CATS OF THE WESTERN HEMISPHERE

Edllt'J ltv

VIRGINIA L. NAPLES

LARRY D. MARTIN

JOHN P. BABIARZ

THE OTHER SABER-TOOTHSSCI M ITAR -TOOT H CATS OF T HE W ES TERN H EM ISP H ER E

Edil(dIry

VIRGIN IA L. NAP LE S

LA RRY O. MAR TIN

JO H N P. BABIARZ

GmpJIics&lirar, H. TODD WHEELE R

THE JOHNS HOPKIN S UNIV ERSITY PRESS

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,«:on",.<'''''' oflne en",. ' "'m, 1(1)0'' ''''')' (d"'W'"8 "'.'yTOM"I

PR EFACE

W hen ..,menne "ys ".., I:>< ' -' OO1 h COl: ,he fi"" image ,hat u,ually rom•• tu

mind is ,h. fic",e-lookmg, ......ll·k nown genus S,"iloJo~. from ,h, to< _p'

of R.ncho La IIr• •. It i. ,he purpose "frni, boo k <0 bro.Jen <Al' urKk....'.ndingbeyond ,II.. genu. '0 indude • rar< g roup of ••Nrc.... which ar< nOl only contem·ror.ry w;,h S",iWdo" but had. u n ique appe.rance . nd ""h.. io•. Utili! ,he 1"'''rouple ofde.<de., our knowledge of",imi"'-'OO1h c.". ,he O1h....b<r-'OOlh., .......'<elY hmiltd, Ind.ed, ,he genu. Xc"",""l., ",as only ,«enlly d,scm,.",d. Vinu. tlycompl. .. skeleton. of ,he.., an im. lo. derailed in ,h i, book. Jem""",..,. how in«,­."ing.nd 'p<<<.<ul., ,he scimitars WeTC, w••rt able ,,, ",,,.1 •• much .bout ,hei,.n""my, appe.r.ne<, and b<h.viol .. i. known ahom S"'IOO"". induding ,h, his_tory of,hei, di•••", •.

Ou< book was in,p;r<d by 'he 19J1 d ,,,;<, Th, frhJol< .fR"","" La B,<a. by Mer ·riom ond Stock, We provide romp""ble iIIus, ... ion, of skeloul an. 1Omy by ."i"Ma<yT.nn". Life·lik< ,...,omion. w.,.. doneby pal.""ni" M.rk Hallott . The text

"' .. mostly written by the thrtt principle autho..., Larry D, Martin, Virg inia L.N.ple., andJohn P. Babia,.. . l'ou<o<h...OIOO... ront<ibOl, to the <h.p'..... bringingaddition.l h,..ad,h .nd <ompl.te""" to the .ubject, Jonena M, Heo"t di>covered a

...m.rkoble . keleton ofH..."'~m"," in ,he PUocene of Idoho and included 'he Ii",deSCription ofthat 'peei men in h..Ph.D. d i"ert arion. H . Todd W fie<,1<r. 0 mechan i­

<01 .ngin••r. "'.. "",rui,.d by Lorry Mortin ond Virginio Nopb in 19% to build amecnonieal device '0 ,." the v.rious hypothetic.l bi,. models of ,h••aber·tomhS..iIOO"". I'our yea.. I..... thi.de'i~.<.lIed '" Robo<.,; "'.. filmc-J by ,lie BBCand

f...ured in ,lie Di.covery Chonnel ..,rie" Bruce M, Rothschild. a procheing rheu·motolog;', and a le.d mg figu,.. in the <rudy ofpal.opatholugy.•umma"'" ,h.di..,a,., and ... ,ultan' pa'hologie. found in fo-~,;] ..be ...." . !'tter E. Kond..>hov. a

p.leonlOlogi" form.rly a..aehed to ,he Russi.n Ac.demy of Science. in Moscowand an expert on the Cenozoic faun . ofCcn'rol A,ia. describe, a lIo....I,,'i....l ulldi,cove,..d .Iong the rout. t.ken by ,h... ca" wh.n ,hey .ettl.d in ;'oJoc,h

Amene• .

Xll . ... Ae.

SmilO<!<mfo '~li, d.lend;ng ;" , ill, I B" o" coif. I",,,, I "'001D'" wol..,_ Id""iog M", HI li.,,)

The ;nni.l eh.ptor eon.. in. on inrrOOll<t;on ro ,.be"." .with d;scu,,;on of ,h. ,h..e pr;ncip.l mo, phm y!""" dn,, ­roo, h. ",im;"<-Io<"h.•nd cookie-cu tter. WCofTcr, modd'" expl.in how ,he", c.ts mOj' h... pa" n ioned th . ;r"", imn_men', incl udtn~ diff.ren«s in rheir mod., of prey cap'u«• nd k;Ihng beh.viol-.•nd how , beiTOTi~; ns • nd..01 ution..ytrend, or< rel"ed ' 0 ,h. bro.d 'I"'"",m of dim." ch.ngedm;ng rho p' " seven million )'C'r'S, Induded or< , ideb. r'S o nd.,ing lhe f" " il record .nd a br;efb; >tory of ' ."",omy

The ,."""d eh.pler 1001<>",he ,ober,r",,,h h;t< f,.om , he' -,ewpoin' of'imple meeh . n i", We descr ibe lhe working.of I meeh.nicol h;1< , imul.,o, ,hat was te"ed on ani m,1cad.,'e r'S , Th.,e oxperi men" illu""," lhel; m',.. ~,n.of ,hedirk·roorh and ",im itar_,"'"h <comorph. ;n p« y1.,tli ng. • ndwhy ,helwOm0'l'h01ypr, m lY h.ve emh,.d

The ,hlrd chapter" a ,u"'.yofthe "iden« forp•.,hologyon , he ,kel...1rem" n.of ",imilar -'OUIn catS.nd a rompari .

son with S.. ilodoll_ On« ' )I;.in. w" h" e in 'i)l;h, imo ,heI><h" ior oflong-e"in« p....d.tors. includ;ng lhe d,sco ,'eryof <viden« fo,- , ube rcu loSlS in , he "" m iur·too,h HomOlk

".m"".. , Thi, di,., a.. wos prob.bly "ni'O". 1in ,he :-JonhAmerican bison I" 'pu l" ion. which , ug;."'''' th .. b,son we",• prey i'em fun h.. 'pre;."

Chapter fum pro" id., ,he firsl JeuileJ Jescrip1ion uf lhe, Icele..1• I\,,"myuf ,he cook'Nu,, '"cat XC""''"il"J hodJo.a, _

The ,hott-l'l!.Iled, I><ar·like propot'1 ion. of X'""'m,I....ecom pared '0 ,he Ion!:, ,Iendc<, .1m"'t ch....h-like I, g, of aPI"""ne 1I0..o,h",".. , .how In)l; how di(f."",1Y,h< so rel. "dform, J..h w nlt problem, of cap,uring lar!:, prey_ Thi.chapter. along wi, h chop", , i•. e".hh.no, a h",i, for com­I"" ison, w'ilh olher pred.,ors_

Chop'er Itve explo,m ,he rel"i"",h;p of ,he muscul.,u ....to the ,1.el,wI\ .nJ how' th i, <an be ""J to ", >tore th e ' p'!'<.ran<e of eXlinct . n im. l, and ~ain in'i!:ht into the ir be­h"iur_ The >nato mi,·. 1"'U<lU" of x",.,,"il., i. d.,,,,ihcdand related 10 ,he biomech.nical require ments fur ,he life·

>1yle of. robu"ly buill, ,hOl1·limb<d .mbush pr«l.wr, it i,then com p'r<d wIth the .n..omy of5,.iI00"".

Chapt« ' i' d<",ribos in d<..il one of the mos, compl..., kel.ton, of lIo,"o'/'m"," tK/'y..." . Th. prop<>Ttion, of thet"orth Americ.n Pliocene sdmit.NOOIh c.t.r< comp' r<dto tho~ of the "out. ,ho rt·legg. d cat. Xt o", ,.il,,, /'oo' ooat.Tb " cbap'er . 1>0 indud" ••i<kb. ron the di><overy .nd , ite" ..tig"phy of110"",/',"" ," i""y"u from Id. ho (110. 6.l~

Ch'pter .."n i, . ,umm..y of the t..onomy of Ifoml>­

,nm",. in NOl1h Americ., Itcondude' tlu t the Nonh Am<ri­can Pliocene ..mple con, i", of two lo ng.legged ' p<<i" . II ,io<Ily..." • nd II. <lC"'"poll. Indud.d i' the ' ku 11 dtKriplion ofII. c",,,,joo" from Ddmont , South D. kota, These .peci..can he "p<l ..t~ from •.,h OIher on .i.e .nd ct.ni.1 ch..•. cten . wi' h II , muapo'i boing ,m.ller, .nd m<l..e hke theI.ter 'pecks II. ""' ,",

TIl< eighth chapl.r d<scribo, • hOmOlh... , kull from thePliocen. ofceo".1 A,i• . Thi ' ..bore.. ..mpb the Beringi.n11""",,11.,,"'" to.. found .long the high I.." ude p<ltbw.ycon·ne<'ing the •.,h.ng. betw••n Eur..i• •nd NOl1h Americ.Wh"" <OmporeJ with the contempor.ory r-,'on h Americ.n<pecie' . it ..em, ..oIution.rily preeodou '. , ugge>ting ' north·em <enter ti>r ;><l..n«ment in the H"",,,,nm"," hn••g.,

.....<0 . " ,

Ch' pI. r nine pull. together ' he common thread. ofho·mo! here evolution ;nto an unde", .ndable po,tem ofdim.. icclung• • nd di"ributional ..·. 01•. Th. reCOTd of ..imitar­' ooth <au in South Americ. i' .1>0 discussed,

Th e . ... ngement of ,he.. ch' p'e" progre ' ... from .g.n...1diKu"ion of ..imitar·,ooth ..bert.., in 'ht Plio·Plei"ocene, indud mg how we determi ne 'heir 'pp<.rance. nd boh.v;or. '0 ' det. iled d<Kription of the . n..omy ofhom Olh.re" Thei r 'p<ci« Jiv...i' y .nd b;ogcog..phic di,·t ribut ion i, rd..ed to theiTw,y ofhf••ndth e overall imprin tof .lim.tic chang• . Their ' uce.., and uh im... extinctionmU>1 , in >Om. W'y, ,,1... '0 tb". f. cto...

A, in . 11 ba,ic scienee, tr u,h i..lu>lve. Som. of wh at we

h." 10'.y i. 'p<cu I..ion, ' upported by b;"ofe, iden« fromdi,.....nd of'm unrelated >ourr... A, such , , h. "udy ofscimit..·tOOl h c.1. i' incomplete. W Ith ' hat cave.t. we now.ubmitourcor.du,ion. f",your re'kw.W. beli." ,hi. bookconrribut., to the und."tonding of ' he rep<. ..d cyd. ofevolution . nd , ubsecju.nt ext inction of..boN ooth ..... We. 1", hop< 'hat it will in,pi... more in""igation, in ,o th ...f.sci natin g am mal•.

ACKNOW LEDGMENTS

W .woold like to ' hank ' he followi ng for . " j".o« . nd p<rstverane<: MuyTanner, who p r<:pu e<! ,h••h.ded illm".tion, .nd graph\<:> for x,""''''".,

~oJ","",. 110.....""... i",~yr"" II. ,.,.,.UlJi'.h•• nd H. ",nat";,"" M.rl< Hallett , whoprovid«t.n f,.,-,hc l>ookjockn and . dditional art work;Jerry Surk, who took manyofth . photograph" Don Hurlbutt. who prov ided tl>< d igita l im age of[he X",,,,,,,iI .,hd""'.ecr.ni.l mus.;le recon,,'u«ion; II. TOOd Wheeler, who did <ompu'tr gr. ph­i« and im'g<" edi'ing ; 1', ,,10 M. Ott. who d 'd ,h. f'O,"",mial reconstr uction of x,Iwd",••e; Bill Ak<"'.n. who g.~m.d the flr" t.".ble mooel of •• l>c'-'OOIh hio­me<hanic" D.vid Kronen. who did ,h. mold ing and <..'i ng: Mik< K".,h, whomgln....d,h. X, .oJ,".,,,I«I,,.1mount;th. lit. Phill ip V. Well•. who ge nerou sly

provid<d ,h. Norlh America" L... P1ei'lO"e". vegem i"" d.....nd Mary Brook.fOf the rolo reJ m.p; O"d "",h<r, for rriti,,1 rcading of the manu ocript ond forproparing ubi.., imag.., and the g1o>.. ry: Tia Eaton. for t"" hnical ",vicw. wordprr><:e"ing. •nd form.tting of.ll <hapte.. . nd tab"".

F<>r u>fful Comment>, d ,>cu""o n,, photog"pby. imag<>, acee" to co1].,.;'i"",.and .."ing We wou ld .1.., like to ,h.nk: Mauricio ",ntOn; Kenny Bader: lIemiclIaner; II. 1I. 1Iesio; 110" Barnttt 'Japh<th Buyct: David Burnham , Matt Chri" ophe"G. Como" Sbollcy Cox: [)(mgla, Dut: Elizabeth Ebert ;Burkhart Enga, .." A. F.l k:A\Shng F.rrell M. Frank: Ted ). Frtmd; Henry G.Ii'"":Joh,, M. H.rn.; " eto .."Hooijdonk: G<>rdon Hubbell: R. C. HulbertJr" Cri, Ja", Gr<>rgc: T,Jdfo""", ChfTJ..emiab: tbe la.. Bjorn Kurton: W L.ng"on; W. nn Lon"on Jr.: Wil", v.nlogchrm: E. Lundeliu>: IIru« MocF>dden: II. McC. rty; loi , McKonn " Cuy M,,·wick:T.J. Meeh.n, Di<:k Mol; Lyndon Murray: F"ndoco Pttvo" i: "' ,conio Rincon;HoIhston L. Riv,..., TimOlhy Row<: 'he I. .. C. B. Schultz: Kevin Se-ymour; Chri.Sh. w: Ron S"blr r: Ce"'!!e S"venson; Craig Sundell; " em Sunddl; II, Tedford ;BI. i« Von V.lk¢nburgh: M, Voohi.., D. Wobb, Lon Wrr drl in: P. Whi stle".ndMIke Wiggcr, who a"i"ed the study in a v"i<ty of way'

We ' ppr<cia.. . nd th .nk the folloWing in"iIUtion. for .llowing us .«t" to' 01"

'VI AClNOWLlDGN lNH

loon' from. 'heir =1I«'io"':.~m",ic.n Mus<um Df N.'u",1Hi.,ory; Bone Clo"",. 1m,; FI',riJa P.le"",o1ogi<>l 5o<ie'y;Id.ho Mu><um of N.'u",1 Hi"ory; John D.y fos'il B.d,N"ion,l Monu ment; !AI. Angel« Coumy. Goorg< C. P.geMu~um : M..k 1l. 11e" 1'. 100.,-t; Mu..um ofN.'u,..1 Hi.·lOry, Lyon: N.,uc.ll li'lOry Mus<:um . L""don : " .'umi",,·risch.. !>1u,." m, B.,.I; Ru"i.n Ac.demy "f 5ci.nc...Moscow, School ofOentimy. Oregon H.,I,h . nd i>cienco.Univeni'y ; 5ilve, Riv", 5p, inl/:' MUkUm ; T.." M.m",;. lMu..um; Unilrd S,.,.. N.,ion.l Mu..um. Smith",n ian

In"i,u,iun; Univeni,y of FI"rida Mu><um uf N.,u..1 Iii,·tory: Uni...."ity of K. n, as ~lu,.um of ".tu"l Hi"01'y.Vert.br.r Palro Lab; UniversityofNeb"'h Sr... Mu~um:

.nd a ,prei.1 mem ion to BIOI'SI ti>r ~n.n<;. l 'upport,We rsprci. lly th . nk Drb L. Hod'on, Joan M...m. and

Vicki L. Wheele< for their forbe..a""e throughout thi'rndca,,)c,

T"" Robo.., .,n, ", bow ,.., ~.,",. (lop). ..." ofS"'_lAC ' 2001-249.,.11 ,nd ""nd,ble w,,~ •compo,• • """"'....•••I II,""'.tI< G""'P' P' p "'v.."'" (~". "'). leo.'10>1v;,.,"' ,.,>

2H . TODD WH EEl e R

Experimental Paleontologyof the Scimitar-tooth andDirk-tooth Killing Bites

CATS ,~ND CAT-LIKE cunwo,..n• • ,..,heultlmatehypercarnivore•. •nd

,;1>« Martin (1980) d ....ifi.d 'hem in,., differem <eumorphic group•• inde­penden, of phylogeny. ,hei, fun«iona! morphology h., b<.n defined by ,h.

>lup< of'hoi r c. nin... Each ofthe ,h",. b.a,i<; type' ofupper can in< morphologies­. on i" l·toot h , K lm i.. r·'OOIh• • nd dirk-'ooth-are .,wei.,.d with un;que killingbi, • •. The ..bcNO<lIh ,,1> (boI h fdid ,,.,, ond ,at-Ilk< mammal. ) or. aptly named.

In .ddIlion to ,he visu al "mil. 'it yb<twun 'h<ir cur~d uPl'<' " nine . (, . bers) . ndthe "adi,ion.1 , ..airy w••pon. ,he y shar• • \>;0,;.: f"n<lional ,imil.,;,}'. 'he -drow­cut, · As.n ."emi.t !"''' of ,h...be,_tooth killing bit•. ,h i. >. t ;on is .,soci. ..dwith 'h, <"" oJ 're mthe ,.be, bl>Je,in which ,h. blad. mo....m.n' ",ralle]10,h.long u i. r"ili,.«, • Miring movement rrrp<ndieul., to 'he bl.d. (Bu rto n. 1884).Mach.irodon' (f, lId or ·'tr u. c., ' saher ·'ooth) upp<r conines .... nOt ,h.rp enough

'0 cu' .nim.1 hide or other con nerr t" . nd mu:<cul., ,i"ue, unk" 'h.... i. OOme."""iared ,I;':ing movemen' .Iong the 'ooth axi•. Both c.ni". morphologk....defined by how ,hi. i...hieved.

$<imiur·'nnrh e' " bit into their pre y " 'it h .hort, broad. «,mod uWn ra n in<s.I' i••ugg<sted ,h., ,hey used an inAerr.d .re tr aj<.:,ory. not unlike ,h., of con;.:.I ·

'ooth cat< and other c..nim.... Once 'he rrerh "up "'oving, ,hey "'ust rhanged;rerrion .nd .ithe, be "'it hd" w n by orrning the mou,h , ....' an' cats do. or bepul lod frr<o rrrprndiruiu to ,h. too,h ..i., •• piranhas do.

Dirk·'ooth cat<.with longer.nd r.l.tively .Iender, finely «"..ed uppert.nine.,

,pP'..n,l y used another "'ethod to rut 'hrough hide and ,i"u•. They cuvod,hrough p..y in. con,inuou. are . kerping ,h. incisor> free • • th.y CUI lhei ' w.y

OUI . Foree w' ''Wlied rrrp<ndirul.rtothe 'ooth axi. a. ,h. bit< prog",.srd, pulling,he tooth into a .liring." posterior to ,he tooth ax i•. Thi. ptodu.... wound«vn. ltim.. ,h. wid'h of'h' 'oo'h. and the rr' j«tory re,,,I,, in ,he upper con ines cutt ing

lhei ' w.y out of ,he ptey "'i'hou' .n~ nred '0 open 'he mou'h. Thi. bile becomesa Auid. even mov",,.n', wi'hou' app'",n' ch.ng. in ,he direction o{'ooth or h. ad

mo"men'Ku"en (1963) fir>t appliod the '"m' scimitor-'OOIh .nd dirk-'ooth when dlS,in ·

gui.hi ng be, wern ,he M.ch.irorlon' Trihe. Homo<heri mi. nd Smi lodon'ini.M."in(19!!O) " I"nde d ,h. u« of ,he "imitar· . nd dirk·toolh term, '0.11 vari<ti.. of car·hke •• her-'ooth e",mo ' ph•. n.ing 'hese de.ig".. i"". 10 eh.racreri...nim.l. th.l.hore n ani.1 .nd postrranial a, well •• den..1katur... Prim ro the idrn,ific.,inn

10 THO OTHU " OU -TOO TH'

of XttW.!mil", ~oJ,ona, (M.n ," ~" l.. lOOO), >eim" . ...oothfo<m. '0" '" ",elu,;v. ly ••"",i01<d wi,h • mo« ,ursori.lpOSt, ,,ni. 1m"'l'hology •• ,omp."d '0 Ih. shortt<-kgg<ddi,-k-""nh ~mTl'_ Hnw.v.r, in ,hi. ,h.ptee I . dd",,, oolycr.ni.1 . nd d<ntal morphol ogic••nd ,h, killing bite mod<l,, hey f",ili, .t< . .., , h.. ,Il< t<rm•• '" u«d ",du.i""ly in ,h.".nio-d<n..I<""""_From ,hi. p<np<et iv•. ,h, di" inet ioobe' w«n $Cimita r- . nd dirk-loo,h i, b. ..d only on ,II< 'yp<ofkllHng bit< , h. m"'l'hology , upport._In . ddilion m ,heuPP<' <onin. propo!'tjon, .nd distinc,ive ,h. ",cte,.. on th,cutt ing bl.d. m"gin•. ,h, . h.p< of ,he «mento·.n. m.ljunction (CEJ ) i • • k. y di. gn,,,,ic fe..ure in ,hi. an.ly.i.(Rivie" . od Wh. eI.., lOOn

THE CONICAl·TOOT HS

A11 "t. n' pantherinc"ts."'00 ie.l·'oo,h" with oval uPP<'<onines ,hort.. ,h.n tho" of the ..beNOOIh.l)'pical Crownh.igh, '0 . nterio r-posterio r di. met" (Af'D) , .,;o. ;n pan ·therin« . '" . rou nd 2.'01 . •nd (APO) '0 1.bi.l-lingu.l di. m­e", (LLD) are .round I.l1 (V. n V.lk,nbulgh . nd Ruff.1987), The.. values , on lDrm '0',11 '0 ",,,nt mod.l. of "I',i­mu m 'oo,h .'pect ",tio (F" em. n . nd I.emen, l007. , Th,uppoe cani ne••'" ..par..ed by a di.",m. from. row of, m.U, un..' '''ted, som,what . patulat<incison_ E". nl pan_,lI<,ine cats u.. one of ,wo killing bil", dcp<nding on , herel. ,iv, ,ize of,h, p" y .nd ,he skill.nd <>p<'i.",e of lhep~'o, {McDoug.lI977: $unqu i" . nd Sunquist,lOOl), POT• qui.k kill. if til< c.. i. confid,nt of being ,uc",;s[ul. • hit<'o,h, n. pe of,he ncek, with ,h, intent of"v"ing ,he .pin.,will be u«d. If'he ,01 i. unce!'t.in ,h., i, e. n do ,hi. 'ucce,,­fully, ,II< def.ul, killing bite will be • ,hroa' 0' no.. clamp.wi, h , h. c., holding on'o ,h. 'hroat 0' munle. "phy.i"ing,he prey o'er. p<Mod of ..'eral minut<.,

THE OIRK·TOOTHS

The di tl<-'ooth. ' re ,h. most .pectacular ..hr,.'ooth•. withupp<' <on;n. crown h. ;gh" « aching 180 mm . Dirk-loothmach. irodonts have .Iong..... hrn .pproaching . n ApDto Crown h.igh, ra'io of .:1. wi, h 'ypk.1 ApD '0 LLD of l :1(fig. 2.1 ). Th. upp<rc.n;nes h."" • •h.", m..gio (by d.ntal" .nd.rd•.•ppro.im..ely O,l-O,J mm radiu.) on both ,h.posterior and inci..1h, lfof tho .nt« io, m"gln. f",~u<ntly

with fin<..,.,.arion. (. I' hough ",.,.arjon, m.y be 10" urly iolife ,hrough w. ar). Th. inci..,,.. .re l. cgrr .nd mo", robustwh.n compared '0 conical ,,oot h " , nd. rd. (Biko<viciu, .ndV.n V.lkenbu'gh, 19%). The ditl<-,oo,h, .1.., h.v• • """.­who, proc-umbam iocisor row. "'P""t<d from ,h. uppe'canin..by . prunoun«ddi.st.m• . In ,onical,'oolh <0" .,II<uppcr .nd 10"''' ;ociso" d;"",ly occlude I' on I,. I' on I,. •nd.., on, In 5..,loJ"", ,h. arch m.y . pproach ,h. str. igh' lin. of,h.conic. l·,oot h•. bu' ,h. incison im«ioc k(AkeN,.n. 19S' ),

Th. I ' . ~ , hr'w«n ,h. I", so ,h. t ,h. incisors foem. zigz. gpalt<rn hr,w«n ,he upp<" .nd lo",ers, fittiog hr , ,,,,,n <a,hO\her",.igh, up .od down .nd in . ",.igh, row. Th<y .'"post,OOrly fCCun.d, with much larg" roots ,h. n . conk.l·'oo,h. Aken'en (198') "''' convinced ,h.t ,hi. impli.d •function.1 rok in ,he killing b" • . An . It" n, ' ,v, <> pl. n01ioni. th., ,h. incisonof. " hrr-l"",h muSt..'ve . 11of ,he pull­ing. 'ugging. g"" ping. •e«.s>ory functioo. in prry c.ptu...f..ding , cub tr. n. port.•nd .., on, ,har .'" norm.lly ,up'porteJ by ,h. can;n" of . >t.m "1S_

Typically,,II< d<, iduou. uppe, canin.. . pproach >eim i,. r­,h.pe. wi,h • crown hdgh, . bou' h.lf 'hat of'he "cond.ry(p<, m. ncn')upp<rc. n;ne_ M " h.;rodon, d.o,. ry-m..il1.ryg'p<, .t< .bou, 90° (Ake,..ten, 19SJ, ..qu iring . shorr.nedbit. , .diu. with .ubstanh. l . kull .d'p,at ion, . The 10'0'«c.nm. i. dram"ic. l1y ",du«J. particul.rly in 'Ol' wi,hlonger uppor canine, 'uch a, 5..iloJo",

Th. highly d<,iv.d nimravid,. /iv''''I", lI~plorJt"""',

(Eo, ..il",) "",rio•••nd also Ba"","",ji1i>, approach . n ApD'0 LLD r.tio of 4:1 . nd m.y ><hi."" . g. pe of 120 '_ Gape...ceeding 9O" .n:-><sociated wi,h hymicomorphi< f.atu ....(N. pl...nd M. n in. lOOO). Th. ..tan' cat with ,h. long. "uppc r c.nin.. in proportion '0 sku ll ,iz. i. N,oI'Ii' n<"b. losa(d oud. d I.opard), which h.. . ,rown lI<igh, '0 APD ratio of),28'1. I' h•• been argu.d ,h" Nwftli, ;, more " milar to , h.m. ch"rodonts , h. n .'" . ny of 'he o, her e...nl "big'"<ors(Ch, ;" i. ,,« n. loo6; lOOS). B.ca u« ,h. low. e c. nin.. ofN,cfrl" .re . 1.., proport ion.lly long". ,he dental morphol.ogy i. mot< like • •m.ll cat punc,u« 0' . kewer bit<.nd ,h•onlilhe.i, of ,II< culting func,ion of , .hre-'ooth uppe ' c.­nines. Th. strung crani.l ,imila'lties of N'~frli, .nd 'hemach. irodom • •" g.pe ad. pt"ion. (Sl" ero od V.n v. lk.n ­bu,-gh, l00 8),

A distinctivr ch.,act<,i"ic of di,-k_'ooth mach.irodon,sahrn i••n invrr,ed bell-.h.ped CEJ.Start ing .. ,h. ant<no,m•.-gin of ,h. uppe' c.nio., ,h. CEJ on both , h. labi. 1.ndlingu . l , id" cu"',, ' ow. rd 'he ,rown ID, .diSl. n,·e .bou,<qu.1lO,he ApD of,he 'OOIh. A' .bou, ,h. midpoint i, cu",...pic.lly ('ow.rd ,h. roo1) . nd "'urn. '0, h. post<'iOT mar ·gin.•' . poin' . few millime,.n .w.y from ,he . 1"..,lar hon.on. fully « Upl<d upp<r cooin. (Rivi"••nd Wh..I".looJ).The ling ual , id. of ,h. uppe' can in. CEJ of nimravid. i,,;milar. bu' nor n. « ssarily ,he l.bi.l.

In,.,,,,,ingly, ,he mOT< d<riv.d nim••vid, ond /la""'"",­frli, (Bryant. 19,5 I h. ,.. d.d dunu...hr n .imilar in profileto , h. perm.nen' upper <oninc, .nd ,h" .1.., approach , h.mio crown h.i ght. P,ior to ,h. eruptIon of the pe,m.n. n' .. .hr,.. . , ubst . n,i.1 W<a' on ,he 'eSt of the perm.nen, d<n,i,iooh., . I" . dy occu.....d (Bry. m, 1988), Thi, d.lay.d «up'ioncould be beo.ficial. Shou ld • •uhadult c., b..ak • d.dduou.sahr r in' h. cou.... of I••m;ng ,II< killiog bi'e t<chnique. ,II<cat will ge' • f".h " .rt wi,h ,II< "uption of ,he perm.n.n'cao",..,

f 'IU" 2.1. Tn. f',o!i l. of. Smil<xicn UP~' CO",", '~""" ng

<~..",.,''',' I.,,",,, oIt~. eEl, l ACMHC 1000 R·~ l :

(;<o'g. C. P.g. My",um <01'-<,,,,,,. (pnoto A,. I'" I F..",II,, ,," ' " ' ' Tn. G<o'g. C. P. g. "'u'. um)

ROBOCAT AND DIRK-TOOTH BITE MODELS

Not ,urpri'i" gly. d 'rk· ,oo< ~ 'Ol' ~.V( alw. y. r«<i,..,d 'h,most ' ",n,;on from re«"eh,rs . nd th, public .111<0 .T~.jr,h"i.m..i, n"u", "",wi,~,..nding. , ... f", ..m.'n. ,~..the . bs.nc. of .ny living proxy P"<c!ud.. r"h"i, in.igh, emhow'''' de-n'i'ion fun"iOl1ed from di..ct ob«rv.. ion_Thish.. led '0 ""m,h. ,heori .,. "'''''' di. pUled . more of..ncomrove..i. l. 1:><11 . 111.",ely ,he<>mic.1 ., '0 , ... mcthod OTme,hod. of d;.p"ching p"'y,

The.. i••n ,,"n, m.mm.l ,,'i,~ "...,~. ' h" ,orne du«in e"ern.1 'pp",..",e ro 'he dirk"OOlh;, h. Chin,« w. ' erdur lfyJtop," " i",,,.i, (fig, 2,2A). The« ....h are betterde-""bed as 'usks. because t~ey ~..'e no "I'I""'ing lower... ,h . nd O<Ily ...,,;gi.l en.mel 'o.>'i"g, Tbcy .",only"..,11de- ..\opeJ in ,he m.ks . nd serve "" ",.fuJ function '1"'"from m..ing ,nd di'pl,y. T h '"'~.... used wi,h a <10>«1mout h ,o J" . " .. ,'" w;' of"""'r m.1<o during"ruggk.to ....b];sh do min.",e, Th. t u,~. h. v• • very ,hort roo' ,

wi'h .n n,,,,,,,,,ly ,hic k periodontaJ lig.m.m (fig. 2.18)• • l­low ing ' hem '0 fo ld b,, ~ in 'hri' ""k... (fig , 1,lC) . nd not..",ict "OTm. l beh.. ior . uch as g ru ing. 8<" use ,hey pro.vide • modern . n. logue of wh.. . brerdi"g di,pl.y ..ber i.li1<o , ' be« ,u,ks refu.. 'he hYJ>U'hcsi. ,h....be r·tooth , at, .be" h.d • primary bre.ding displ.y pu rpose, . , ,hey h..elow «xu. l dimorph ism (V. n V.I Lenb"",h.nd So,co, 200l) .•nd , ... m.,,;ve l'OOI' demunstr. .. ,h.. ' hey .pply ,ubstan ·,i.l furre in biti ng

Dirk,'O(>l h' h...been ,h. focu' ofmos' p,coced ing ~iIling

J:>;.."uJi.. (M. " h. w. 1901 , Bohlin. 1940.1947;Simpwi' ,1941'Kurt "n. 1952, M. rtin. 1980, A.eme". 19.1, Bry. n' , 1996),ALe....n b.aseJ hi. m<><lel "" • , pedfi, 'p«imon (LACMHC1001·1. G«>rg< C. p.g. Museum ,oIl. ction from Roneh" L.Bre., RLB1. con'ideri ng i' '" be ,ypie.1 of S..iledo", .nd"ipul>teJ m...u" Ne pa,.. m<1' ..,,h".by c",ating . testablehypmh..i. 'ha' could be ,'eri fieJ.,perimen.. lly, The initialfocu, ofthi' rese.",h w., '0 "" ,he Ako,,,on killing b"em<><le l by replicating i' . nd com p.a ring it ro gene..l;zed'hroa' killmg bi,., u'ing • m«h.n;e.l de'i"" (Wllul<T.2000, fig. 2.l )

The pract ice of phy, ic.l <xpe. im. nution i' more 'i 'iblein ot h.r field, , u,h . , . reh. eolog y (Cole,. 1973).wi,h. pub­Ii,hed e...gory of "' omb.a ' u,h..ology" involving ,.", of

repl;c' "..,'porI"'""'MU' replic..ed .rmor or pro, ie. (Mul­loy.2008).The.. h"'e bee" rome ."ell.n' qu.n'i' ative l.bo·n ",.y "udi.. in the paleon tu logy fi.ld yklding Tyra."o"'.rJ:>;, r force ( Eric~",n.' al.. 1996) .nd .ffe" of «rr.,i"".;.e,shope.•nd ,h."P"'''"" 'u, ,;ng ,J:>; Ii'y in prey pro, ymateri. l(Able•. 1992),Abler constructed a fixture c.p.able of.pplyingme••ured force perpendieul.r '0 . te" bl. d. , whi le applying.n ind.pend.nt me. ,ured f""e p.a"Jlelro , ... Node. adding' he d,.w_eUl '" ,he oth.. qu.n,;..,i" ,,,;.blco. The inten'

>.>. T H O OT H " !A ..~-TOOTH'

ri~".. 2,2, A ,,,,,o i,,,<noft~< C~ in... , " .... d<<<, H~"'""",

i"'''Hi!: A, Th<,",., in ,i>< fo'w" d comb.a.~i' iM. B. Tne," , ., in .ne nin, <od b.a". ,,,,io, ~i'ion. C (oppO$i"). T~e,~ ;,k .ho,' "",,""on,. I I;,.m<n' "",,,, it>oin8<·I;", m~"'<m.nd i, , ho" n by> ro.", ..pli... ofto... ",< dim<n, ion, .. '~e

'''''' ,"'." on .~e 'i8~' .",t (<<'"rI.., Vdi Wheele,)

of my de.;gn w•• prim.rily quali", ;v., <0 ....bli.h which..~" .nd ><quenc< of movement> wcr, wo,kabl, . nd..lll<~ wcr, not. Th, cap.abili.y.o m,..u"" [h. m.gn;,ud..00 d;""e'ion Qffot-<:.. w•• a me.n. of ......;ng [h. ""lal;".ff«<iveJ'l<» of, he vuiou. propos.dbi.. modet, in .h<exi,,·ingli.....'u"". My...umI"Km w.,,h .[ .n ex;"ingbite modelwould "",r\:,.nd ,h. """ would not provo fe..;bl• .

Thi, m«h.nie.l ..be'-[OOIh eo[ .equiKJ .he nickn.m.

'" Roboc•• ,· I. u>«l both " ..I.nd resin '""' .. in"rch,ng.'. bl< SmilDJ"" ..<th (LACMHC Z(I<)1 ·249j from RLB . Th;,'p«;m.n w.. "ho,.n bee.u.. of .he ov.i lobili.y of high·quali[y e..to• •he p,<>en<e of.n oci. ..d m. ndibk wdh

good fit .nd ocdusion.•nd . h. rep n..[i". eoodi[;on of,h.d.m;,ion '1"" from . he"be" ( L.~CMHC 2001·249, ilke

'C' "'''''TOO''' ."0 O''' ''OOTH . " " .. " ", n . 2J

lO.4mm

mo" of ' he coHoe'ion. w•• ",rover«! withou' ",1><,,). The••1><"u..d fO!" ,h..." .pecim.n, fi' ,h. we k." (, ugge " ing'hey . re prohobly a bIt .m,ll) md r..ult in ' he' ips bein~dose'" ,he r.pr• •• n.., iv. cond ition .nd 'hor of lAC MIIC 1001-Z.alt hou~h the CEJ f,11. too f.. from ' hc .Iveolu. by . bou t Zem_~~eometry ofour re.ul'in~ model i' idenric.1 ro rh. ,of ' he original"'ull (I t\CMI IC 1001-Z).od m.odiN. (il ker­" en. 1985). redu«d by J% '" co,,,,,pond '0 ' hc differencein overall 'ize oflACMHC Zool·Z49(see figure on p.a~e 18).Inte",h. ng<.ble ".el . " .. II....' in ro mpo, ite •• nd c, " ..I><..Wet'< modeled from OIher RlB'pedmen,. 'imto LKMHCZOOI -l. reduc .d by )%_The ro,,,,,uc,ion of multipl. sets ofdlfferen' interrh,ngeobk teeth w., done'" ,I", we could"art with 'he "ronge" .Ihteel se" m d, .. our , kill im·pro ved. prog"' '' '0 increa'ingly more . « um . ly derail.dbut w..k<"T ..", Ultim.,e1y. ..... intend '0 det«mine ..peri­rnen" Hy and qu,n'ify ,he effe" of diffe",,,,, .. in .....tion,,nd w...on ,h. for«. involved . The fur....ndob>e".,iun,,n 'hi' ch. pter reAe« re,ul" wi'h u"'e rrated. " ..I.edgedcompo. it< ",1>< " . with O.t to O_Z mm ..diu. ,h.rpne". P.r·.icul.. c.re w....ken ' 0 m.teh ,h. erupted upper c.nineheig h' . nd . n~le , det<rmining ,he 'ip loc>tion rel.tive '0....n i.1.nd mu,",u1o, morphology_ Robo<., i. mounred on• mini " 'ck hoe ,h.. employ. hydraulic foree '0 move 'hemod.l of'he he.d and m.ndlble independently 'hrough thefull rmge ofmovemen' ,h .. . din: -lOO,h c>l cou ld ..hi.v.*hik .ttempting '0 bite ple y. I' i, de,i gn«! 'o u ett me..u,­able forr•• ,ubstanti.lly in ...." of 'hose a hVing..' roold..,hieve, R..he, ,h,n .. ,empt ing ' 0 lim i' ,h. d.. ic. ,o .n

r---j [ 3.5 mm

[6.8mm

_ I

estim ated "rengIh of .n extin,,, Cll. Ih. d.'ign rould . imu·I. .. v. riou , bit< model • •• nd ,ubse<juml .n.ly... rould .howifthe 00...nd model ,re pr..'ic, l Th. first test ",,,ionu..d . 559 kg f.mal. 8i, 0. ~i... ..d.v•• ••• prey pro . y for8. ".'i9""" an identifi«l prey .pedes for SmiloJo. >t RlB(J. ff..",n and Goldin. 1989, Colt.. in et al.•Z0(4)_ 1replic..eda ·c. nin• • n, ..-bi..•• a, de scribed by il kc,,'en (1981') on ,hemim.I ·, abdomen (fig, Z,~). and. for compari",n ,. simil..bi le on the th""",- Observe" p..sent included John Harri• .l orry D. M..,in . Virginia N.pks• • nd Chri. Sh,w,

Th e fir" observation ", a, tha t Smilodon ..I><". re not>h.,p enough 10cut througli bi"'" hid, unk.. ,h,.. i' re l.­live mOlion be'ween ,ne cutting ed ge of the uppe' canmcand the hid._ A. long a. thi. mOlion included a componentp.a ..11.1 to ,he ''''''h .,i• • • draw -cu,. ,h. "be" , li..d ,hehide, No ,.."""blo .mount offorce (gonem cd by • livingcat). appli.d perpeodicul.. '0 ,h. cutt ing .dg. ofth. tooth ,will cu' in th.. plane wi' liou' ."""i..ed up Ordown m""e­m.nt . If the proper trajectory i. followed . the ' abe" cut,h rough the prey in on' con,inuou , . rr wi,hout ,h. corh,ving '0 reopen in mou,n (Ake,," n. 198J). Howcv«. tnedraw ·cut " all, if ' he inci''''' hook into the hid. or iffor . nyrea"", ,h. ''''''h "OP' mov ing in , he vert ical pl.n• . With •mou 'liful ofti"...,. tlie ind ",,, n..·.n·t reach«l ocdu,ion., od the ,h.ar p.a" of,h. upper ca n;n. sh.ar ·bite doe' nOlo<:cu r (Ake",,.n. 1985)_ The bite " all.d a. ,h. ilKi,or, en·g.ged 'lie hide in , lie.I>domcn . " . mp' (fig. Z . 5A~ we appli.da force of 5.400 Newton. p",pendkul.. to the upper c. nin.axi, .... itho u, pulling the ",be" througn the nide (fig , 1,5B)

F"u'" n. A. Th" Rob<><., k,ll;n, b,.. <e<' n.,U'" e. Rob<><.."","n,ed 00' 0 . 80bc.. X·ll1 ",od.1 h, . .."I" "ock00..(,ou"", VO<k; Wh••I..)

f ;,u'" H. Th. Rob""" ,. " n"u.. 0.;01: po,;' ;on.d fo, m"io, .coo;n" ,~.. ,-b,.. on 'h" . b<I"",".. of. bi,,,,, .poe;"'"... «"""..,Vi<•• W....I..l

.n<! diocon,inu.d ,h• •".mp' . ft<r ,h. core.., , u " ed to hf,off t~. grour>d

E..min"ion of, he"'.ul1 ingwound lng,Ulindic.,ed ,~.... !'hough ,h. ,.b." dod p"r<:' , he hid< . nd f.ay ........ 'heydod no( po........... ,he .bdomin.ol c>,i,y d••ply ...-.oug~ to

Fi," ", 2.1, A. V;d«> ,.q".nc.l,o", • c.o inc,h...·b ;'• •"""pI"" ,h• • b<I"",.n 01. b;",.. 'poe;""". 8. n , b;,. , ,,II • •• 'h'i. c""" ".., . ",. Thob,,,," .",,, '" 1,/' oll ,~" "ou"d .. ' hofo'eo ;. ;n' ....ecI '" 5,_00 " ...."'0., (CO" ,..., V;<~; Whocl..)

produeo.n immi.....,dy to".1 injury. The un.n imou. ron..n·.u, of ,ho.c p...",n' wos ,h< b;,<a. ,.,,<d wos not ~i,bJ<,

Sub,",!u.n,ly. Ak<"t<n (2005) poin'<d ou' ,h". hoc.u",,h. forelimb. "f5miloJo•• ro •• ,,,,moly robu", ,h< ....ull.n'fo,a on ,h< ,.b." wos nOl!imit<d '0 . for<:. <qu,! '0 ,h<prry w. igh'. Th. ca' could apply foro. in.xo...of'h. w.igh,of ,h. P""y by pu.hing .w.y from il> P""y. During , ~< 2003S.b.,·,,,,,,h Symposium oTg. ni"d by Ak."ton, w. , .pli.c.t<d , h<.h<ar·bit<5<q"'""e on '~e . bdomenof • btl11 C"""-'(elk)c.",... wi,h • body mo., of .bou, ISOkg (fig, 2.7).

On , h< ini,i,l . ".mp' ( fi~ . Z.S). w. brok< • compo.iI•..1>«.Th. n..' .".mp' w•• '00 .upornci.1 ",seve"he in,e.·tino. bu' we subsequemIy.uc«<d<d in«<ou'ing .n .bdomi·n.1 bit<. Ikcou",of,h. sm. ller di.meterof,h• • lk .bdom.n.

'he bi'e pone""ed ,he .bdomin.l c.~i'y .nd ""c!'Cd ,hcin,<"in<s,Tbi. wound migh' no' I>< imm<d,..e1y d,..blingbu' would c.".inly I>< f."LTh< for« r.qui!'Cd to pull ou'of'h< wound woso"or II ,000N. ..·''''' ., wi,~ ,he ~idele"ing

mhe, ,h.n king Cut by tho ,. 1><" . As ,h. m... (1.14S kg)uqu,,<d to k<. p ,h••11< on ,h. ground w" o,'er six hm..,~.. ofthe P""y. we used t~e mock hoe to hold it down , Fr«·m.n ond Lomm (ZOO7h) npelim<nuny d....minM ,h.

f.,"", 2-6.An. M"",.n.1w""nJ 0" 0 b"on ,_'men ,It.,1><,", <0'o~n 1'0, e"m;n,,'on. de", on",..in, ,"it '~e ""und;n '~e ";de a"d f,n, li"oe wu ,op«f>c;aland did not . ...nd;nto ,~• • bdomin.1<....y. (coo<l"1 V;,,;nio N.p....)

f;, ""' 2.7. Til<op"mo", po, ;'ion fot "" , ;n, a coni... ,". ,,·bi'.!><on, mo,,,"d On ,~• • >do",." of on . 1. ,_,,,,.n. lcoo,""VI,k ' "'~eelo'J

. n,enor-po" . rio ' "It;m'" " ..ngt~ of S""l<>J.n "ppe' ,,­nin., to bo 7.000 ~.wton,.

"h" '!Udy of [h. photograph;c ,,~o.nc.. (1 f"m. 1"condl of th. , ·.[;OU, b;" . " .mpt> , ;t w., app... nt th ..the <m mc> had "opp«! ,Ii<ing and " alled (~g. 2.9M wh.nth. oppe' inci",,, ... ach.d [he . lk·, h;d• . W h. n the ,I;c;ng"opped. ,h. "I,, ;v. "",,,m.nt of ,h. fi"o... , . 1", "opped,lifting the <.'c." off,he groo nd. Oor n." a".mpl. wilhbo.m, to hold th. elk down. "t"<tchcd th. hi<!< with th...n;n.. bot ron oot of"..·.1bdor< r.ac~mgt~. elast ic hmit(fig. H Il),

W. th.n repo>it;"".d the ....... ",i,h I. rger b..m.do",to rhe wOllnd. At a pe . k fo,a 0"" 11,000Ne"'ton" t~. h;d.

F'8"'" 1,1. to,de of t~. abdo"'en of ' Oe ..po' i",e "'" el, ,poei·meo unde, ten,,,," f,o", t~e "ailed ....r> " t~e pointof co",·PO" " ' 01><' fo"o,• . (."",t.., Vicki"'nule,)

to r." ,h. Canmc>. When the hide r..led. it w.. st ill pinchedbet ",een the opper and 10 "" , r inc;""" and..n be".n [0 pollf... with""t co";ng lfig. 2.K:~ In th. next r"m. (fig ,1 .9D ),

th. in.." in. ;, >eV . ...d as , he <m m., te.. ft«.Sobse~ oent wound •••minoti"" show.d th., ,h. ;n.;",,,

did not Co t th. hid. (fig. 2,10), Thi, indic..., th.. the foree

...quire<! to "all th. bi,. "'" ..1"i..ly 10'" .nd prob.blyo<curr.d w~~n the ;",i""" " .rt.d pulling ,h. hid< .I""g inth...m. rdati" mUljon as ,lie upper canine>. O ur ..st>

... v. alecl th . t cutti ng by .p..ul. .. inc;"''' doe. not 'ppe.rto be po»;bl•. n...... '1, Orhelpful to fr.. the upper".nine>,. nd wou ld ' pp".r '0h.ve no mOf'< role in tho dirk·toothkill ing bit< tli.n ;t doe,;n the bM, of~"'nt ..... B...o.. ofthe I.rge forrc> imolved. " ..I <an;ne, and ;nei",n ",e,..u..J. Althoo~h ... ,in ..st inci..".. . re , harpe r. the "eel rep·lie.. w.... no mOt"< round.d th.n m.ny W'om ;nei<Ot'pee;.m'n' in ,h. RLB col1«tioo. Wh.n w. rete" ",,,h sharper;n"sot'> , I doubt it will ch.nge the re,ol". Th. "eel ..be,m'rgin. o"d W~re , h. rper ,h.n typ;c . l 'pecim,",;n th.RLB ool1«t;on,

Therefore. [h. c.n;n. ,heor·b;t< mod.l.s p""p""d by,\ k."t<n (19Sl. l OOl),;n ",hich th. inci",,, arc occlu ded and,hear th rough the hide and Ii"ue un,,1 they cut free of thebody ofth. prey. i,""' . worll.bl. hypothesi,. no! on. bi"'nor '''n thin ner-,k;nn . d prey ,uch .. e",·.,. Add ;t;on.lly.be<.u.. the bIt. fore. of IO.JlO Newton. n«d.d to ocdod<[h. inciS<>f< f.il . d to prodo« <utl;ng. ;t ;••1", d • • r , h.t the;n.i"" row i, nUl••pabl. of ,h""n&throogh ,h. h id. of •lorg. prey . nimal.

A d"k·tooth throat bite m<><kl ,; mil.. to th .. of Mit1 in

,6 TH O OTH" , ....· TO OT H '

F"",,, 2.9. .t, ...,,",ond ."empt ' 0 m"e • c.oine ,.e"·.,,. ,, ;,"" ",I...." On an , I• • !>dom, n, ,., ""'0' ......" ,., .'t. " all••nd 'nO""". hi" • • tn, ,nc;. o" ,ng' g, . 8. Af'" "'1>O'" ,on·,n, " " . "am, to .old ,., el~ """n, ,he aw l;c"ion 01 mo',fo'e<, C, F., ,!>.,"po"''''ningw" n la,S" ".m. ,,,.It. in ' h.nid...a' ins a' a "..~ fotc. of 11,00<1 '1""" on>; ,., n,d, ,,"".~" 01,", OCCI"d'd loc;>o" " ,' ho"' ,h,,"ng. D.ne 'n,""ine,be,n, tom by '~e un ine' .. '~ey".II&ee oft.e n,de, '1ote ,~ "

, ne n;d, n" 'o'n f"~er , n,n ..in, c., by ,~, " all«l '""",(court.., Vdl Wn. ,I..)

(1980)and Kurten (1952) wa, ..plic"'d .u«, ssfully. wit~ thepmviOO , ~.. ,~. in<i..,... d id nO< .ng.g. t~. hid• . [)(m. w.lI,,h. '<sul'ing wound . (fig. z,ll) 0" de«p'i,<ly .m.1l cleancu ts bu ' , ..ult in the ",,,ring of 'h , jugu lar Of",o,id a" ,,·ie, .nd would be ,1m"" in." mly f".1-

Thi, d ,.w-<u' bil<On ,he ' h,oa' Can be penor me<l expe riom' ntolly, but ",~ui..... '",nl< pr«i'ion. u'ing nn more­Mce ,h.n • Iivedirl<.' uo<h .., roo Id, . ..... n.blygene"" (Iig,Z.lzA ), A, our ,«hn;que impro"u I am ici!" l< 'h, force re-.~uire-d will , ub"anli.lly doc",a... The p..cise '«'0' mustbe ,pplie<l if ,h...ber i' ro <lice ou, of ,h. p,.y (fig, Z.l ZB), If' h, bite '''Jeeto,y i, '00 , hanow. , he wound will be , upern.ci.l . If the tr'j«,ocy i' '00 d••p .nd follow, ,h. n"""l arcof 'he 'ooth ui., 'he bi" will "all. and ,he COl will he luckyto .",ape with ,h...he" inUc' (fig. 2.Be). A. long " ,h.prope' tr 'j.ctOfY i. m.in"in.d. ,he ..be... will.1'( 0'" ,h.bor'om of ,he """k (fig. 2.I)DJ.

Real-world applica'ion of all 'h, bit< model , i. difficultdue '0 ,h••moun, of di>to,hon of ,h. pl'(y ', bodY" ,h. bi,.

p,og~"OS, L..g. ·se.1< movem.n, of'h. P"'Y ",ta'h'. '0 ,h,p'N"O' con be ron",.ined by pow.nul fo...nn" b'" fin. ·",. 1, adju"mcnts in uppe' c.nin, pl.«m.n' requir< tactileinput from "'hi.... ". hp,.•nd n,'V" in ,h. periodontal lig• .men, .nd pulpco'i'y, Tac' ile inpu' from ,h. 'oo<h ce,vi • •nd'be question wh..her gingi.. could ex i" on i' (Ri, iel'( .ndWh..I.,. 2001) i, . ,ubjec' "ill unde, study (W, rd. lin ondsardella. ZOO6~ Thi' i,,~. i' not e.m... l '0 ,h. propose<! bitemodel bu' m.~ly '0 OUI und.'stand,"g of ,h. ",,,,ce(,) of,b...e,il< inpu' '0 "h;,.ve ,h. b;,e.

A' ,he ini'ia'ion of'he dirl<·'oo,h killing bit<. ,ubstan'ialjaw .dduc, ion fot-c, i, in,-oJ..d " high g.p< angle wh.n theuppe' conin., pi.= ,he hide .nd und.r1ying ,i"u• . A, ,hebite ptogl'("", .nd ,h...ced ]>',h of ,h. <lieing 'abel di­"''!lOS ro".rio, to ' h, proj.ct.d.re of the ..be, u i" ,h. jawoddue';on componen' ofthe ,«to"um..'ing on 'he uppe,coni"",di mini.li••, A. 'h, h,addept,ssos. n<ekand fo.-.limbmuse"lotu", becom. ,h. p,.dominan, "'''I'(' of forc.Ther<fo,•. M dirl<"oo,b,. j' w .ddue,ion mus<~Io,ur< .ndmandibl. beam streng'h are ..cy pootindicOl"" of"strong"'.."u, "'w••k. ..be, ,, 00<b killing bi,. "reng'h. $<>phi"ic"e<lcompu'er ' <ehniqu.., , uch a. finite .1ctn<n, analy,i' (FEAl.no rna".. bo", w.II ,,<cut<d . ar< no be".. ,h.n the moth ·.",..ie.1 m<:>del on which ,heY'r< "><d, If ,h. mod.1 in·volve, a ..ric, of m",',rn.nts, ,~ch •• a ..he, ·'oo,b killingbit<. ,h..., ul" can be no be.,.. ,b.n ,h. a" um. d ",~u.nce

. nd given condi,ion• . As Cool< (Cool< . , . 1.. 2(02) "ot., in

'he in'roduc'ion '0 bi' "Xl , "' I, i, po."bl. '0 us< FEA \'10'

f~l.lO- ,."._~ ",,,,,.-..__

""-.... ....-- -""-_.. ""fa; (Ioc-.:l b, "" .. -.. and ""__01 01 _ .. _01 _ ,."._ .. ""~ _ _ "" __. +c-on.Vo<k>

""-

r...... Z.11 ,,,_ r.om •WC<n.fI.l _ ..... .... _.--. ,.". ..... 9 .. . . ~ _ <ut, _ "" ,......,_

and....u_.-,-.+c-on, V..10~

g...... ""hll<......,ng hIl l<k_io<d# "" , he .,..Iy_ .....hod

Of ,he pcoblcnl to ",-t""h It pph.... ,n' '' 'ngronocq""""'",hot: m.I~ • •~ from cmbo" '" ,od,u" : hm in •• i n l woricI . , he ~ lor C1pcri l polton,olog~I ' ng ....l lortt "" 1boo!og;c. l ',,>IX

(EricklOR .nd OIIOR. 1_ Eri<klOR 1.. 1_ T1Iorn.>.IOR

., . L.lOO1, M.nnlng" . 1.. ZOO" Fm<m.ln . nd u ....n. UIO/i:

Z0070: ZOO7bI, Oet" looi ,he m",h.,rodon, d,tk·,_h kllhngbile .....,,11 'gmg. .nd ,he ;mphn ,ion. for [~ nlm ·

n."'••nd "" "'fr1... ",'lIh ll'pe" C1c«d,ng 00"• • nd hY"<rimmo,plnc mnrpho,k'llY. . ... 1'')I.ly un."'d,.... Funh..••penm.n..1",-ntk .hould help ;n ,he de••lopm.n' "" ;m·pro..d bite mod.l•.

Ifou,b". """,.11 ng...fi"," ... lily. ,he . ....nc. of. I".« i·".1d.r.ul, kIlling bit. w• • • ""Iou. probl.m for dir\:,"OOlh

r...... l-ll . ,,""'" _ 0I.__~ A."'" , 01....__ boo< .......__... ,.". ___

_ .... .-. __"","""",," boo< "'""""" ...........--H. Todd )

r 'I Yro 2.lJ. "0"" '"""...01• 1>.10" "I'<~ : C. 1<1. 1,.""lOry t1,.,IS lOO ....p t>«."," ,,,".., '••11owo"l ,,," ..be' '" follow ". "".

""I.." '"'0'h. "'"' ""Old of l>." "1 '"_I"....~. 0, ".....P"'I'''''"lon • ,.,,,b" "'pK"'" ,,, b,.. '"""'I" ,nd ".' "flh•prr, in • "om;"",,"• •r<. (d"w;"1 .... Todd Wh....,j

,8 THI OTH" SU"-TOOTH'

COl popul.,ion., lffm sume re.sun 'he COl could nOl perform'he spedahzed 'hroat bi'e th., w.. mod ele<! succe"fully. OT'here weeo nosui,able large preya"n.ble (a"uming sm.lIerprey species were .gile . nd difficu lt '0 capturel, «avengingwou ld be 'he only eomamingoption_The required prod'ionofthe dirk,'oo,h bite is under><ored by ,he pre..",e ofdirk ­'ooth 'P"'imens ,h., h.ve bro1<en one Orboth upper can ine'«th . Clearly. 'he.. indIvidual< sU"'ived . nd continued '0u.. ,he canine•. beeau« .pedmens of..n show wear on ,hebroken , urface,. Thcrefore, it i, rea""'ablc '0 a" um. 'hatdirk-too,h ..ber r .... ,uch a. S.. iloo"o, could .urvive on ani<><lividual b..i. as .,;>venger.... do injured. ill. OT aged e.­,an' relid•. A. a tamn. howeveT, ,he dirk -'OOIh killing bite i.aPl" rently so prey.specific ' hat ",'en reb li.ely .man<h.nge.ur d isrup'ion of ,he suitable prey supply would r••ul' in e.­,inc,ion for ,h. cat,

THE SCIMI TAR·TOOTHS

In rontr.st '0 dirk ,' oo , h. , 'hero is ..ry li"le publi.hed li,_. rOlure on 'he Kimitar-'ooth killi ng bite, The i" ue of prey..I«,ion an d pu" ui' has bee n addre"ed in term. of ,h.""'eo cu"orial ecomorph. bu' no, ,he .c'u.1 bi,., desplt.'he consi<!crable cranio-dent al differences be'w«n dirk­and .dm" aT-too,h morphologt<,. 1, seem, logical 'ha' 'he..olu' ion of a dirk·,ooth would in.olve .,a.,ing wi,h a"'imilOT-!OO,h , bu' '0 date, ac knowledged tra"'i'ionalfOTm.in ,h. fo"il r<cot'd ha.. not been r<cogn ized_ A po'­sibl. candid... migh, be 'he ""goo";O" ,pecim.n OO OAlS41) from , h. Ari ka..ea n of ,h. John 0 0)' Ba.in (Bry.n'and Feomd. 199$)_ S->be" wi, h en.mel .nd. cle.. CEJ'rcno , com monly <!cscribed on nimr>vid saber-'oo,h Cat>,More often than nOl, ,h...be"of nimravid sku ll, arc .i,herm i•• ing. foun d broken. ,,"'OTn. or dam.ged to .uch 'n ..,.n,'hat ' he e EJi, not avail abl. for"udy, However,JOO.~ l841i. rc ma<kably well pr... ,,'ed_ I' .how. a level ofdetail notcom munly pr..en, ror nimravid• . The ,pecimen', , .ber>have a Coa""Iy serrated anter io r maf gin over 'he fu1llengt hof'he enameled crown . but the hng u. 1CEJ ,ho...s remark·.ble converge"'e with S.. il";oo (Wheel.. <t . 1.•2(04). Po­S,,,,,,<lo. is . rem arkably long-lived ..be r-'ooth genm , bu' i'h.. not been ..ten.i.ely ,,,,d ied.nd maycontain .urpri,inginte"p<cific d ifferenc.., pn>sibly induding bo'h "'imi' u ­and dirk-'ooth for m•.

Be ,h .... i' m.y. ,he SCimi,...,oo,h i. a vi.ble morphol­og y in it> o"'n righ , . nd nO, jU'" pK<ur"'r '0 ,he dirk­'oo'h_ The tadt a" ump'io n ' hat th e ",imitar·tooth i. aj un iOT 'e"ion of ,he dirk-tOOlh. and ,h., .i mil.. knting bi' e'heorie' oWly to 'he "'imita<-too'h, i, "rioo,ly fi ....ed, ..examination of ' he la rgely non·overla pping difference. in'oo,h .nd " .nioden,.1 morphologies indi<..e, Scimitar­'oo,h, ha.e ,horter upper can ines, u.ua 11y wi'h coa""lyse,,"ed rutting edges, an APO '0 crown hClgh ' raho rang·

J__

F;8"" 2.1 ' _The IinS", 1p,oiil. of. H"",o,.",.m UPP'" con;,.snows ,t>c <n, rWe r;" ;cfe. ,",,, of ,t>c CE). ind . d;nl ,n. ·"ep"';Ih!" P'OfIO""""- LAC"'HC 70001, CoorBtC Page "'''$e'''''(co",te.y )ohn "', H. n' , )

ing from 2:1 '0 2 .6:1• • nd , ypi<. 1APO '0 LLO r"io, ..ngingfrom 2,1;1 ' 0 2.S:' (fig, 2,141,T h. ,harp. se"ated margin ont he • n..rior and po" ..;"r edge of,he uw< rcan ine . extend.over 'he fuilleng,h of ,h. en amded Crown . The inci'ur>arc larger.nd more rob"" 'han 'ho.e ofdirk"ooth mach·.irodonts .nd a,.. separated by . di."ema from ,he uppercan ines ( Bikne. iciu ' e' .1 .. 1\>96), The procumb.n, in6,o,arcad. and .hort mandible result in an ...·cn ,hort...ff«-

ti.. b" . , ••bu. for , he uPf"" ~.n'no•. A. II ,he u " (or , hed,r\·~h•. , he low.,. ,.ni"" ;, d"ma'i<.lIy~. • ndJ>P< .np,••~ abou' 90" . ... ,he1l>on.,."!'P<T c.n"..........""'" <1 n« for bo,~ plac~_n' .nd ,he "'1ull,d....heI, 1

n.. 01. the .n: 01. the ~h (am'" 0I.1IJ'POf~.n,,,. <u u..~ ..h><b ;, ..,......l """' on to

,hewlI.- ndlbula,j<:II.-lTMJ lin S " . (A .....,,,, ) and ..1In d,r\....... m.. ..clooo to the TMJ in .ho ""m­....,·"""'h.u.ihke ,he dut.-<OOdl ......pI........' . • n 'n: drown"""" "" "P 01.the IIJ'POf ~.n nd« ,ee"d. tho TMJfolb_h", ' ho IIJ'POf unmr Th,", m....,..__h

'hofull.v..g.h. ......,.,......-.t "Pn mplr_._ 01. 'ho hoad.nd ....ndlblr to<UI aI 'ho h.-..or un,,1 ' ho il'fClOCn can pcnrrr«< no bnlln

n.. CEJ oI.'ho """,,,.._ h "I'P<T c.n ;, d,."nctlyd"f...... in oh>pr """" ,he d,r\_h can" 10 ' ho dll'tin« frum . ho ..~'"g...-a<e>t ., ' ho . nlnior 'II:,n . nd>loptng opoe.11y up '0 • ><>mo<WM ""'"'zun'.1 "..",,- . nd fi·...11y oharply oIopong 'pio<.lIy up at 'ho poOl,nor ....'II:'n.Il<>Ih 'ho "".l.nd -",,,,," hoigh,, ,n proport lOfl ,o.ho API)

may v.ry bt'"""n "" ",,,.NOOIhc." . nd a" '1u.mi' .,;."Iyd.." " gu ..h.hI<bt, ....n .\f4<lt.ircJ...nd , ... morr deri"edIIMIt.'~'"... IWhcok., 1004).

Whil. ,h. function41ba'i' fUT.hco. CEJ pr<,fik... ".li,..,i••,,11 bting .."died fUT d' rt.·,oot.... CEJ",". m , n •• " ... f,,1,. .....ic·k".1 d,. gno"ic f•.,,,,. (W. tdr hn .nd s..,d.II•.

1OOlI), For ",'m" 'NOOIh•. ' ho cr m.n'"m·covrrrd " gion .,' ho <rrv" (nrc k) of , he <.ni".. bt, ..con 'ho CEJ and tho.lvruI., tx-. is con..n" on.1 ."""gII ,n ""po . nd ~1.,i...... t _ ..n ~• ...,.bIy ..poet it .... rov....d .. ,,10 pn·

pv ho ith .U '" <au.nd moat .. m,la . c.,-n.-. ,h«;odont ....m ls, Tact'" fr<dbock (rum ,10..

"P'" ...... Idboo.""., ...." dunng ,he ..im". ..._h k,ll·"'ll bole. n.. IUt >top '" tho CEJ profilo perm'" 'ho "I'P<Tun".. to boo ........... to "" full <roWn~ bri>rr ,""(tUf«I'" Id"""aa"" P"'Y. '1'U"'""'ll' ..Be n,ndland 10 """""& n.. pndomi ...., 01 P ,he......- 01. !he ~h would t , "" ..Be.......Id ia1~ """.-"'. 01-..~.WOUnd·W1<lm,ng """"'" b,-.ho cd poll"

nor ...."Pn, .. !he COtWI< .......0lh0I....... E ho 10_·

~ of ,"" ""m.u·~h.h.o< Iargn.nd ""'" po<>­<umbo............n:odro. .nd '" ,he .... .s.n-.l ...... . hopogtV. bn...,..., !he "pp" can ......nd tho ,_•.-cod< ;,

""""........ to ' n ggn- ' ho r<fIc.;..., ..10.''''1 n,,,,,h .........tho prry iI pi""hrd bn " PP<T can,,,. .nd il'fClOCn(~ Z.1lA), Th.....AA"'" ,he ll'fClOCndo _ play • rolr,n , ... k ,Il,nl bot• . In ,he morr drnvr.:I Hornoohoriini . ,10""not . he ~. ... and ,ho ,nc;""", brcomr P"'l':" ......ly It>O«

,m-oIvr<l ,n m. ki" l ,he tn,. in ...". lOtm•.Th" P"'ll""1Oft cul mi".,•• ...-it h x..",..,I., 1t<>J........

IM",in .. . 1.. 1(00): , ... inci.......~ lu ge, .nd ,he ,,,,,.d.

mot< procumbam t n ,n . ny ",he. "'m,,.r·,_h d. ·

..rib«L n.. "PI'" tow... inct...... h ..-rntcd .....go....p<Omi...... IirtguOl . nd Iobt.ol "''l'''Io nd hnlo or no dia·....... bnwrrn ' ho "PI'" can.....nd ,"" ....,...... (fig. Z.IIB!.

A. full <eduooon upprr .nd ........ irlcIf;<:on (i ( ,,""""")

mnl> '" _n 'loa ,...,. yha", h<rn ablr to <UI .hn• .-or"'" 01 ....... prry ( <uok,,'cuncr" "0'" pi, bole model;..apI<o andMa...... _ , .......h,n, .... .....'ng ..............tho "'lin in<isot .",adn ca_ ""- lol. rnn .nd~-

coobr-cunrr bot Id__. , h>«hlY oprao1>o:lft. prrodJ."ll

CKprnm<TII" ....,6< lOr !he .......... ~lri-ing pronin 'M ..........tully ,la.~h " ..ploW-ogyon'Rm,L"ruh_.SharL-tI ..c..d -..­....... ""'h w;,h ll"""""11y Rmw. """rh<tk>cI to tho It>O«

drnvrd"""""",,", mciounandproduc<d<v......tng noloooprryol ry,ng .... (Fn.zrt , _ , Apan frum ....

...... c pro:a JiI~ 'po••n...., fig. 1,16~

..-hieh bony ;a.......... ;,k--.. "' bot. ·....bolu.from prry moch u 'lI" , han ...If (F,nk . '''''J:Pau ly. 1994:Frttma" .. . 1,. lOtI').

At ...". Iovr!' ..'. know t"', ,he cool""'''''''- bi« i< fr.·.'hI<. bon frnm previou. bitc m<>Je1• •ponmrn" . w< .1",k""",' th. , " n " gen' """f'hol<'lli<.1 eti,m••ppIy, In both",. rt.. and pi,.nh.. ....h.r< repl. <rd••nd bOl, capabilih..1"""."m. "nworn , und.m'!l'J ....10,Thrpiranh. ;",hebo"..­li ving pro>:y, having bony j' w, , h. , .Jo "'~ pro,," duringbiting (AIf. ro " . 1.•WOl~ In piran.... ,II< ,ec, 10 .Jo....occludebu' h. ...n ."rKlrtb;«; wl, h ' he;o w poantoned 10>Iodr or...... 'ho lower ...,10 ~.. 'ho "\'l'<n(lle .......u. lOOOc ..do

......."".10(" Wor b1adc.~ n.. coni,,1Ofl in ca...."h bladed,rriIc>n i. lOt...pp<n.nd Iowoe.. '0 ""'" blodr to lUJr 10-out ...rrlap• • nd .. morr ..m,l.. ' 0 ..,.. or \>011: "",....

An "nfun........ ' 'P'C' 01. bot h ..pm nd d,"'"oborn- 011"',"11 ...... on .. ' ho d'fl\<ully in ".hng "Ia·'i>'< up or down. \Jtlt.1 fun. 10bole pc' kit ",.d._hi_b...cnn •••..-. dep: ·.h.nd..._ .Ic "pp" conll.... ,"'1uooIo-d. _ """"'" deorrm, ...

,"" pr<tia< fnrcn and c.opohl. on 01' ho bole. AI' ho po",nIto"".10. !he c:ookie·ao..cr bole .....u. and . Ic brurr ............. _bog .........hfu] , he c• ..uacb_.

A Rrong~ oI.ho -<OOdl be tho arpa-bob,,- 01. drf...1r be•• ""'" ,he uudr 'T'tm......Id havr .ffordrd "",m•••~_h ~ 1ft l""""'l .....H .nd Xt= P' '" partocula. , .ho lI..ibil..,. ~'.k,ng llr-rP"J ..-hen fl<tt>N.ry or ..kmJlargo I""Y 01• lU< that bty<orocI.ho~odI oftI.n. upp<t' , kiltmg

biotc b,- innic"ng. · wound,n, bt...- I( ,"" rnough..,....... ....n, 10,..... poch, ..",n'ng'n un""".can km largopr<J in 10.. 'i.... ,h.n" ,.ke• • 1_ to "or' ,10,..,., ~Iomp(Kruu k. 197l) ,

Ant6n.nd G.Iob,,, (11'99) ha" . pphed , ... dir\·,,,,,,h..nino......-b". model of Ak rcn 119" ) ,n ""im".,·."",hmach.ir<>don...urh •• " /1,..;... , Roboca' ..porim.""

JO t HOOtH" U ..~ - tOOtH'

f ,g"," 2 U . A, Tho",,"<.>1M.,"irod..,di"'.m" (EJ. , od p"' _,"mod gin,i, , 1,.4>tion, n,p of ,he ,.be, .nd ;"" ;'0' .." doIW~..I. ,.200(1), e. A mo", de<i~ ·coo' i.·cutt.,· di.,'.ml ,eEl. 1M p""um'" l ,nli..1,.Iobon,nip of " b., OM ,n<i, ,,,10<><1. (Wheel. ,. 100<)_ Not. Mr.",,,,,. in ,n. '''''' he"nt '<&"'";n ,n. eEl in o,n.,wi.. , imd" ..be, p,ohl... (cou"." V"~i

W~•• I· 'I

ho".., shown ,h., . fi>r ony ..be,·"""h wi,h . ,e mOlely <on­v.n,ional inci"", row, one< ,he inci""" ne eng. ged into ,he, kin of ,he pr.y. ,h. ca, ..nnOl f.... it..lf wi'hou, ...open ingin m"",h_ By P"Xe.. of . liminOlion. ,he bi« model fo,><imi"r-tOOlh, >t.rI> wi,h bi' ing dov.'n wi,h ,he "pp'r c.­n;n... which have coa... ",wion, ,h., f..ili..,. the drew·

cu' erre" on ,he downmu1<e. On<, ,he c" h., c1o><d iumou,h a, fn a, po..,bl., it ha, re..h. d whal J 'erm • "'poin'ofinfle"ion"' . nd mu« mo"e ,he uppe,<>ni"". in. J irreten,dir«tion '0 w;,hdraw. If ,he c" p"'1I, back •• i, open. iumou,h, ,he l""l<riOT margin, of the " " ... d "erh wouldal",d,.w,cu ' 'hrough ,h. 'i .."",of'h. preyon 'he "P" mh.

f 1: 1.16.n.. _,.,....of """"". ...... _ ,.; ("''''''''I'.....tf ~: '0t", _PIN" . .... _ d.", ;.. " mee<."I:C. .. , full do>ed poo."". _ ""',""_ d odpH' , _', p<od"""'1 I "'.. _ '" ( -~ ,...• fI'<o<\. ("""lfl, y ,d; W......'l

}' ' H' OTH " ,n"-'OOTH'

p",enti.lly le,ving. wound h. lf ' g" n to double tbe .nt. ro·po"erior width of the tOOlh. Prol""rly . it<d on tho no< ~ of, uit.ble prey. thi, ...ou ld be .1mo" ., dev"'."ng.s the dIrk·<oolh bile. bu' would h"e , he .dv.n..ge of not '"<luiring'he '.m. high dogr.. of ..etik input .nd puei'ion,

BITE MODELS AND FORCES

Sub".n,i. 1bit<forr. i, T<qUiroJ to effo<:' the Kim ," r·'oothbit<_Unli'" dirk-tOOl ho.Kim l..'-too<h ,'"" .mployed m.n­dlbul., oppo>ition throughout th.iT killing bit< in . ddi, ionto mi ng h. >d.dopu>sing fotc. on tho down" roke_Thi' w.,followed by . pp1 ie>tion offo..e. gene"'t<d by the mUKul.­tu« of tho fur<limb .nd no<~ to widen the wound on thoupstro"'_ Mode rn pucti,,1 bit< (o,co " udi« bog. n withM'ic "in vi,o" "udie, on op<»Oum,. "'i'h 10"" g.p< . • ndm. ximum forre ..qul«d .. .".c1.,..,o occlu, ion(Thom.sonet . 1.. IWO: Thorn..on. 1991). Thom..on·, opossu m,.•rodm",' ext. nt c.mivo,.,. opply fotce wi'h the uppcr c.nin«' gai"" ' he p, ey th.. i, «l u.1 to . nd opposit e from the oc·c1uding fotce .pplied to , he Io"'er <.nine,_ Thi' i' not theC's< for , .bo,- tooth' ins . neral . nd dirk-tooth, in p. rIicuI. r.

Tho- "'ull . nd m.ndible . ... not an isolatro free body.•nd,he.. i' no n«<l fOT tho for« on tho m..ill.ry demition tobo equ. 1. nd opposit< to the for« on 'he m.ndibul. r donti ·'ion . Tho " t . nd puy . re int..ronnocted by tho po",..fulfo.-.orm' . nd body of tho c>t. Unli'" conic. I-tooth ca.. , ..­bor·toot h, h.v• • n ..ymm<trical d.ntition. with bbdeduppotc.n ine. oppo>ing • m.ndiblo with ..due«l in<I""-Iikelow.. c.nin." The foree that c.n be .pphed '0 the low..dem ition i.limited by the ,,'eng,h ofthe m. ndiblo .nd ron·n.ctiv. , i»ue , bu' 'he upl""r limit of fotce . ppliro to theupporc.nin.. i, only limitro by tho" «ngth oftli< , kull.ndbody of'he <at . nd ,he strengt h of its gr. , p on the p...y_

Aprim. ry issue with "bite for.. dub" ,hinking (w ro< ..

. 1. 200s) . ri"" when )'Ou ,tart to compa.. , .ber·tooth bit.,'0 roni<>l-'ooth <.mivore bM,. u.ing m..imum bite for«nlu." The us<of ,implt£ro I.v.r arm model, £ts "ry wdl.t 0< ne.. fullocdusion. Howeve,. bi.. fore< " ri<.with gap<.•nd i' i. ,he <h.pe of ,he gape'v.",u,-fore< curve ,h" di"in­gui'he' ron ie.I-.scimitar-.•rod dirk-t<X>t h bit<model., Unit...conic. l·tooth c.... scimit.r·tooth, h.... . TequlTement furhigh bi,e fme< from full g.pe '0 "",lu.ion,The prob1<m withI<v.. arm, i. th.t the m«h. ni,,1 .dv",..ge of the t<n'ionel<men, (mUKI<) g...., ' '''''y '' ' futlCtion oflhe ,ine of the.nglo be' .....n the mUKle .nd tho lover orm. lde.lly. thi..ngl. i. 9O".t the POlOt wh. .. maxl lOum fotc. i...qu i..d_For.n b.rbivo.... th i. i. pr:Kti"l. in which "'" the cosine of,ho g.p<.ngl. h.sthe ..me ..lue ."he ,ine "flhe I.""rm.ngl•. which wou ld be 1.0 .. 0" or full ocelu"on_

E.pand ing ,hi. model '0 larg.. """i<. l_looth <.mivote.,uch., """,.,," 1,0(Afric. n lion) rai,., roncem. bec.u,. ofg.pe itIC ....'" . nd uncort.inty . bou t tho force ,ver<u•.g.perequiremen,,_ In . <.,ni,'ore ,uch .. P_ko. maxim um foree

for the carn. " i. l would be ...qu ir<d .. . g.pe ,uffici.nt '0de.r the <arn i. I, plu•• foldofhide .nd .ny itICluded fie' hthot , h. lion ants to 'w. Uo.... M..imum fotc. foT tho c.-nine, would bo >t . gape .ngle ,hot de •.., 'ho c.nin••. plu.a ,.. rtebra .nd .ssori..ed mu,.ubtu« for . r.erk bite. but a,.condary «qulr<ment " for .uffici. nt fotc. to pinch thewind plpe.t des< to maximum g.p<dUring . thro " biteE."n, «mi'",•• ,ypic.lly h..e .. Ie." 60" of g.p< (Rw in­. ky. 19SZ). Ifthe im. gin.ry lion....h.n ..« uti ng . bite. i. or.l(l"of g.p< "'ith its <.nine . eng.ging 'he venebr. i' w.n"to cTu,h or "parate , . nd tho lev« . em .ngk i. s<t .. 90". tho....ul..nt tr igonorn<tric {unction (sin 90")i' 1.0_ .~, the g.pe.ngle in<re... , '0 60". the lion "'ill h.v. 0.1\66 (. in 60") .,much furre .vail.bl. for ,he throat eI.mp. The"f",.. thefu nc'ion.1 lion bite i. n·t th.t dIfficult to .chiev._

Ifwe h>dbeen dr.fting. blutprint for . bear.•nd ."ned...ith. m..imum foree "quiroment .t occlu'ion of ....n­tia Ily0"g.pe (bo• ..,boing omn ivorou, h..e bunodont <rn.h·ing mol.,... not ' ..nch. nt slicing c.m."i.I.). 'hon ., 60" ofg. pe tho .v.il.ble foro. would be O.f (. in JO"), You c.n ...",he.. Ihi. "gumen, i. le.ding.•nd it i' ,he re'son we oo"th... "'y ..ber-tooth be.,.., Becau.. "flht ini'ial g'l"" .nglt...quir<d jU" to ckar the c.nin< o"rl. p...ber·'ooth, ".rt...ith . g.p< ..quirement of9O". ",hich incre.... t" 120" foT",m. of the mo.-. d.riv.d nimr.v;d•. For our hypo<li<tic.1..boNooth bear.• g.p< of 90' "'ould re.ult in 'he Ie'" . rm.ngle being por.llol '0 ,h. "n,ion ele-m.n, (muKlel.•nd . In0" i, o. Thi. i••lso ,h. ... son dirk-'ooth•. with g. p....cero ­ing 9(I ' .....d some diffe..n, mUKle m achment point> ju"'0 dos< th.iT mou,h" E"n foT generalized c.rnivo,., with, m.ller g.pe•. the mUKI< fiber< . ... on.nted . t mul"ple. ngle> with multiple "' '''' hment poin, .. The .".nt b.."provide . n mpl. of >orne of 'he ad.pt.tion, thot m"chmotphoJogy ith g.pe.v. " u" foree curv••

If we look .. ,he differen, . kull ,h.p<' of.n om nivorou.brown be.r IU" . , a"'",) compared '0 . trn ly h,rbivorou,p.>nd. (.4i l~",poda mel.Mlt.",) indorsal ,iew, ,he greate"izeof the tempo.-.h, muscul"uTe i• •pparen' in the baCT.How­eve,. the' e i• • g....t.. proport ion. 1...idth in the p.>nd•. Notonly doe. ,he ....mboo di<t'"<lulre. high bit<foree. tho fore•requiTel1\<n' i• ., ocdu.ion. For ••ber -too,h•.• 'arlety of. ddi, ion. 1comprom'''' in tIi<.ttachm,n' poin.. . nd .ngb.... requi d_ T he.. "riation.... .. differen, . nd cornpli.c.t<d n in tho mul"pl. 'ype' of fossil rots . nd c.t-like<.roivo bot ,hey . ...n differen' from e".n, c.,,_ Thecooki< -cu"er ••ber-too,h c.ts requi.. high bi,e fore<. at oc­elu""". "'ithou' los, ofb"e foTC• • t full g' P<. re.ulting in aun ique morphol ogy_

POTENTIAL PREY

Th. ide. of. dirk-tooth .uch •• s..iloJ"" killing ho.lthy full­g, o",n ,\1.In""h.,(m. 10 moth)h., porn laT .ppe.]. but Iial.pr",,'i<. Uty_ There I. I'\CI fossil.,idene<foTtheirh.vingta k<n

ma,ure mammoth, (N. pl. , ...1.. 2000; M.rtin ...1.. 2001~

A, a w.apon for ha"."ing bison. ,ho dirk ,'ooth. • , .mbod·i. d by S..,loJoo, given ' u llic;"n, cover. w.. unm."he<l un,il,ho develop"",n' of the Sh.'P' riA., However, for any proyI. rger ,han amammoth calf. dirl:·tooth .." h.d no def.ultme.n. of inAicting bi« wound•.

E"an, lion •. giv.n .uffiden' "me . nd enough lion •. I:illjuvonile and adul, .I.ph.n" Ooube" . nd Jouberr , t9\l4;19\17) th.. wou ld be beyond ,h. re><:h of S""loJooindividu­ally or in group', Wh .. i. norable abo u' thos••ceou n" i.,h .. ,he: ,,,g...d .nimal h. , to be d.bili",.d (r. duc. d '0,ho beinI:ofcoll.pse)by drou ght Or" ....., ion, Becaus<lion ,are not .bl. '0 use a l:iIIing bile onjuv.nil. 01" larg« .1. ·ph.nt>. they r..emi,lIy c., ,hem 'u death , u.ing a I:illingm..hod more hk< that ofhyen• • and c.nid. (V. n L.w icl:and Goodall. 1971 : Kr uul:, 1972).Thi. m.,hod i...yp i" l fo rf<lids, I' ,horefo« kern' «.>onabl••h.. Ho..", ,,, ,,"", couldhave use<! .imilar ,• • tk> . nd •• I:en <Ven he.lthy ju,'enilemamm",h, by .,,";,ion_ There i. direcl .vide-~ at Fri. ..n·h.hn C" ,. , T. n . (Grah.m, 1976). th .. H. so,... ,,,,,I:calve.oreven juv. nil. m.mmOlh•. • nd m.ny ",h.r .i t.. indicat .( M. rt in <t . 1" 2001 ) juvenil. m.mmo,h p,eJ.. ion by Ho·"olll<ri"",_ How",'''' mat".. m.mmOlh.V«)uld h,v. beenun.uilabl• •• "imiu r ·'ooth prey, .bsent som...vorelydebilitating roodi,ion.

SUMMARY

C." and c..·ULe <arn ivora n, rem.,n . h. ul"m'l< hyper­carn iVOl"<', and, he: . hape of,hei r uppe r c. n i..... defin<•• h.irfunction. l morphology, Each of, he: , h«. b<o.ic rype. of c.­nine morpholo>:i..-eonic.l.' ooth,",imira..,ooth. •nd d i,k­tooth-i. a"""i..od wi,h • unique l:ilIing bi«_ En . n. """i·cal-toot h c' " . ppear >till '0 be u.ing ,h.ir "adi'i"""l, .ndttrt.inly prov.n, bi« and cr ush te<hniqu. , It has carri . dthem .hrough rcpe.«d . >tinction ..... m,.nd <on,in"". '".upport their current .uce.ss, Bite modd.imul.. ion experi­men""" full-.i• • cad... ...h,,'. confirme<lthe ...."ion that..!>cr-tooth. us< a draw -cu, ' 0 produc. wound. wi,h ,h.ioupper ..ni..... _An e,sent i.l fe..ure of, he •• ber-'ooth kiIlingbIle, . h i. d,.,.,-<u, ..,ion i• • •000i.,eJ wi,h .h. cu" . d .n;of the ..!>cr bl.de, in which ,he bl.de movemon' !""lld ' 0,he: ui, f..ili,.,., • •licin~ mov.ment perpendicular.o the

,c".. .. . . ·ro o, .. . .. 0 O''' · 'OO T .. . "C, .. C ." " JJ

bl. de, Sabe<-toot h upperc.nin...« nut ,h.rpcnough 'o<u'. nim. 1hide .nd tissue unIe.. ,he« i. >Ome , ,,od . « d , Iicingmc"'.men, . long ,h. tooth ..i. , E. ch of the 'h .... upper<anine mOfJ'hologi...<hieve. ,hi. diff.rent Iy, Conic.l·,oothbig c."h,,'•• w.ll.devdoped poi nt . nd ..pre. ratio, perm i,­' ing a 'pine-sev. ring n.pe bit. O1".n anchored .u{foe.'ingthro•• d.mp,

S.<imilaNOOIh ca" hite into prey with .n innceted orc" .j.cmry . nd mu" open th.ir mouth, to withdraw or, inthe .ped . 1 Cis< of ,h. cooL;,,-cu l1er COl , fully <x<lud••bladed incisorar<:.de ' 0 cut out of prey, Sdmira"'ooth, use,hei, olton. broad , seroar.d upper c.ni".. to bite down inlin. with the tooth . ... . . they do>< ,heir mou,h,. TIli...­,ion prod",., .n .n,,)' wou nd ,he width of,h. tooth, Ad­dltion . 1injury occur< a. 'he:se ..he" .... ""'i,hd " w,, , Wh.n... I. ..in>: the m.ndible . nd " mulu neou.ly pulling in ,hepo".rior di rection. the se... ted upper c.nin.. furt her en ­la'S• •he wound On , h. ni, mok<_ Mor. d.riv.d "imil. ,...toothc."cou1d. 1", inAi« wound, w"h tlieir ioci>orb<onory.. . d.r.ult bite whcn the pr.y '0'" '00 la'!!. or nth."",i..umuil<d to ,heir .prci.lized Lilling hi«.

Dirk,tooth c.". with Ion>:" and ..l..iv<1y .Iend.., Ie..se" ...d uppe r c. n in•• . use a,,"'her m.tbod to cut rhroughhide- .nd ,i..u• . Tky carve ,hrough prey in . con, i" uou.arc, k«ping the indsor> f... . ., 'hey cu, thei, w.y out, They.pply fortt perpe ndicular to ,h. 'OOIh..i, • • ,h. hite pro ·gre..... pullmg the t"",1i imo a .tieing ore po",";or ro ,he'OOIh..i._Thi . prod",•• • wound « ......1tim.. the . ntero·po"enor w Kith of the tooth. • nd , he: ,,,j.,,ory ' <oult. in ,heuppe r c.nin.. cutting ,hoir w.y out of the prey wi,hout . nyn••d 'oopen the mou,h , Unlil:. ,h. ",imitar·tooth. the dirL·tootli bite w•• .., ,i«- .nd prey -.preific thOl ,hi . <at I.ck. dan . lte rnOl iv. ordef.ul, mean. of..riou.ly wound, ngpreyun,uired ' 0 ,h. ir primary killi" g bite_

The re.,on for Roboc....perimen" in • virtu.l FEAworld i. ,h., comp"'« .n.lyse. tend to reAcet the initi . l.»umpri"". , boIh ••plici' ."d implici'. withou, re. 1worldIomita,iom

Phy'ic.1 • • perimen" not onlypro. ide.m """'" to.he quos·,ion,p<:>kd. bu, f«quondy . uAAC<t het rerqUCOtion._Tb. y,.11you when your . " umption• • re wrong or when e...nti. 1i>­, ue. h.v. he.n overlooked in u" .mbiguou, """yo. .uch ••h.ving tlie experi"",nral b;wn .p«im.n lift off ,h. ground_

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