ON A COLt;B1U D w nll VE IlTl CALL Y ~IO VAI3 L E l\IA X I LLA .

83

7. On a ('oln hrid Snake (~rl:'Jlodon) with a v('rt i (:~,lly Ill oyable

)la xi llary Banc. By E. G. B On L}~ NG EB , F .Z.;;:; ., Curato r
oC Heptiles.
[ Hecein)d No\'clllllc!' 10, l!) I'l : Bead F ebruary H, 1915. J

(Text-fig m e 1.)
lsO E X.

StructUl'e or ) fOl" pitolog.\. i\la xi ltlrY bOlle ill Sua kl's .. .

Physiology. h uportance of physiolog ical action of thc
vcnolll s of Snakes Oil "li\!'siticatioll ... .. . ... .. . . .. ... . .. .. . ...
jE tiology . E I'olutioll of t he maxillary bones ill SHllkeS.. .

I'age
83

83
85

In Vip81's t he ll1ax ilhil'Y hones, t o wh id ! t he poi:son -fnngs 1\ 1'13'

fi r mly )'I,Ltached, are mova.bl), 1l.1tic ula t ed to the pl"ch 'olltals all(l
ectopterygo ids, th e po i ~o n -ffUlgs bei ng, when n.t I'est, fold ed
Hga inst th e roof of the mouth a nd becoming el'ected, 01' eyen
t hrllst for wa rd, when the nni mal is a bout to ~k i k c . T his \' l"t ica l mobili ty of t he nmx ilb ry bon e, w hich gi ves t h e~e sna kes
s tlc h a mechfln ical ;)Ch ':lntage whe .. th ey ;)I'C about to ~ tri ke , h<1."
:dways bee ll leg:-l.Ided ) 15 e:sse ntia ll y chamcteri &t ic of t he members
of t he fami ly Vipel"i che. T he Society I"ecell t ly I'ece ived fr om
1\[ 1". 'vV. A . S mi t he n :, O. 1\-1.Z.S ., a genel"OlI s don ol' to iL:s coll ect ion ?
a specim en or X elwdon mel'l"emi, a n ngly photl on t co lubrid inhabi t ing Brazil a nd Pa l"ag ua,y ~ wh ich is chal":1.ct lized by a n
ex tremely shor t l11:lx ill;wy with on ly six Ol" .<;e\en teeth , followe d
a fter fi n intel'space by:1. pail' of stl'ong l), en l:lI"ged but likewise
sol id , ungrooved fa ngs . On ta king t he snake from t he box in
whi ch it was packed and cfi tch ing hold of it be hi nd t he head,
1 wa s most s Lll" pl'ised to see the Cl'eatUl'e, on ope nin g its mouth
in a n a ttempt to bite, er ect nnd de press i ts Ltngs in a t horough ly
Viper ine ma,nlle r . FU It lle louSe l"vationss ll owe,1 tha t th e mobili ty
of its nUl. xil ln. \\";l S so grea t t ha t t he fa ngs co uld be not mer ely
eI 'ectec1, but t hrust fonv:t rcl an d sideways, r e\'en ling the fact t ha t
t he mecha,n ism i n t his sna ke is mOI'e per fect t han in it la rge
lI um be l' of V iper s of silll ilOl' size.
Th is d iseovel'y of a. solid toothed Oolubr itl wit h a vertically
movabl e maxi lla. is of s peci:1 1 inter est, as I thin k it goes a long
way to w:u ds settling t he l)l"oblem, so often d iscus:sed , of t he
der iv:l,t ion of t he Viper ine max ill a ry bone. Th e V ipel'idre were
form erly bel ieved t o hnve spr ung from t he .P l"otelog ly ph Oolu l)I" id:s. I n t he Ca t..'lJog ll e of t ile BIitish Mu:se ulll , published in
18!J;3, m'y fa.thcl, G. A . Houl enger, F.R.S., exp ressed t he opini on
t hat the poison a ppa l:"1.t u.s of t be Vipe rs was in all proba bili ty
del'ived fl'om t he Opisth ogly phs. L ater , in a pa pel' p ublished in
th e :P Ioceedi ngs of thi s Society, h e poin ted out tiwt, from t he
A gly ph odont form s in wh ich the teeth iucl"ea.:': c ill :-.ize po~tel' i orly,
we a re gl"1l.<J uft ll y 1Nl to t he Opisthog ly phs, wllich Cftll be (li fier ent iated only by th e presence of 1l 10l'e 0 1" le!-i.s d eep gl'oo\,es on t he

6"

84

Mn. . 1::, G. BOL-U;XGEH

O~

posteriOl' fang-li ke teeth, the sCI"ies (' u\mill:-1ting in such fOl'ln s as

show th e maxilla IT bone much :"lbl)!'cvi:Lted. the soii cl t eeth \'0du ced to two Ol' till"ee on ly, nnd t il e fnllgs extre mely large and
gl'oo\ell.
Th o htest co nt l'i butioll to the .':,ubj ect is one by

1\11-. J oh n H ew itt *, who nttempLs to s IIOW Lh rlt th e Vipel'ida: :1,1'0

not of Opi :-.thoglyph :'\,ncestry, but, ::lT e 111 0 1'e ('losely r elated to t he
P l'otel'ogly phs. The most impOl'L:l ll t Hl'gu llJent foi he uses to establi sh hi s point fIt'O that, in t he fll"~t plnec , in the Opi sth ogly ph ~ the
Text-figu r e I.

orb.

prf

mp!.

epl.
A

pr/:

...... plf

rna::(

s
:'! l axill:l I'Y (111,1'.) of X ellQdOIl merJ'emi at. rest (A), :lIld erected ( 11).

ept., cctOJllcr,\'goid: orb ., orbit; l)rj:, preft'onlal; pif-, postfroutal.

fallg- ben,r in g po],tion of the mn.xilh is sitllnted f~l' hehind t he

preft'on ta l, find cO ll ~ eqllentJy tk"t :1 S t h er e :-tppefll's t o be no
t enden cy :lmongst Opisth oglyphs fo]' a. fOl'\vnrd movement of the
fang-bear in g pOl,t ion , it is difti cult to conceiwl how the evolu t ion
of t he Vipel'ine charllcter commenced; secondly, t hat in th e
Protel'oglypils t he fang-ben l'in g por tion of the ma xilla is some.. Ann, Trflll.':'\-flal

i\ l ll~ ,

iii , 1911.

CO LUUR I I) WIT II VERTl(JALLY MOVABLE MAXII, r,A,

85

whn t enlnrged, often in a ve l 'tic~11 tli l'cctioll, showi ng a l'esenlblnnce

to t he stat(' of t hing:>; found in (/(f.,U8lt8 .
Th e fil';..t a rgument is ell:;; il y di:-.posed of, as in :t l1U mhel' of
Opisthoglyph:-. the fnng1'l :u 'e situ:tterl just helow t he pl'erl on tal
(J/ iodon, Pole1l1ou , B /'(u:liyophis), ~ow ill Xellodon the pOI-tion
of nHlx illn, herll'iug the f:lllg-li ke teeth will be found to he much
enhu'ged, a nd in a 1I10l'e 0" I'ss \'eltical di,'eelioll. :llH l it Olliy
I'e mn in s fO l' the last two teeth . to be flll'lli shed wit h gl'oo\'es to
t l'an :-.fol'm X enodoll into an Opi:-. t h ogl~ ]Jh with the fangs f.. itllnted
helow thl' pl'erront)\. l?ul'the,'; we ha,'e on I," to compnre the
maxillft of X eJIOdOIl with t ha t of the least specia.li zecl of the
Vipers, CCW81tS. to see tllat mel'ely fI sl ight t ilting I1p of the
m::t,xilh of the f01'J)I(,I' snnke, with the lo ~s of t he few fr ont
teeth and n vcry slight mor/iucation of th e hone, i,... neede,l to
bl' itlg ;dJOllt:t conditiion ~ im i lnl' hoth in stl'lu.:tUl'e altd me('h[t ni ~ m
to that of Vi PC"8, Thu s _fenodon with it ~ "el'ticnHy nloynble
lIHL x iH~c elHl,bl es us to trace t he pl'Obalde e"ol l1t,ion of th is bone,
and th e old view, recen t ly j,(>\,j" ecl. that Yipe t'$ ;u'e d e~cclHled
from P t'otCl'ogly ph s nt lJ ,..,t, in my opinion, be <lbnll(loned,
1\(", Hewitt in his papCl's states th;1 ~ the v:lriolls experim e nts
on snak C-yelH)lll see m to show t hflt t here is more in common
between t he Pl'oteroglyplls and the Vipers; , than between t he
Opisthoglyph s awl cithel' of the othel' diyj::;ion.:-. That this i.::i so
in th e ma.jo rity of C:ls;es h:1s bee n delll onsb'n tecl oy P h isnlix, It
should be hom e ill mind, 110\\'e,-el', t hftt. flS has I'ece ntly bee n
showll by l?itz:,;iHlolt s, the poison of the most h igh ly ,'enomOlls
Opist hoglyph , Displwli(llls typus, i n its phys iologicnl fiction is
p<ll'ticulndy cJlftrnctel'ist,ic of that of the South Ameri can Vipers
of the gen us Lachesis. The physiologicft l nction of the " enom s
can, t herefol'e, h:we little impo ltallce in the settlement of the
bl'onder })l'oblem of' t he class ification of Snakes fl'om the point of
view of descent.