indarctos Pi lgr . being probably r efer ab le I to a common an ces tor and to a contemporaneous date of origin a n d extinctio n, yet th ey constitut e two independen t stocks. T h ei r an cestor is to be searched for in t he Miocene among the multitude of sma ll forms belo nging to genu s Ursacus S ch loss. Differences of size excepted, th e mo lars of Ur sa- v us come so nea r in sh ape to t hose of In dar ct os that so me species of

Ursavus

were or ig in all y r eferred to H yaenar ctos (Fa lc.

&

Cau tl. ), as for ex am ple Ursacus bre'&irhvnus Hofman n , 1887 , by M. Schlosser in 1887 iden tif ied as

Hu aen oirct o«

m in utu s Sc hloss., .an d Ursav uJs d epe r eti Schlos- se r , 1902 in 1895 by Ch. D eper e t referre d to H ua enar ct os arc toiien s D ep.

Individuals fro m Agrioth eriu m hav e sometimes abtaine d con side rable dim ens ion s. Th e skull of A . sivalense (F alc.

&

Cau tl.) is by

R.

L ydekke r r epor te d to h ave been 482 .6 m m in le ngth , whic h m ak es the size of the w h ole bo dy of th a t for m e qu al the dimensions of the la r ge st indivi duals of the cave bea r . The skull of

A.

in t erm ediu m n. sop. could not h av e be en m u ch sm all er . Since, in A . siv a7ense, with le ngth of M' and M2 at 57 mm, t h e length of skull is 482.6 mm, then, with t he same ratio ap pl ie d to

A.

in- term ediu m , in which t he le ngth of a number of teeth attain 51 m m, t he l en gth of skull wo u ld h av e to the 431.4 mm .

Th e sl t r ong ly dev elope d carn ass ials in all species bel ongin g t o the Hyaer. arcto s group in dic at e that . r e p r es en ta tiv es of t h is gr ou p were all carniv or es wit h po werful bodies . Th e l arge si ze of the uppe r fo urth premolar, shaped v ery much like that of the Hyae na, must prob abl y have ma de easy th e crush ing of the bigg es t bo n es. T h e m od e of life of A . int er- medium ma y , . t h er ef'or e, hav e r es embled that of hyenas. Its h abi ta t w as per ha ps one of ste ppe s wi th a few trees an d bushes clustered h ere and t here partic ularly in the vicini ty of wa te r reserv oirs.

Zoologic a l Institute ot the Po li sh Academy of Scienc es

Cr acow Bran ch Krakow, Sept emb er 19,56

REFERENCES

DE P E RET CH. 1895. Re sultats des Iouilles pal eon bolo gique s dans le miocene su pe- rieu r de la colline deMontredon. C. R. heb d. Seances Acad.Sci. ,121, 433-434, Pa r is.

DEPBRET CH. & LLUECA G. 1928. Sur l'In darctos arctoides et la phy logenie des Ursides,C.R. Soc.Ceo!. France,4, 149-160.Par is.

ERD BR IN K D.P. 1953. A review of fossil an d recent Bears of the Old Wor ld.Inaug.

Dissert. Mi n. Ceo!. Inst. Univ . Ut r ech t ,1- 2, 1- 597. Utrecht.

14 JAN ST AC H

FALOONER H. & CAUT LE Y . 1836.Note on the Ursus sivalensis, a ne w fossHspecies from Siw ali k Hill s. Asi ati c Research, 19, 193-200 .

FLOWER W. H. 1877.Note on the occurrence of the rema irns of Hyaenarctos in the Red Cr ag of Suffolk. Qu art.J. Geo!.Soc., 33, 534-536. London.

FRICK CH. 1921. Ex ti n c t vertebrate fau nas of Badla n ds , of Bau tist a Cree k and Sa n Tim oteo Canon, Sou therm Califo rnia . Univ. Cali! . Publ. Geo!., 12, 277-424.

Ber keley.

1926/1930. The He rnic ybn inae and an American Tert iary Bear. Bull. Amer. Mus. Nat: Hist., 56, 1-119.New York.

GERVA IS P. 1853. Descrip tion des ossements fossiles de mammif e r es rappor tes d'Espagnepar MM. deVe r ne u il,CoUomb et deLoriere ,Bull. Soc. Geol,France, 10, 147-1 67.Par is.

HELB I NG H.1932.Uber einern Ind a r ct os-Sch adel aus dem Pon tien der Ins el Sames. nebst einern Anhang : Hyaen ar ctos sp . aus dem Pliocen von Vialette (Ha ute- Loire). Abh. Sch w ei z. Pa l ii ont . Ges., 52, 1-18. Basel.

HOFMANN A. 1887/88.Uber einige Sau g eth ierreste aus derBraunkohle von Voit sberg und Steie r eg g bet Wie s., Steierma r k . Jb. k, k. geol. Reich sanst. , 37, 207-2 18.

Wien.

LYDEKKER R. 1884. Indian Tertiary and Post-Ter tia r y Vertebra t a. P. VI: Sivalik an d Na rb ad a Ca rniv or a .Palaeont .Indi ca,ser .10, 2,202-23 9.Cal cutta ^& London. MATTHEW W. D. 1929. Third con tribu tion to the Snake Cre ek faun a. Bull. Am er.

Mus. Nat. Hi st. ,56, 437-5 60.Ne w Yor k.

OWEN R. 1840/45. Odontogr aphy .Vol. 1- 2. Lo ndon.

PI L G RI M G.E. 1931. Ca t alog ue of the Pontian Carn iv or a of Europ e.Br it. Mus. (Nat.

His t.), 1-174. Lond on.

SCHLOSSER M. 1891. Die Affen, Le mur en , Chiroptere n , Insectiv oren, Ma r sup ialier, Cr eod ont en un d Ca r n iv or e n des europa.sche n Tert.ars und der en Beziehungen zu ih r en leb end en und fossilen ausse r-europaisc he n Verw andten. T. 3. Beitr. Paliion t. Ost err.-Ung. u. Orients, 8, 1- 106. Wien.

TOBIE N H. 1952. In dar ctos und Ursavus (Carnivora, Marnm .) aus den Unte r p li o- zan en Din oth erien dand en Rheinhes sens. Nbl, Hess. Landesam t. Botlen j. , 6, 7'-14.Wiesbaden .

VIRET J. 1939.Mon ogr a phie paleontolo gique dela faune de Ver te b r es des sable s de Montpellicr.3: Ca rniv or a Fissip edia. Trav. Lab. Geo!. Fac. Sci.,37, 7-26. Ly on. WAGNER A. 1837. Pa la on t ologische Abhandlu n gen. Gel. Anz. Mitg l. k. bayer . Akad.

Wi ss., 5, 168, p. 320; 169,327-328; 170, 334-33 5. Mtinchen.

JAN ST A CH

AGRIOTHERIUM INTERMEDIUM N. SP. Z PLIOCENSKIEJ BREKCJI KOSTNEJ WYDOBYTEJ Z MIEJSCOWOSCI WEtZE W POLSCE

Streszez en ie

Autor opisuj e szczatk i Agriotherium Wagn, (= Huaetuirctos Falc. & Cautl.) z brekc j i kostne], wydoby tej w miejs cowosci W~i.e kolo Dzialoszyna (woj, loozkie), porniedzy ktoryrni zachowaly sie stos u n k ow o dobrze pierwsz y i drugi zab trzonowy

AGRIOTHERIUM INTERMEDIUM N. SP. FROM THE PLIOCEN E 15

gornej szcze k i oraz nieuszkodzony pierwszy zab trzonowy dolnej szcze ki tego same- go osobnika, dozwalajace okreslic z wszelka pewnoscia rodza j tego zwierzecia ,

Autor zestawia wszystkie (24) opisane do t y chczas ga tunk i ^q; pierwo tne j sy st erna- tycznie grupy Hyaenarctos, rozdz iel a n ej obecnie na dw a rodzaje : Agri oth er i u11!

Wagn, i In d ar ct os Pilgr., pod ajac, wedl ug danych zaczerpnietych z prac roznych autor6w,rozrniary zebow P!-M2 g6rnej szcze ki iM₁ doln e j ora z miejs ca znale zienia tych gatunk6w.

Szczegolowe por6wn an ie zebow trzonowychosobni ka znalezionego w Wezach wy- kazal o, ze nalezy go za liczyc do rodza ju Agr io theri u m Wagn., bowiem brzeg doj e- zyk owy korony pierwszego zeba trzon ow ego g6rnej szczeki jest u tego osobnika kr6tszy niz brze g zewnetrzny, gdy tymczasem u Indar ctos Pilg r . prz ekr6j pods tawy korony tego zeba jest niemal kw ad r a to wy, a w dr ugirn zebie trzonowym M2 brak zupelnie talonu, ktor y u In clar ct os jest na tym zebie mniej lu b wie cej wydatnie wy - ksztalcony.

Nieco trudniej okres lic do kl a dnie gatunek Agriotherium zn a lez io riego w Wezach. Dotychczas z pli ocenu Eura zji pozn ano cztery gatu n ki, kt6re na pod s ta w ie ksztaltu zebow trzono wy ch zaliczo no do rndz ajuAgri ot h er ium Wagn, Autor por6wn uje ksztal t zebow okazu'z W~i:6wz ksztaltem zebow tych czterec h gatun k6wi docho dzi do prze- kona ni a, ze pornied zy tyrn i zeb ami zaz n a cza j a sie wyr azn e roznice, Ok a z z W~z6w

pod tyrn wzglede rn zajm uje jakby pos r ed n ie stanowisko porniedzy A. insigne (Gerv.) poznany m.z Fran cj i (Mo ntpellier) a A. sivalense (Falc,^& Cautl .)zSiw a lik w In diach. Dla zobrazowania roznic wystepujacych w ksztalcie zebow trzonowych porn.edzy osobnikiem z W~z6 w

a:

innymi gatunka mi, autor zes ta w ia na pl. II ksztalt zebo w czter ech pokrewnych gatu nk6w Agri otheriu m Wagn. wedlug ry cin , podanych w opi- sach tych gatunkow przez in nyc h autor 6w. Auto r nadaje gatu nkow i z W~6w naz we Agriotherium in te r medi u m n. sp .

Obszar zamieszkaly prz ez grupe zwie r za t obejmow anych wsp ol n a nazwa Hyac- narctos byl bardzo rozlegl y , zajmowal bowiern niemal cal a polu d ni owa i srodkow a Europe, Iacz yt sie dalej ku wschodow i z siedzibami ich w AZlj i i rozprzes trz enial si ~

sze r ok o w Ameryce P6ln ocne j.Pozw a la to przypuszczac,ze w pewnym

,

okresie czasu warunki i:yciowe na duz e] przestrzeni w Holarktyce, przynajmn iej w pewnej sze- rokosci geograflczne], byly dla tej grupy zwierzat ⁰ He nie [ednakowe, to bardzo do sieb ie zb lizone, i da w al y poczqtek powstawaniu form geograficznych czy ek ologicz- nyoh mal o sle ad siebie roznia cyc h.

Jei:e li na w et z tej wsp61ne j grupy wylaczymy formy, obej mowane obecnie nazwa rodzajowa Agri otherium Wagn ., to wystepowanie A. insigne (Gerv.) we Francji,

A.

intermedium w Polsce i A. si val ens e (Falc. & Cautl.) w p6lnocnych In d iach (Si- walik) wskazuje na istnienie w pewnym okresie czasu nici wiazace] formy bardzo do sieb ie zbliionena dl ugim szla ku od Fr a n cj i do Indii.

Ok res lenie czasu wystepowania grupy Hy aen ar cto s napotyka na pewne trudno- sci, nie zawsze bowiem zgodnie ustalany jest przez geolo gow wiek zI6i:, w ktorych

16 ^{JANST ACH}

znajdowano szczatkt przedsta w icieli tej grupy i waha sie on w granicach od gcr- nego miocenu do gor neg o plio cenu . Zlo:i:a, w ktorych znalezio n o szcza tki Agriothe- ri u m Wa gn., datowane sa zpliocenu; srodkowy lubgor n y pliocen zdaje sie bye okre- sem najwieksz ego rozwo j u gat u n k ow tego rodza ju . Na tom iast szczatki Indar cto s Pil gr. spoty k a n e sa czesciei w dolnym, anizeli w srodkowyrn pliocenie, i Indarct os uwaz an y jes t (G. E. Pilg rim. 1931; H. Tobien, 1952 i in.) za cha r a k tery s ty czny ele- ment fauny hip paric nowej w poludniow ej Eur opie i Azji.

Pon iewaz drapiezne , pozna ne dotychczas dokladniej z brekcji kostnej W~:i:o w,

zb lizaj a sie do elernentow wymarlej fauny, zn a n y ch ze zl6:i: Mon tp ellie r we Fr ancji, a czas powstania tych z16:i: przesuwany jest z dolnego do srodkowego pliocenu, prz e- to i szczatki Agriotherium inte r m edi m n nal e:i:aloby datowac jakopocho dzace ze srod- kowego plio cenu.

Jak k olw ie k gat unki rodzaju Agri ot he rimn Wagn. morfol ogiczn ie Z'blizajq sle ba r - dzo do ga tu nk 6w rodzaju In d arct osPil g r , i prawdopo do b nie mialy wspolnegopr zo d- ka, a takze niemal ten sam czas swego powstania i wygasniecia, to jednak ich 1'0-

dowe linie biegna oddzielnie. Pr zod ka ic:h szu kac nale:i:y w miocen ie i kryje sie on wsrod rzes zy drobnych form rod za j u Ur savu s Sch loss. Pomij a jqc r6:i:nice wlelk osci, zeby trzonowe Ursavus Schlo ss. sa tak zbliz one ksztaltem do ty oh ze u Indarct os Pilgr., ze niektore gatu n k i rodza ju Ursav u s byl y pocza t k owo oznaczane jako nale - zace do rodzaju Hy aen ar ct os (Fa lc. & Cautl.), np. Ur savus br ev irh i nus Hofman n , 1887, opisa ny przez M. Schlossera.w 1887 r. ja k o Hy aen ar ct os minutus Schlos s., a Ursavus depereti Schlosser, 1902, opisany przez

c u.

Depe re ta w 1895 r. jak o Hyae- nar ctos arctoidens Dep.

Osob niki rodza ju Ag rio t he rium dos iega ly znacznych rozm iar6w cial a,gdyz czasz- ka A. sivalense (Falc. & Cautl.) miala , wedl ug R. Lyd ekke r a (1884) 482,6 mm dhrgo- sci, czylizwierze wielko scia zbliza l osie do bardzo duzy ch ok az6wniedzwiedzla [aski- niowego. Niew ie le mniejsza mus iala bye czaszka A. interme dium, gdyz - biorac pod uwage stosu ne k dlugosc i zebow MI i M2w gor rie] szczece u A. siva len se do dtugosci czaszki tego gatunk u (57 mm:482,6 mm) - przy dlugos ci tych zebow u A. in te r me - dium, kt6ra wynosi 51 mm, dlug osc czaszki tego gatunku wynosilaby 431,4 mm.

Silnie roz w ini ete zeby tna ce u wszystkich ga t u nkow grupy Hyaenarctos wskazu- jq, ze pr ze dstawiciele tej gr u py byly dr ap ie:i:nik ami 0 potezn ych roz m ia r ac h ciala . Duz y zab przedtrzo nowy czwarty g6rnej szczeki, podobny kszt a ltem do tegoz zeba u hieriy, ula twial zapewne zwier zeci u kruszenie nawet duzy oh kos ci. Spos6b zycia A. intermedium mo gl wiec bye podobny do zyci a hien. Mogl y to bye zwierzeta ste- pow, z wystep uj acym i gdz ie n iegd zie sk upienia mi drzew i kr zewow, szcze go ln ie w pob lizu zb iorn ik6wwody, kt6re z cala pewnos cia rnusialy wystep owa c w pewnym ok re sie czasu w okoli cy W~:i:6 w.

AGRlOTHERIUltf INTERMEDIUM N. SP. FROM THE PLIOCENE

OBJMiNmNIA DO ILUSTRACJI

17

PI. I

Fig. 1. Fragment prawe j str ony g6rnej szczeki z resztka przedtrzonowego czwar- te go i zeb a mi tr zon owymi pierwszyrn i drugim.

Fig . 2. Siekacz tr ze ci z lewej stro ny gorne] szczeki,

Fig . 3. Fra gment doln ej szczeki z lewej strony Z resztk a czwar teg o zeba przed- tr zonowego,

Fig. 4. Pie r w szy tr zo n owy Z lewej strony dolnej szczeki, wid zi any od str ony do-

j~zykowej.

Fig. 5. Pier w szy zab trz on owy z prawej strony dol ne j szc zeki: a od strony ze- wnet rzne], b od strony dojezyko w e],

Wszystkie okazy wie lk osci natu r alne j

PI. II

Fig. 1. Sze re g zebow gor ne] szczeki pt -M2 Agr i oth er i u rn in si gne (Gerv.), wg Ch. Fricka (1926-30); niece zmniej szon e.

Fig .2. Szereg zebow gor ne] szczek i p4 -M2 A. sivalense (Falc. & Cautl.), wg Ch. Fric'ka (t. c.) ; nieco zmnieiszone.

Fig. 3. Szer eg ze bow gor ne] szczeki p4 - M2A. pal aeindicum Lyd ., wg Ch, Frick a (l. c.); nieco zrnniejs zon e .

Fig. 4. Dr ugi zab trzonowy go r ne] sz cze k i M2 Agrio ther i u m sp. wg H. He lb ing a (1932) ;wi elk. nat.

Fig. 5. Pie r wszy zab trzonowy dolnej szczeki 'M_l Indar ct os sp. z Sa mo s, wg H. Helbinga (I. c.); wielk. nat .

Fig. 6. Te n sam zab, widziany z g6ry, wg H. Helb ing a (I. c.); wielk. nat.

Fig. 7. Pie r ws zy zqb trzonowy doln ej szc zekiM_l A. insigne (Ge r v.) z Montpelller, widziany z gory , wgJ. Vireta (1939); wielk. nat.

5lH CTI\X

.4G RIOT HE RIUM INTERM E D lUM N. SP. 113 lIJIHOUEHOBOr:I HOCTHOn BPEKYHH H3BJIEYEHH Or:I B MECTHOCTH BEHlKE B IlOJIb lll E

ABTOP OHHiCblBaeT !ppalI'MeHTbl3y6oB Agriotheriu m Wagn. (= Hyaenarctos Falc.

& Cautl.), 'H3iBJI€''I13HHbIX ill3 nJllHOl.\ e HOllCH HO(" I1HOH 6p e.K 'IHiH II MecllHOGTH Bemxe

;; Floasur e IHonpenenser HX JIlpHHU'AJIem:-!ocTb H HOBO:l>lY BHAY Agrio the rium inter- medium, HOTOpblH3a HliJlWeT Bc1'poeIllm HOp OHblIRope HlHblx 3y60H, Hoc06eHHocTHlV1²,

!Wilt 6bl npoaenryrosayio cacreaeraxecayro nO~H ~HIO IMelfW A. insigne (Gerv.) .lI3

Mo a nen se (M on tpelli e r) BO CP,pa HI.\HH li A. sival~n se (Fale. & Cautl.) 1113 OH8<·l.1lJl~a (Siw ali k ) B HH~lIlli.

Agriotherium intermedium 6b lJl cocrasuoa 'I<lCTblO clJa y H bl 6J1:Hi3ROH (paYJllc iH3

Monnense H ern reoJlOI'H'l00K1IHso3pacT 6b lJl Tacrrnte sepoanao cpenne-nnnouenoau n.

06pa3 iltH3HH A. intermedium 6blJl He.pOfl11HO 6J1H30 X oop as y iHH3HlI Men, 'llO~06110 OC1'aJIb llbl M HpeACTaBHTeJlIIL\1 aron npyrmst,

Acta Pala e on t ol o glca Pol onica - \"01.II/I 2

Fig. l.

Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

PI. I

Fragment of th e righ t maxilla wit h damage d P4, M' and M2.

13of the left maxilla. .

Fragment of the left mandible with P, strongly damage d. Mi of !!he left mandible, lingual view.

M, of the righ t mandible; a labial view, b lingual view.

All f·igu res of na tur a l size

PI.II

Fig. 1. P~_M2 of Agriother iu m insigne (Gerv .). aft e r Ch. Fric k (1926-30) ; somewhat reduced.

Fig. 2. J>4-M2of A. sivalense (Fa lc. & CautI.), after Ch. Frick (1. c.); somewhat re- duced.

Fig. 3. J>4-W of A. pa l aeindicu7Jl. Lyd ., afte r Ch. Frick (1. c.): some what reduce d. Fig. 4. M2 of the right maxilla of Agriother ium sp., after H. Hetbing (1932); nat.

size. .

Fig. 5. MI of the mandible of ln d m'ct os sp. fr om Sam os, after H. Helb in g (l.c.l; na t. size.

Fig. 6. The same tooth, top view, after H. Helb in g (1. c.): nat size.

Fig. 7. Mt of mandible of A. insigne (Gerv.) fr om Montpelli e r, top view, after J. Viret (1939); nat. size.

.ACTA PALAEONTOLOGICA POLONICA, VOL. II

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