-'
C
\ '01. II
T
A
I'ALAEONTOl.O
GICA
POLO"';'
, 957
CA
No. I
JAN STACH
A GR I OTHERIUM INTERMEDIUM N. SP. FROM THE PLIOCENE
BONE BRECCIA OF WEiZE
Study on th e Ter tiar y bone b r eccia F auna from WE:ze
near Dzialoszyn in P oland
P ART VII I *
.4. bst r act . T he writer describes frag me nts of den tition of A g riot he rium Wagn .
( = H ya ena rc t os F a lc. & Cau tl.) fr om t he P liocene b one breccia re covered a t Wp,ze
n ear Dziat oszyn, P oland. He refers t hese remains to a n individual of a ne w
species fro m genus A gr io t he rium i n te rme dium, which occupies an in termediate sys tem ati c position between A . insigne (Gerv.) fro m Montpellier in F rance and A . sival en se (Fa lc. & Cautl.) fr om S iwalik in India , in w hat conce rns the sh ape of th e
crown of mol ars, th at of M2 in partic ular.
DESCRIPTION OF MATERIAL
The bone breccia discovered at WE:ze n ear Dzialoszyn fr om whic h
th e writer has already described ArctomeLes pliocaenicus, Ursus wenzensis and Nyctereutes sp. ', all belonging to the group of carnivores, has also
yielded the remains of Agriotherium ( = Hyaenarctos), 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 considered are small, mostly detached fragments,
m ainly referable to one individual.
1) The largest of these remains is , a fragmen t of the right upper jaw ,
showing remnants of the fourth premolar and t he first and second mola rs in a damaged condition.
The other r emains consis t of:
2) .lingual edge of cro wn of left upper secon d molar (M 2);
" P arts I-V - see A ct a Geo!. Po!. , vol. II-V/1952-55 ; parts VI-VII - Acta P a!aeon t . Pol., vol. 1/1956.
1 Arctomeles pliocaenicus n. gen. & sp, from the Melinae s ub fam ily. Acta Geo !.
Po!., vol. II/195I. Ursus we nzensis, new species of a s mall Pliocen e bear. Ibid., v ol.
I II/I953. Nyct er eu t es (Canidae ) in th e Pliocen e of Poland. Ibid., vol. IV/1954.
JAN STACH
2
3) b r ok en off ex ternal cus p of right up per firs t molar (M');
4) posterior portion of the right upper fourt h p remola r (P4);
5) fragment of left ma ndible with re mnants of lower fourth p remolar (p.) ;
6) a detached, well preserved lower third in cisor (13);
7) a likewdse beau tifu ll y preserved complet e left lower first mola r
(M 1 ) ;
8) ri ght lowe r f irst molar (M1) , sli ghtly damag ed.
T he fra gment of right maxilla, mentioned a t the head of the her e
specifi ed remains, is, by far, the most diagnostic specimen:
The upper fourth premolar of this fragment of maxilla is b adly damaged , th e crown and the anter ior part bein g altogether m is sing. The
ou tlin e of the base of the cr own in the 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 shape an d size . The only conclusi on of some prob ab ili ty is tha t the len gth of the to oth h ad b een 29 mm.
In this it agrees wi th the size ascertaine d for that same tooth in Agrioth eriu 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 account the 12 mm length of the
broken off pos terior cusp (metacone) of the tooth under item 4, an d b y
com paring it wi th the same ousp in A. in sign e (Gerv.) fr om Montpelher
whic h is likewi se 12 mm long, as fig u re d by Frick (1926 -1930, f ig. 20)
in a na tural size drawin g of the 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 the four cusps are missing, having b een
partly broken off after the animal had perished and partly w orn 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 measure its size. The
base shows to be of sub-quadrate shape with corners somewha t r ounded .
Ex te rn all y the tooth is 26 mm long, 25 mm medially and 22.2 mm in ternally. The crown of the tooth is thus narrower lingually an d with inn er
border m ore rounded. The 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 erior cu sp (m etacon e) in this
toot h may be figured out 'by measu r ing the b r oken off cu sp of an ot her M' ,
und er item 3. This cusp , as measured externally,is 15 m m high inclu di ng
t he cingul u m, 10 mm with out the Cli n&ulu m, an d 13 mm w ide. The summit
of the cus p is pointed, the edges consp icu ous, the s u rfa ce wrinkled , this
b ein g better recognizable on t he external si de of the cusp.
Th e upp er second m ol ar (M2) is in a fa r bett er state of preservation,
the paracone and th e external border only b ein g m issing. Perfectly pres erved are it s secon d ex ternal cusp (metacone) an d the whole ldngual
AGRIOTHERIUM IN T ERMEDIUM N . S P . F R OM THE P LIO CENE
3
part of the crown, including the cingulum, and th e ridg e running lengthwi se through the centre of the tooth. The crown of this tooth is Like wise
sub-quadrate in shape, with rounded a ng les, The length here is 23.5 mm
ex ter nal ly, 24.5 mm at mid-len gth and 25.2 mm on the lingual side ; the
wi d th is 26 mm a n te r iorly, 25 mm m edially an d 24 ' m m po s te riorly .
These three dim ens ions are onl y ap prox im ate sin ce the external b order
has not been p reser 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 metacon e as m easured
ex tern all y, w ithou t the cingulum, as this is missing, ds 6.8 mm, the length
of t he ed ge d escending lingu all y is 8 mm, t hat of the posterior edge, measured 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 hape of an almost regul-ar pyrami d, wdt h well sh owing
anterior, lingu al an d po ster ior edges , an d fine ly wrinkle d surface. This
cusp was separated from the anterior cus p (pa r ac on e) b y a rather p ro minent de pressio n . A gently cu r vin g ri b, in the shape of a diagonal
ex te n ding lingu ally to the poste rior angle of the s qu a r e, an d well p r eser ved over its complet e length, branches off from the po sterio r border
of the missing paracone. H is difficult to in d ica te the exact position of
the lingual cusps on this r ib. The grea te st h eight, 3.8 mm, is at ta in ed by
the ri b medi ally, opposit e to the depression sepa r ating the parac one
fr om the metacone. It is als o her e that the r ib attains its maximum width
of 6.5 mm, while a t it s in itial and termi nal part it is 3.8 and 3 rom respective ly . Auelong a ted, more or less tri angular area separa te s the r ib
along its entire lengt h fr 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 covered by den sely arrange d bu t
s ha ll ow w ri nkles . Ther e is no li ng ual extension whateve r of th e pos te r ior
end of the crown, i. e. no talon. The circumference of th e to oth consis ts
of a si ng le cin gulum, particularl y high (6.8 mm) on th e lingual side of
the crown .
A detach ed fr ag ment of the lingual edge of th e left upper se con d '
molar (item 2) is probably r ef er able to th e s a me individual even though
i ts length (26.5 mm) exceeds some wha t that of the lingual borde r of the
just described M2. In .this fra gmen t als o is the dense transverse w rinkling
readtly recognizable over the s u rface s eparatdng the rib from the cingulum.
A fragment of t he Left mandibular branch has been recovered consi s tin g of a part ex ten ding from the posterior end of the canine to the
be ginnin g of the first molar (M1 ) . The length of this fragment is 48 mm,
the height - 51 mm. Of the dentition a trace of the lower third premolar (P 3) has been preserved, with fragment of a single root e mbedd ed in
a rather small alveolus, 5 mm in diameter, al so of the lower fourth premolar (P 4 ) directly adjacent. It is this remnant of the fourth premolar
4
J AN STACH
w h ich ma de it possible t o asc erta in t he e x act length of the to oth to be
18 mm an d i ts m aximum m edian b r eadth as 10 mm. The area between
t he 'c an in e t ooth w hic h is m iss in g h e r e, 'a n d t he alveolus of t h e thi r d
premolar does no t s how any t r ace of teeth.
Very w ell preserved is the crown 0.£ a de ta ch ed left low er th ird incis or (1 3 ) , In s hape it mark edly resem bles t h e crown of this in cis or in
b ears. A s m all e r accessory 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 separated from the
cen tral sh aft b y a sh allo w groove, w h ile linguall y three nibs des cend
from the s um m it of the crown to its base, na mely along the inner edg e
of the sh aft, mediall y, an d fro m t h e external edge,. uniting a t the b as e
of the crown . The h eigh t of the cen tral s haft measu redexternally f r om
th e b ase of t h e crown is 17.5 m m , that of the slide branc h 13 mm; th e
su m m it of the shaft overtops b y 4.5 mm the s u m mit of the s ide cusp.
The wi dth of t he crown measured at the b ase is 11 m m , w h ile th e t h ickn es s of the to oth at dts b as e is 12 mm.
In a very sa tisf a ctor y state of p r eservation ar e . also two' de tached
s pecimens of the rig h t an d left m andibular fi rs t m ol a rs . They both, prob ab ly, belonge d to the same individual as they differ neithe r in shape no r
in di mensions . Their s ize is r at h er large, the basal len gth of the cr ow n
:b ein g 37 m m, frontal wi dth 17.5 mm, that in the mi ddle al ong th e Iine
o f the central CUSIP (protoconid) 18.7 m m, while at the e n d, w ith in t h e
t alonid, i t is 20.5 mm. The p r in cip al cu sp (pr otoconid), w hich is stro ngly
d ev eloped, attains 21 mm w he n measured fro m the lower border of the
cingulum on th e outside . Its b asal 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, runs from the s u m m it anter iorly,
uniting directly with the posterior r id ge of the preced ing ant erior
c usp. T he s h or ter (4 m m) posterior ri dge b ifurca tes only justa little bel ow th e s um m it t o f or m two s harply ou tline d ri dges , 8 mm in length,
wh ich enci r cle th e fla t triangu lar postenior wall of the 'cus p , s t eep ly
d escen din g to the dep r ess ion of the talo ni d. The lateral extern al
ri dge is p oorly disc ernib le, w hereas numer ous fi ne wrin k les are well
shown at this s ide , arran ge d a lmost p ar allel to ea ch other , from the
s um mit to the cin gul u m. The anterio r cu sp, the pa r aconid, measured at
the base , as 12 m m thick an d 11.5 mm lon g . Its h eigh t m easured from the
Uo wer marg in of the cin g ul um is 15 mm. It sh ows on ly t w o distinct
ri dg es, the anteri or one descen di ng fr om the s u m mit to th e b as e w here
<it unit es w it h th e cin gulum, and the p osterior one, 6 mm in length . The
ex ternal surface of that cu sp is als o fin el y w ri n kl ed. On the li ng ual sid e
the summit of the m etacon id r ises b eyond the main cusp. Its h eigh t meas u r ed inte rnally f ro m the low er m a rgin of the cingu lum is 13.5 mm,
w h il e m easure d fro m the side of the talon id it is 6.5 mm ; t h e meta-
AGRIOTHER IUM I N TERMEDIU M N . SP . FROM T H E P LIOCE NE
5
conid is 7.75 mm long. Ant eriorly the me taconid to uc hes th e base of the
large protoconid, forming with it an d w ith the posterio r cusp th e trigon id , 28 mm long. Re ally it fo rms the beginning of the internal d elimitation of the talonid. Beyond it, on the lingual side is the conspicuousl y lower e nt oconid, w ith h eigh t of 10 m m measu red from the low er m argin
of cingulum, followed by the still lower entoconuli d , which pa r tially
closes up po steriorly the taloni d by its arcuate low summit ri dge . The
hypoconid rises externally, opposit e the two la st small lingual cus ps . Its
,heigh t measur ed ex te rnally from the lower ma r gin of th e c ing ulum is
14.5 rnm, th a t m easur ed f ro m the talon idal dep r ession bein g 6 rnm ; it s
thickness is 12 mm, the posterior su mm it r idge is 5.8 mm long, w hile
the anterior, reaching to the base of the cardinal cusp is 8 mm long. No
cusp or th ick ening is discernible on this r idge.
The talonid, being thus lingually delimite d by three cusps - the metac on id in clude d - and labially by the hypo con id an d its anterior r idg e,
is re lative ly b road and rather long . The ci ng ulum surrounding the crow n
at the b ase is high (7 mm) on the outer s ide, b elo w the h yp oconid, that
is to s ay posteriorly. It is, h owever , relatively weak ly marke d, near the
s u mm it lit ascends and becom es almost entirely obli ter a ted, to be again,
though very indistinctly discernib le, nea r the anterior cusp. Lingually
the c ingu lu m is almost completely obliterated.
GENERIC AND SPECIFIC DETERMINATION
Th e shape and size of the here described teeth lead with perfec t s urety to the inference that they formed part of the den tit ion of an indiv id ual belonging to a group of species or iginall y k nown under the
common generic name of H sjaenarcto s (Falconer & Cau tl ey).
Re mains of an individual from this genus were first recovered from
Pliocene deposits i n the Siwalik Hills of India. They consisted of
a s tr ong ly damag ed right mandible wit h P 4 , M 1 and M 2 , also with alveoli
referable to two p remolars and M a, which are all missin g. The nex t find
was th at of part of a skull w ith preserved canines, P4_M2 in a dama ged
condition, and alveoli of P 2_P3. Th ese r em ains were in 1836 descr ib ed b y
H. F alcon er a nd Cauil ey and r ef err ed by the m t o th e skeleton of a b ear
which they calle d Ursus siv alem is F alc. & Cau tl .
The autho rs themselves, an d la ter in 1837 also A. Wagner , laid emphasis on the dentition of the des cribed specimen not agree in g with th at
typical of the gen us Ursus, so muc h so that th e ge ne ric n ame of Agriotherium was suggested by A. Wagner fo r the in dividu al from the S iwali k
H ill s. So me ti me af ter th e n ame of Huaenarct os Falc . & Cau tl . was introdu ced for that g enus by R. Owen who ass erted that the auth ors h ad
6
JAN STAC H
o r iginally intended to call their specimen by this name gui de d b y the
resem blance in shape of P ! to that tooth in the hyena. T his n ame, a ccepted
by H . Falconer (1848) as s ubgeneric for this type of fossil m ammal , h as
b ee n since used by palaeontologdsts as generic.
Gen us Hu aemarctos (Falc, & Cautl.) has been subsequently s plit up
in t o g enus Agriotherium Wagn. with Agriotherium sivalense (F alc. &
Cautl.) as genotype, genus Indarctos P ilg r im with In 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 h as not b een generally accepted, with Hy 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 ap e of dentition , of t he upper jaw particularly. T hey
h ave been summed UiP by Ch . Frick, W . D. Ma tthew, G. E. Pilgrim and
others. The chi ef on es are as follows :
1. In Indarctos Pilgr. t he anterio r extern al Hceesso ry cus p, the parastyl e, on P\ is poorl y developed while it is prominen t in Agriotherium
W agn.; in Agriotherium this tooth is large an d longer than the next
molar, MI.
2. In In darctos Pilg r. t he b asal se ction of M! is sub-quadrate, w hile
i n Agriotherium W agn. the lin gual border of the crown is s horter than t he
labial an d with angles distinctly r ou nded .
3. In Indarc tos Pilgr. th e labia l poster ior bo rd er of M2 extends into
a m ore or less prominen t ta lon, whil e in A griotherium W agn. t here is n o
tal on on th is tooth .
4. In Indarctos P ilgr. M, is rela ti vely long, with t he t alon id s omew hat shorter t han t he trigonid, a n d the h yp ocon id lower than the ento conid , w hi le in Agriotherium W agn . M, is r el a tiv el y sh or t, with the
t alonid m ark edly s h orter than the t rigo nid, 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 s pe cified, 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
g en u s Agriotherium Wagn. N amel y:
1) MI is wi th the li ng ual bo rd er distinctly shorter t han the labial
a n d with angles r ou n ded ;
2) t h e ling ual posterior b orde r of M2 d isplay s n o p osterior con v exity ,
which m e an s t hat t he crown is witho ut a talo n ;
3) t he an terior p art of M l , the trigo ni d , is cons iderably l on ge r than
the p ost erior, the taloni d; t h e crown of this t ooth is larges t w ithi n the
ta lo ni d, a n d t he h ypoconid is h ig her t han the ento con id .
It is fa r more di ffi cult specifica lly t o ide n tify t h e re mains of Agriot herium, fo u n d in the W r::.ze b r eccia.
AGRIOTHERIUlvI I NTERlvIEDIUlI1 N . SP . F R O M T H 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 even ted th ei r undoubted id en tification wi th this or anoth er speci es of
Agriortherium Wagn. or of Indarctos Pilgr.
For this r eason such r emains were mostly referred to some species
within the group of Hy,aenarctos w it h out generic differentiation into
Agriotherium and Indarctos. In this wa y, a score or so of sp ecies have
be en es tablished - part of which are probably id entic al - or th ey were
s imply called Hyaenard,tos sp.
The r ecove ry, therefor e, from the sa m e depo sits , within so small an
a r ea as that occupied by the W p.ze brec cia , of fragments of both the upper
a n d lower dentit ion of an in divi dual, undoub tedly re fer able to one species
fr om ge n us Agriotherium, provid es impor tant evide n ce for the knowledg e of the dentition of this species.
Since d ental dimensions s er ve as s up plementa ry evide n ce in spe cific
di ag nosis and s inc e th e gr oup of forms known under the common name
of Hyaenarctos has a ve ry wi de r ange of dis tribution within the Hola r cti c area , it seems mos t ad visable to t abulate data concerning the
di m ensi ons of the here studied teeth .in spe cies thus far ide ntified f rom .
the ge neral group of Hyaenarctos (see t able on p . 8-9).
A revie w of the sp eci es li st ed in the he r e annexed t ab le shows th e
predom in ance, within the en tir e H olar ctic a r ea, of species referred b y
their au th ors to genus Indarctos P ilg r , The predominance wo uld , perhaps,
be still gr eat er, should the u pper se cond mo lar be always present in t h e
r emai ns of forms b elonging to the Hyaen arct os grou p, recorde d from
different sites. On the pres ence or a bsence of a t alon in this to ot h, the
remains could without doubt be referred to some spe cies w ithin gen us
In darctos or Agriotherium r es pective ly . Sp ecie s tentatively determine d
as Hyaena rctos sp , would then also disappear from the list.
The assignmen t of a fo rm to genus Agriothe rium or to ge nus Indarctos
on the firs t molar on ly , does not always seem quit e justifie d to the writer .
A comparison for exa mple of this too th,collected fr om the W p.ze breccia,
with th e same to oth recovere d fro m the .islan d of Sames , the shape of
wh ich is excellently shown on ph otog r aphs in a pap e r by H . Helbing
(1932, fig. 3 a-b) , here reproduce d in pl . II , fig . 5-6, shows their al most
iden tical s imilarit y . In both , the h ypoconid is high er than the entoconid
an d a distin ct entoconulid is p r es en t in additio n to the en t oconid. An d yet ,
on the a bsence of talon in the upper second m ola r in the Wp.ze specimen ,
it is ge n us A gri otherium, while the prese nce of a p ro m inen t t alon in the
same toot h b elonging to the Sam os s pecim en refers it t o genus Indarctos.
The fo llo win g are, aft er Ch . Frick (1926-3 0), sp eci es undoubt edly
r efe r ab le to Agriotherium Wa gn .:
J AN S TAC H
8
T able show ing dimens ion s of P '', M ' , M ' a n d M ,
in species w ith in th e g ro u p of Hyaenarc tos (in rnm j
pI
Spe ci es
M'
Patria
M,
in t er m edi1!111
n. s p .
25 / 25
24 .5
X
37
25
X
W ~ze,
20.5
Pol a nd
in si gn i s
G e rv a is , 1853 29. 1 X 21
27.4
27.8 Y 27.5
v
26 .5
41. 5
i nsi gni s
40
38
Gerva is, 185 3
sp . He lb in g,
X 22.5
X 21.5
24
25.2 X 28.5
1932
Boutonne t,
F r a n ce
I
sp. G erv a is,
30 \, 23
1853
',
I
Montp ellier' l
F r a n ce I
1
I
!Ge n u s
i
Vi alette,
France
IAlCOi, Spain Agr.?
sp. Fl ow er,
1877
p on t i cum
K ormo s, 1913
30
X 30
I
41 :x 19.5
M e- \
neghini, 1863
Felixtow,
England
A gr .?
Baltavar,
Hungary
Agr.?
laurillardi
a r ctoides
Deperet &
I
Ll euca, 1895
1
24.6
24.6
X 22
I 28 .2 X 22
urct oides
Deperet &
,
Ll euca. 1895 \
IMon te B a m- Agr. ?
boli , It aly
33
i
27
)<
21
27
X
Or ignac,
F rance
25
29 X 21
!
30 X 24.5 1
23.5
utti cus Dames, 1
1883
Ind .
I
IMontredon,
France ,
I Pikermi,
Greece
,I
40
I'
IJir e ti Villatta
& Crusafo n t,
Valles-Peln edes, Spa in
1943
sp . H elb in g,
26
29.1 X 24 ,3
1932
29
X 27
34
X 26
141.6X 22.2 II
I
silJale n si s
palaei ndicus
L yde k k e r ,
1878
:G au - Wein h.
Ge r m any
25 .1 X 18.3
I
I
I
F al coner &
C au tl e y , 183 61
pu njabiensi s
L y d e k k e r,
1878
Greec e
I
s p. T ob ien ,
1952
Sarn os,
32
>< 22
29 X 29 .9
29
32
~<
30 . 28
31 X 27
28 X 23
22 .5
27
I
X 25 .5 I
X 30.5
27 )! 2 7.5
38
S iwa li k .
Ind ia
Ag r .
42 .5 ! 2 1.5
Siwali k ,
I nd ia
I nd .
Siwalik ,
I nd ia
I Ag r .
I 39. 5 .
21
AGR IOT llER lUlYl INTERlYlEDI UlYl N .
se.
F R OM T HE P L I OCENE
9
T a ble s h ow ing d im en sio ns of p I, M I, M ~ a n d M I
in species w it h in t he grou p of Hyaenar ctos (in mm )
(con tin ued)
I
pI
Sp e cies
MI
M~
M,
Patr ia
Ge nus
44 X 23
M a r agh a ,
P e rs ia
Ind .
mara ghanu s
Mequenen ,
1925
sa lm on t an us
I
35 X 27
29 X 25.5
P il gr im , 1913
Salt Range ,
Ind ia
I
lagre lii
Zd ansk Y,1 924 27.3 X 21.4
Zd a n s ky , 1924 1
42.8 X 21.4 Pao-T e di Sh " 1
Chi n a
,
-
32.3 X 21.7
35.4 X 25.8
36.5
I
I
I
30.5
X 32.3
i
S ou t h Ch in a Agr .?
I 43.5 X 23.5
,
I
oregon en si s
M erriam,1916 31.7 X 22.6 ,
schneideri
Sellards, 1916
sp , Matthew,
1918
I
\
sp , L yde k k er ,
1844
Frick, 1921
37.5 X 20.8 P a o-Te d is tr .l
Ch in a
i
sinensis
grego r i
32.6 X 23.4
27.3 X 25
I
I
sp , Freud enberg, 1910
-~_!~
. -
-
35.:1 X 27.2 '
I
I 29.8 X 30.2
41 X 23.5 1
I
26 X 24
,
: 46.5 X 24
i
E d en,
California
Agr .?
Ore gon ,
USA
Ind .
Florida,
USA
A gr.?
Snake Cree k A gr.?
USA
I
Tehuichila, Agr ·?
I
M eXICO
1) Agriatherium insigne (Gervais, 1853) ( = HyaenaTdt:os insignis
Gervais) from Montpellier in F r ance;
2) A . sivaLense (F alconer & Cautl ey, 1836) ( = U 7'SUS sivalensis F alc.
& Cautl.) fr om Siw al ik in India ;
3) A. palaein dicum (Lyd ekker , 1878) ( = Hu aenarctos palae indicum
Lyd ek.) from Siw ali k in India ;
4) Agrio therium sp. (Helbing, 1932) ( = Hy aenarctos s p. H el b.) f ro m
Vialett e in F r anc e.
The fo llowing are a lso regar ded by Ch. F r ick (l. c.) as b el on gin g to
Agriot heri um :
Hyaenarctos sp. Gerv ais , 1853, from Alcoy in Spain (fragment of
upper jaw wi th P ' and a r emnan t of the ante ri or part of MI);
H . " schneideri" Sellards, from B r ews ter , Flori d a, US A (M ' only) ;
JAN STACH
Hyaenarctos 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-M2)'
Thus truly but one single species fir om Agriotherium Wagn. h as b een
recor de d from Europ e, namely that of Agriotherium insigne (Gerv.). This
species 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 publishe d in 1853 and 1859 a na tural s ize drawing of t h e fourth premolar and of
b oth molars. Th es e are t eeth with s hape which offers particular interest
ow ing to their 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
h and , described in 1939 b y J . V bret on evide n ce of three mandibular fragm ents colle cted from Boutonnet, M ontpellier. Excellen t photographs
sho w n in table II , fig. 2 a-b of J . V ir e t's paper, enable 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 better comparison of th ese te eth the following drawings are
reproduc ed in pl . II of the present pap er: upp er dentition (P4_M2) in
species A. insigne (Ge rv.), A . siva ben se (F alc. & Cautl.) and A. palaeindieus (Lyd.) , M2 in Agrioth erium sp. (H elb.) , an d lo wer den tition (MI) in
Indarcto s s p . H eLb. and M 1 in A. insign e (G erv.) described by J. Viret.
Wh en comparing th e size and shape of the maxillar molars in the
W Gze. speci men with d imens ions in ot he r spe cies of Agriotherium s p ecifie d in th e an nex e d ta b le , a lso with the d r awing of tee th by Ch . F'nick
inclu ded in h is desc r ip tion of A . insigne (Gerv .), it is observable that the
teeth in the W ~ z e sp e cim en a re some what smaller and the crown in the
b et ter preserve d mola r s omewh a t differen tl y shaped, to say :
1. The l ength of t he an ter ior e x ter n al CUSp2 in the W ~ze specimen
p r actically eq uals th a t of th e next cusp, the metacone (12.2 : 12 mm ), the
b ounda ry line where th e ir b as es m eet is at mid-length of the t oot h, w hile
in A. ins ign e (Ger v .) th e anterior cu sp is di stinctly longer than the posteri or (15 : 11 mm) and the b ases of t he cusps meet beyond the mid-length
po int of the too th.
2. Th e anteri or bo rd er of crown in M2 of th e W~ ze sp eci m en is a lm ost
parallel to the post erior bo rder , whil e in A . insigne (Gerv.) the borders
are inclined to each other at an acu te a ngl e.
2 Th ough in t he W r:ze sp ecimen this cu sp is broken off , w e can see outlined
the re th e cours e of t he cin gulum fr om the edge of w hich the m easurements w er e
ma de. Th e ling ua l edge of th is cusp is dis ce rn ible on the crown of the tooth, as w ell
as the lin e wher e th e base of thi s cusp m eets with tha t of the n ext cus p, the m etacone.
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 osed to the lingual border, with an inconspicuous
d epression at its mid-length, while in A . insigne (Gerv.) the posterio r
border is 'b r oadly r oun ded lingually an d displays a distinct cavity upwa r ds of its m id-length.
In m an y of th ese ch aracters M2 of the W ~ze specimen approaches the
s ame tooth in the s peci me n from Viale tte, b y H. Helbing (1932) defined
as Hya enarctos sp. In the Vialette spe cime n, however, the rib extending
obliquely over the en tire length of the crown is quite distinctly divided
in to separate lingual cusps.
Much closer than to A. insigne (Gerv.) does M2 in th e W ~z e 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 identify with A . insirrne (Gerv.). In A. siv alense (Falc. & Cau tl.) , h owever, ther e is some slight t en den cy to develop
a t al on , ow in g to w hich the lingual border of this too th is som ew hat longer tha n the la bial (27 X 25 mm) , according t o dimensions shown by the
dr-awi n g in Ch . F r ick's pap er.
The lower fi rst molar is less diagnostic in the m atter of comparison
a n 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 t aloni d is 20.5 mm; 1n A. insigne (Gerv.) the se
dime ns ions are not mu ch g reater, being after J. Viret (1939) 38-40 mm
an d 21.2- 22.5 mm respecti ve ly. The t oot h is, ho wever, shaped s omewhat
di ffe re n tl y, to say :
1. In the W ~z e specimen , o n the li ngu al si de of M I , tw o distinct
cusps ri se in addition t o the me tacon id , that is th e entoconid an d the
e n tocon ulid, as an exte nsi on of this border of the crown . In A. insigne
(Gerv .) the ento con ulid is miss in g in this tooth, th e entoconid onl y constitu ti ng besid es the m et aconid the lingual wall of the cr own .
2. The cingulum of the crown on the labial side of the toot h is less
promi ne nt in the W~z e s pe cime n a nd it ,is not discontinuous as in the
tooth of A. insigne (G erv .).
Th e complete obliter ation of th e cr own in M I belonging to th e specimen of A. sivalense (Falc. & Cautl.) im pe des the comparison of its sh ape
wit h that of the crow n °i!I1 M 1 of the W ~z e spe cim en.
On the ground of di ff e r en ces of shape , observed in teeth just mentioned, belonging to the W ~z e s pecimen a nd those in A . insigne (Gerv.) and
A. si'l:alense (Falc. & Cau tl .), an d also in consi deration of the great geo gr aphical dis t an ce separating W~z e from the Siwalik H ills in India which
have yielde d the re mains of A . siv alense (Falc. & Cau tl.), t he present
w r it er believes it advisab le to assign an independent n ame to th e W ~z e
specim en, namel y that 'o f A. i.ntermed-ium n . sp . Some lucky discovery
o f m or e ample m a ter ial , at these or ot he r s ites, will pe rh a ps in th e future
12
J A N ST ACH
lead to the kn owl edge of va r ia tion s, if any, .an d of the ex te n t to which.
they m ay h av e affe cte d the s hape of den tition in in di vid uals f r om Agriotherium Wagn .
GENERAL REMARKS
Fo ss il remains belon ging to animals fro m the g r oup he re considered
an d originally refe rred to under the common n ame of Hyaenarcto s (Falc .
& Ca u tl.) h ave been recor de d from Sp ain, Fra nc e, Gr ea t B ri tain (F elixs tow near Wald fieldr ing), Ge rmany , Poland, Hu ngary, Italy, Greece (P ike rmi, island of Samos), P ersi a, Ch in a, India, also Californ ia, N eb r asca,
Or egon , Florid a in the USA, a nd Mexico. The a rea in habited b y this gro up
of carnivo res is thus s een to have bee n a vast on e, s ince it extended ov e r
ne arly the whole of southern and cen tral Europe an d stret che d on eastw ard to As ia , occupying als o wide terri to ri es w it hin No rth Amer ica.
This suggests g reat similari ty if not ide n tity of li vi ng conditi ons of
this group of animals, prevailing during a ce r t ain p eriod of ti me ov er
im mense territories under th e same ge ogr ap hica l la ti tude. These e nvironmental conditions were probably responsible for the developmen t
of geo gr aphic 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 occurrence of the Hyaenare t cs group. Geologists do no t always agree as t o th e
age of deposits from which fossil remains have b een recovered r eferabl e
to r epres enta tives of that group, the age ass igned t o th em r anging f rom
the Upper Miocene to the Upp er Pliocen e. D epos its which have yielded
remains of Agriotherium are regarde d as of P li ocene age an d it is the
Middle or th e Upper Pliocene periods whic h appear to h ave witnesse d
the greatest specific ab unda nce of this genus . Fossil remains of In darctos, on the other han d , are more copious in the Lower t h an in th e Middle
P liocene and In darctos is by G. E. Pilgrim, H . To bien and others r egarded as th e time-marker for the Hipparion faun a in th e s outh of Eu r ope
and Asia.
The carnivo res from the W ~z e bone br eccia, which have thus f ar
been given closer con sideration , resemb le foss il faun as recorded f r om
the Mcn tpellier deposits in France. Since the age o f these deposits i s
n ow b eing s h ifted from the Lower to the Mid dle P li ocen e it is h ence to-
AGRIOT HE RJUM I N T E R M E D I U M N . SP . FROM T H E P LIO C ENE
13
b e inferred that the remai ns of A. intermedium are li ke wi se assignable
t he Mi ddle P lioce ne age.
Thou gh th e mo rp ho logy of species of Agriotherium Wagn. com es quit e
close to that of Indarctos Pilgr . being probably r efer ab le It o a common
an ces tor and to a contemporaneous date of origin a n d extinction, yet
th ey constitute two independen t stocks. T hei r an cestor is to be searched
for in the Miocene among t he multitude of sma ll forms belo nging to
genus Urs acus S chloss. Differences of size excepted, th e mo lars of Ursav us come so nea r in sh ape to those of In dar ct os that so me species of
Ursavus were or ig in all y r eferred to Hyaenar ctos (Falc . & Cautl. ), as for
ex am ple Ursacus bre'&irhvnus Hofman n , 1887 , by M. Schlosser in 1887
iden tif ied as H u aen oircto« m in utus Sc hloss., .an d Ursav uJs deper eti Schlosse r, 1902 in 1895 by Ch. Dep er e t referre d to Hua enar ctos arctoiien s D ep .
Individuals fro m Agrioth eriu m h ave sometimes abtaine d con side rable
dim ens ions. Th e skull of A . sivalense (F alc. & Cau tl.) is b y R. L ydekke r
repor ted to h ave been 482 .6 m m in le ngth , whic h m ak es the size of the
w hole bo dy of th a t for m e qu al the dimensions of the la r ge st indivi duals
of the cave b ear . The skull of A. in t erm ediu m n. sop. could not h ave b een
mu ch small er . Since, in A . siv a7ense, with le ngth of M' and M2 at 57 mm,
the leng th of skull is 482.6 mm, then, with the same ratio ap pl ie d to A. interm edium , in which the le ngth of a number of teeth attain 51 m m, t he
l ength of skull wo uld h ave to the 431.4 mm.
Th e slt r ong ly developed carn ass ials in all species bel ongin g to the
Hyaer. arcto s grou p in dic at e that .r ep res en ta tives of th is gr ou p were all
carn ivor es wit h po werful bodies. Th e large size of the uppe r fo urth
premolar, shaped very much like that of the Hyaena, must prob abl y have
ma de easy th e crush ing of the bigg es t bo n es. Th e m od e of life of A . int ermedium ma y, .th er ef'or e, h av e r es embled that of h yenas. Its h abi ta t w as
p er ha ps one of steppe 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 reservoirs.
Z oo lo gic a l Institute
o t t he P o li sh Academy of Sci enc es
C r acow B ran ch
Krakow, Sept emb er 19,56
REFERENCES
D E P E RET CH. 1895. Resultats des Iouilles p al eon bolo giq ue s d ans
ri eu r de la collin e d e M ontredon. C. R. h eb d. Seances Acad.
P a r is.
DEPBRET CH . & L LUECA G . 1928. Sur l' Indarctos a rctoides et
Ursides, C. R. Soc. C eo!. France, 4, 149-160 . P ar is.
E RD BR INK D . P. 1953. A revie w of fossil an d recent B ears of t he
D issert. M i n . C eo!. I n st . U n i v . U t r ech t , 1- 2, 1- 597. Utrecht.
le miocene su peSci. , 121, 433-434,
la p hy logenie de s
O ld W or ld . I naug.
14
J AN ST AC H
FALOONER H. & CA UT LE Y. 1836. Note on t he Ursus s ivalensis, a ne w fo ssH s pecies
from S iw ali k H ill s . A si ati c R esearch , 19, 193-200 .
FLOWER W. H . 1877. No te on t he occ urrence of t he r ema irns of H y aenarctos in t he
R ed Cr ag of Suffolk. Qu art . J . Geo!. Soc., 33, 534-536. Lo ndon.
FRICK CH. 1921. E x ti n ct vertebrate fau nas of B adlan ds , of B au t ist a Cr eek an d San
T im oteo Can on , S ou therm California . U niv . Ca li! . Pub l. Geo!. , 12, 277-424 .
B er k ele y.
1926/1930. The He rnicyb n inae a nd a n American Tert iary B ea r . B ull. Amer .
Mus. N at : H i st ., 56, 1- 119. New York.
GE RVA IS P . 1853. Descr ip ti on d es ossements foss iles de mammif er es rap por tes
d 'Espagne p ar MM. de Ve r ne u il, CoUomb e t d e Loriere , B ull. Soc . Geo l , France,
10, 147-1 67. P ar is .
HELBING H . 1932. Uber e ine rn Ind a r ct os-Sch a de l aus dem P on t ien d er Insel Sam es .
n ebst einern Anhang : H y aen ar ctos sp . aus d em Pliocen v on Via lette (Ha uteL oire). Abh. Sch w ei z. P al ii ont. Ges., 52, 1-18 . B ase l.
HOFMANN A. 1887/88. Ub er ei nige Sau g ethierreste a us der Braunkohle von V oit sb erg
und Steier egg b et Wie s., S teiermar k . J b. k , k. geol. R ei ch san st. , 37, 207-2 18.
W ien.
LYDEKKER R. 1884. Indian Tertiary and P ost-Tertiar y Vertebra t a . P . VI : Sivalik
an d N arb ad a Ca rniv or a . Palaeont. I n d i ca, s er . 10, 2, 202-23 9. Cal cutta & L ondon .
MATTHEW W. D. 1929. Third con tribut ion to the Snake Cre ek f aun a . B ull. A m er .
Mus. Nat . H i st. , 56, 437-5 60. Ne w Y or k.
OWEN R . 1840/45. Odontogr aphy . Vol. 1- 2. Lo ndon.
P I L GRI M G . E. 1931. Cat alog ue of th e P ontian Carn iv or a of Europ e. B r it. Mu s. (Na t.
H ist.), 1-1 74. L ond on .
SCHLOSSER M. 1891. Die Affen , Lemur en , Ch iroptere n , I n se cti v or en , Marsupialier,
Cr eod onten u n d Car n iv or en d es europa .s che n Tert.ars und d er en Beziehungen
zu ih r en leb end en un d fo ssilen ausse r -europaisc he n Verw andten. T. 3. B ei tr .
P aliion t. Ost err.-Ung. u. Ori ents, 8, 1- 106. W ien.
TOBIEN H. 1952. I n dar ctos un d Ursavus (Carnivora, Marnm .) aus d en Unte r p li ozan en Din oth er ien d and en Rheinhes sens . N bl, Hes s. L andesam t . B otlen j. , 6,
7'-14. Wi esbaden .
VIRET J . 1939. Mon ogr aphie paleontologique d e la faun e de Ve r te b r es d es sa b le s de
Montpellicr. 3: Carn iv or a F issip ed ia . Tr av. L ab. Geo!. F ac. Sci ., 37, 7-26. Ly on .
WAGNER A. 1837. P a la on t ologische Abhandlu n gen . G el. Anz. Mitg l. k. ba y er . A kad .
W i ss., 5, 168, p . 320; 169, 327-328 ; 170, 334-33 5. Mti nchen.
JAN ST A CH
AGRIOTHERIUM INTERMEDIUM N . SP. Z PLIOCENSKIEJ BREKCJI KOSTNEJ
WYDOBYTEJ Z MIEJSCOWOSCI WEtZE W POLSCE
Streszez en ie
Autor op isuj e szc zatk i A griotherium Wa gn, ( =
z brekc j i kostne], wydobytej w m iejscowosci
W~i.e
Huaetuirctos Falc. &
Cautl.)
kolo Dzialoszyna (woj , loozkie) ,
porniedzy ktoryrni za chowaly sie stosu n k ow o dobrze pi erwsz y i drugi zab trzonowy
AGRIOTHERIUM INTERMEDIUM N. SP . FROM T HE PLIOCENE
15
gorn ej s zczek i oraz nieuszkodzony pierwszy zab trzonowy d olnej szczeki tego samego osobnika, dozwalajace okreslic z w szelka p ewnoscia rodzaj te go zw ierzecia ,
Autor zestawia wszystkie (24) opi sane do ty chczas ga tunk i q; p ierwotne j sy st erna tycznie grupy Hyaenarctos, ro zdz iel a n ej obecnie n a d w a r odzaje : Agri oth er i u11!
Wagn, i I n dar ctos Pilgr., pod ajac, wedl ug danych zacz erpnietych z prac ro znych
au tor6w, ro zrniary zebow P! - M2 g6rnej szczeki i M 1 doln ej oraz m iejs ca zn ale zienia
ty ch gatunk6w.
S zczegolowe por6wnan ie zebow trzonowych os obni ka znalezionego w Wez ach wykazal o, ze nalezy go za liczyc do rodza ju A gr io theri u m Wagn ., bo w iem brzeg do j ezy k owy korony p ierwszego zeba trzon ow ego g6rnej szczeki jest u
teg o oso bnika
kr6 tszy niz brze g zewnetrzny, gdy tymczasem u I ndar ctos P ilgr . p rz ekr6j p ods ta wy
korony tego zeba jest niemal k w ad r a to wy, a w d r ugirn zebie trzonowym M2 b rak
zupelnie talonu, ktor y u I n clar ct os jest n a ty m zebie m n iej lu b w ie cej w ydatnie w y ksztalcony.
N ieco trudn iej okres lic dokl a dnie gatunek A griotherium zn a lez ioriego w Wezach .
Dot ychczas z pli ocenu Eurazji p ozn ano cztery gatu n ki, kt6re n a p od sta w ie ks ztaltu
zebow trzono wy ch zaliczo no do rn dz aju Agri ot h er ium Wagn, Autor p or6wn uj e ksztal t
zebow okazu 'z W ~i:6w z ksztaltem zebow ty ch czterec h gatun k6w i doc ho dzi do przek ona ni a , ze p ornied zy tyrn i zeb ami zaz n a cza j a sie w yrazn e roznice, Ok az z W~z6 w
p od ty rn wzglede rn zaj m uje j akby posr edn ie s ta nowisko porniedzy A . insigne (Gerv.)
poznany m .z Fran cj i (Montpellier) a A . sivalense (Falc, & Cau tl .) z S iw a lik w In diach .
Dla zobrazowania rozni c wystepujacych w ksztalcie zebow trzo nowych po rn.edzy
osobnikiem z W ~z6 w a: innymi gatunka m i, autor zes ta w ia n a p l. II ksztalt zebow
czter ech p okrewnych g atu nk6w Agri otheri u m Wagn. wedlug r y cin , po danych w op isach tych gatunkow przez in nyc h autor 6w. A uto r n adaj e gatu nkow i z
Agriotherium i n te r medi um n. sp .
W~6w
n az w e
Obszar zamieszkaly p rz ez grupe zw ie r za t ob ejmow anych ws p oln a nazwa Hyac narctos byl bardzo rozlegl y , za jmowal b ow iern n iemal cal a p olu dni owa i srodkow a
Europe, Iacz yt sie dalej ku w schodow i z siedzibami ic h w AZlj i i rozprzes trz enial s i ~
szer ok o w Ameryce P 6ln ocne j. P ozw a la to przypuszczac, ze w pewnym okresie czasu
,
warunki i:yciowe na duz e] przestrzeni w Holarktyce, przynajmniej w pewnej sz erokosci geograflczne], byly dla tej grupy zwi erzat 0 He n ie [ednakow e, to bardzo d o
si eb ie zb lizone, i d aw al y poczqtek powstawaniu form geograficznych czy e k ologicznyoh m al o sle ad s iebie ro zniacyc h.
J ei:e li n a w et z tej wsp61ne j grupy w ylaczymy formy, obej mowane obecn ie nazwa
rodzajowa Agri otherium Wagn ., to wystepowanie A . insigne (Gerv.)
A.
we F rancj i,
intermedium w P olsce i A. sivalens e (Falc. & Cautl.) w p6lnocnych In d iach (Si-
w alik) wskazuje na istnienie w pewn ym okresie czasu n ici wiazace] fo rmy b ardzo
do sieb ie zbliione na dlugim szlaku od F r an cj i do Indii.
Ok res len ie czasu wystepowania grupy H y aen ar cto s napotyka na pewne t rudno -
sci, n ie zawsze bowie m zgodnie ustalany j est przez geolo gow wiek zI6i:, w ktorych
16
JAN S T ACH
znaj dowano szczatkt p r ze ds ta w icie li tej grupy i wa ha sie on w granicac h od gcrnego mi ocen u do gor neg o p lio cenu . Zlo :i:a, w k torych znalezio n o szcza tki Agrioth er i u m W a gn ., datowane s a z pliocenu; srodkowy lub gorny pliocen zdaje sie bye okre-
s em najw ieksz ego r ozwoju ga tu n k ow tego rodza ju. N a tom ias t szczatki
Indar cto s
P il gr. s poty k a n e sa czesciei w dolnym, a nizeli w srodkowyrn pliocenie, i I ndarct os
uwazan y j es t (G . E . P ilg r im . 1931; H. T ob ien, 1952 i in .) za ch a r a k te r y s ty czn y e le men t fauny h ip p aric n ow ej w po lu dniow ej E ur op ie i Azji.
P on iewaz drapiezne , pozna ne dotychczas dokladniej z brekcj i kostnej
W~:i:o w,
zb lizaj a si e do elernentow wymarlej faun y , zn a n ych ze zl6:i: M on tp ell ie r we F r ancj i,
a cz as powstania tych z16:i: przesuwany jest z dolnego d o srodkowego p lio cenu, p rz eto i szczatki Agrioth er ium in te r m edi m n n al e:i:aloby datowac jako pocho dzace ze srodkow ego pl io cenu .
Jak k olw ie k ga t unki rodzaju A gri ot he rimn Wagn . m orfologicznie Z'blizajq s le b a r dzo do ga tu n k 6w ro dzaju I n darct os P il gr , i prawdopo do bnie m ialy wspolnego p r zo dka, a takze n ie mal ten sam czas swego powstania i wygasniecia, to jednak ic h
1'0-
dow e linie b ieg na oddzielnie. P r zod k a ic:h szu kac n ale:i:y w m iocenie i kryje sie o n
w srod r zes zy drobn ych f orm r od za ju U r sav u s S ch loss. P omij a jqc r 6:i:nice wlelk osc i,
zeby trzonowe Ursavus Sc h lo ss. s a tak zb lizone k sz taltem do tyoh ze u I ndarct os
Pilgr., ze n iekto re g a tu n k i rod za ju Ursav u s byl y pocza tk owo oznaczane jako n alezace do r odzaju H y aen ar ctos (F a lc. & Cautl.) , np . U r sav u s br ev irh i nus H ofman n ,
1887, opisany przez M. Schlossera.w 1887 r. j a k o H y aen arct os minutus S chloss.,
a Ursavus depereti Schlosser, 1902, op isany przez
c u. D epe re t a w 1895 r. jak o
Hya e-
n ar ctos arctoidens D ep .
Os ob n iki rod za ju A grio t he rium d os iega ly znacznyc h ro zm iar6w cial a, g dyz czasz-
k a A. sivalense (Falc. & Cautl.) miala , wedl ug R . L yd ekke r a (1884) 482,6 mm dhrgosci , czy li zwierze wie lko sc ia zbliza lo sie do bardzo duzy ch ok az6w n iedzwiedzla [ aski-
niowego . Niewie le mn iejsza m us iala bye czaszka A . inte rme dium, g dyz -
bio rac pod
uwage stosu ne k d lugosc i zebow MI i M 2 w gor rie] s zczece u A. siva lense do dtugosc i
czasz k i te go gatunk u (57 mm : 482,6 mm) -
przy dlugos ci tyc h zebow u A. i n te r me -
d ium, kt6ra wy nosi 51 m m , dlugosc cz aszk i t ego gatunku wy nosilaby 431,4 mm.
Silnie r ozw iniete zeby tn a ce u w szystkich g a t unkow g rupy Hyaenarctos wskazujq , ze p r ze dstawiciele tej g r u py byly d r ap ie:i:nik ami
0
po tezn ych r ozm ia r ac h ciala .
Duz y zab przedtrzo nowy czwarty g6rnej szczeki, podobny ks zt a ltem do tegoz zeba
u h ieriy, ul a twial za pewne zwie r zeci u k ruszenie nawet duzyoh k os ci . Spos6b zycia
A. intermedium m o gl wiec bye podobny d o zy ci a h ien. Mo gl y t o bye zwierzeta s te-
pow, z wystepuj ac y m i gdz ie n iegd zie sk up ie nia mi
drzew i k r zewow, szcze go ln ie
w po b lizu zb iorn ik6w wo dy , kt6re z ca la pewnos cia rnusialy wys tepowac w pewnym
ok re sie czas u w okoli cy
W ~:i:6 w .
AGRlOTHERIUltf INTERMEDIUM N . SP . FROM THE P LIOCENE
17
OBJMiNmNIA DO ILUSTRACJI
P I. I
F ig. 1. Fragment p rawe j str ony g6 rnej szczeki z r esztka przedtrzonowego czwarte go i ze b a mi tr zon owymi p ierwszyrn i drugim.
F ig . 2. S iekacz tr ze ci z lewej s tro ny gorne] szczeki,
F ig . 3. Fragment doln ej szczeki z lewej strony Z resztk a czwar teg o zeb a przedtrzonowego,
F ig. 4. P ie r w szy t r zo n owy Z le wej strony do lnej szczeki, wid ziany od str ony d oj ~zy k o wej .
F ig. 5. Pier w szy zab trz on owy z prawej strony dol nej szc zeki : a od strony ze-
wnet rzne], b od s trony d ojezyko w e] ,
Wszystkie okazy wie lk osci na tu r alnej
P I. II
F ig. 1. S ze re g zebow go r ne] szczeki pt - M2 A gr i oth er i urn i n si gne (G erv.), w g
C h. Fricka (1926-30) ; niece zmniej szone.
F ig . 2. S zereg zebow gor ne] s zczek i p 4 - M2 A . sivalense (Falc. & Cautl.), wg
Ch . Fric'ka (t . c.) ; n ieco zmnieiszone.
F ig. 3. Szer eg ze bow gor ne] szczeki p4 - M2 A. p al aeindicum Lyd ., wg Ch , Frick a
(l. c.); n ieco zrnniejszon e .
F ig. 4. D r ug i za b trzonowy go r ne] szcze k i M2 A grio ther i um sp . wg H . He lb ing a
(1932); wi elk. nat.
F ig. 5. P ie r w szy zab trzonowy dolnej szc zeki 'M l I ndar ct os sp . z S a mo s, wg
H. H elb inga (I. c.); wielk. na t .
Fig. 6. Te n sam zab, w idziany z g6ry, wg H . H elb ing a (I. c.); wielk. nat.
F ig. 7. P ie r ws zy zqb trzonowy do ln ej szczeki M l A. insigne (Ge r v .) z Montpelller,
wi dziany z gor y , wg J . Vireta (1939); w ielk. n at.
5lH CTI\X
INTERM E D lUM N. SP . 113 lIJIHOUEH OBOr:I HOCT HOn
BPEKYHH H3BJIEY EHH Or:I B MECTHOCTH BEHlK E B IlOJI b lll E
.4 G RIOT H E R IU M
ABTOP OHHiCblBaeT !ppalI'MeHTbl 3y6oB Agriotherium Wagn. ( = Hyaenarctos F al c.
Ca utl.), 'H3iBJI€ ''I13HHbIX ill 3 n JllHOl.\ e HOllCH HO("I1HO H 6 p e.K 'IHiH II M ecllHOGT H Bemxe
;; Floasure IH on penenser HX JIlpHHU'AJIem:-!ocTb H HOBO:l>lY BHAY Agrio the r ium inte rmedium , HOTOpblH 3a HliJlWeT B c1'poeIllm HOp OHbl IRo p e HlHblx 3y60H, H oc06eHHocTH lV1 2,
!Wilt 6 bl npoaenryrosayio cacreaeraxecayro nO~H ~HIO IMelfW
A. i ns igne (Gerv .) .lI3
&
Mo a nen se (Mon tpelli e r )
(Siw a li k ) B HH~lIlli .
BO
CP,pa HI.\HH li A . si val ~n se (Fale. & Cautl.) 1113 OH8<·l.1lJl~ a
Agriotherium i ntermedium
Monnense
6 b l Jl
cocrasuoa
H ern reoJlOI'H'l00K1IHso3pacT 6 blJl
'I<l CTblO
clJa y H bl
6J1:Hi3ROH (paYJllc
iH 3
Tacrrnte sepoanao cpenne-nnnouenoau n.
0 6pa3 iltH3HH A . i ntermedium 6blJl He.pOfl11HO 6J1H30 X o op as y iHH3H lI Men, 'llO~06110
OC1'aJI b llbl M HpeACTaBHTeJlIIL\1 a ron npyrmst,
Acta P alae on t ol oglca P ol onica -
\"01. II/I
2
PI. I
Fig.
Fig.
F ig.
F ig .
F ig.
l. Fragment of th e righ t maxilla wi t h damage d P 4, M' and M2.
2. 13 of the left m axilla . .
3. Fragment of t he left mand ible with P , str ongly damage d .
4. Mi of !!he left mandibl e, lingual v iew.
5. M , of th e r igh t mandible; a labia l v iew, b ling u al vi ew.
A ll f·i gu r es of n a tur a l size
P I. II
F ig. 1.
F ig . 2.
Fig. 3.
Fig. 4.
Fi g. 5.
F ig. 6.
F ig. 7.
P~ _ M2
of Agriother iu m insigne (Gerv .). a ft e r Ch . F ric k (1926-30) ; somewha t
reduced.
J>4-M2 of A . sivalense (F a lc . & CautI.), after Ch . Frick (1. c.); somewhat r educed .
J>4-W of A. p a l aeindicu7Jl. Lyd., a fte r Ch . F r ick (1. c.): some w h a t r educe d .
M2 of the r ig ht maxilla of Agrioth er ium sp ., after H. Hetbing (1932); nat .
size.
.
MI of the mandible of l n d m' ctos sp . fr om S amos, after H. H elb in g (l. c.l ;
n at. size .
The same too th , top view, a fter H. Helbin g (1. c.): nat size.
Mt of mandible of A. insig ne (Gerv.) fr om Mon tpelli e r, top v iew, af te r
J . Viret (1939) ; n at. size .
.ACTA PALAEONTOLOGICA POLONICA, VOL. II
J . STACH, PL. I
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J . STACH, PL . I I
ACTA PALAEONTOLOGICA POLONICA, VO L . II
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