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796

B. ButT-3m a. a.: Line morphologiscllO und fnnktionelle Analyse

F.: Uher die \Terteilnng der spongiosadiehte im coxalen Femurende und ibre Becloo.
tung fr (lie Lehre vom funktionellen Ban des Knochens. Morph. Jb. 95 (1954) 3554.
Gesammelte Abbandlungen zur ftmktionellen Anatomic des BewegungsaPParates. Berlin: Springer 1965.
Atlas zur Biomechanik der gesunden nnd kranken Hfte. Berlin: Springer 1973.
mk cler Organismen. 2 Bde.,
Roux, W.: Gesammelte AbhandlUngen fiber ieldtmg5me
Leipzig: Engelmanfl 1895.
SCHMITT, H. P.: Uber die Beziehung zwischen Dicbte und Festigkeit des Knoelrnns am Beiepiel
des mensebbeben Femur. Z. Anat. Entwicklgm 127 (1968) 124.
Smuco, K.: Uber die Hftpfanne. I. Mitt. Z. Morph. Anthrop. 17 (1915) 325356.
TmbMANN, B.: Die Beansprueltung des mensebliehea Huftgelenks. Hf. Die Form der Faeies
lunata. Z. Anat. Rntwickl.gesCll. 128 (1969) 329349.
Zur Lokalisation von clegenerativea eranderungen am Femurkopf bei der coxartlirose. Z.

PAUWELS,

Gegenbaurs morph. Jalirb., Leip


zig 125 (1979) 6, 5. 797817

Aus dern Xoturhitorischen Muse


urn Easel, Zoologisclee Abteilung

Orthop. 111 (1973) 2327.


T: Das Beek
en. Bonn: Cohen 1899.
WttuEt, H.: Die Dicko der mensebliehen GelenkknOrPel. Mccl. Diss. Univ. Berlin 1897.

A Functional Interpretation of the Var


anid Dentition
(Reptilia Lacertilia, Varanidae)

WALDEYEE,

Oliver Rieppel

i)r. EArNER, Batun,


Dr. WALTER OBERLANDER nod

Dr.

I
With 14 figures and 2 tables

HnNz.JUReEN KURRAT

Anatomiscbes Institut
der Univereitt zu Koln,
Lindenburg
D 5000 Kbln 41

Received for publication 2. February


1979
aad in revised form 11. April 1979

A. Introduclion

Cranial kinesis in Modern lizards is a


long standing and much debated prob
lem in
the area of functional anatomy. Clas
sical contributions are those of BuADLE
Y (1903)
and Vnstuxs (1912). More recen
tly, FRAZZETTA (1962) renewed the
interest in the
functin of the lizard skull with
the introduction of the quadric cran
k chain as an
explanatory model. IORDANSKY
(1966, 1970) and Gostus and GA5
C
(197
3) fnrther
elaborated the ideas put forward
by FRAzzErn (1962). All of these mod
els, as well
as

the one presented in the prese


nt contribution, still have to he teste
d by clectro
myography. The only electromyographic
data on the head muscles of lizards
avail
able to date are those prese
nted by TumoelniouroN (1978) on
the
m.
ptery
goid
eus
and vi. depressor mandthulae of
Uromaslix.
The adaptive significance of cran
ial kinesis has been discussed by FRA
zZETTA (1962),
IOflDNsuy (1966) and THo
IsoN (1967), but analyses of feed
ing
lizar
ds are rare.
PRAzZETTA (1962) described
feeding in Gerrhoaotus. and TilnocKl
uonr
oN
(1976) pre
Seated a einradiographie analy
sis of feeding in Iguana and Uror
nasti
x,
two herbi
Vorous lizards. The only published
data on feeding in T7aranus are
the
few obser
vations made by BOLTT and Ewu
n (1964). A detailed cinematog
raphic analysis of
feeding hi Fan ens has been
attempted by hIPEY (1967). His
still unpublished ac
I toat differs in impo
rtant respects from the observations
presented here.
The present study reports cinCradiograp
hic analysis of feeding in Papa nus.
The
movements in the head of Paranus
are related to the function of the dent
ition
. It
Wifi be argued
that amphikiaesis is correlated with the
proper functioning of the
beth in T7aranus, which is critical for
successfully seizing and handling large
prey.

798

0. Riurrun: A Functional Interpretation of the Varanicl Dentition

0. Rirppn: A Functionai
Interpretation of the Varanid
Dentition

Varanus is a lizard genus adapted to relatively large prey. This is reflected in the me
chanics of its head. Consequently, no generalisations will be attempted concerning
the amphikinesis among Modern lizards from a mechanical point of view.
The study reported on here is part of a thesis presented to the Faculty of Natural
Sciences at the University of Basel in partial fulfillment of the requirements for the
degree of a Dr. phil.

799

B. Material and Methods


For the study of cranial kinesis on Vurenus, four live specimens of ]7 bengolensis and one
specimen of J7 exenthemeticsis elbiguleris wore available. In addition, observations could be made
on live V. komoclocesis held at the Zoo Basel.
Cinematography was clone with a Canon 814 camera using Kodachrome super 8 movie film,
and with a Baulieu 14-16 E]ectronix camera using Kodak 4-X reversal film 7277 (16 mm). (mi.
radiography was clone at the Kantonsspital Basel using Kodak film RAE 2498 (35 mm). The
analysis of the X-ray film was clone with a Tagarne-35 analyser.
For their use in stress analysis photoelastic plates were made out of Araldite B. The plates
were molded to the thickness of 4mm at a temperature of 120 C. The analysis of isochromatic
fringes and of isoclinics was clone by means of a simple polaroid analyser as described by MNcH
and Forra (1959) at the (ii a-Geigy 0 Brnl.
The apprec cli to the ctructure of tI e head of Voranus is based on the following mueuio spe
cimens. Specimens marked by an osterik have been dissected, the others are dried skulls.
Vei-onns ecomithuros (FNNH 98935); V. beegeleeois (three imeatalegued specimens555, MBS
13 162*; V. dutserilii (1\lBS 3721*); V. duwerilii heteropholis (FMNH 1457]]); V. exoothcosaticus
olbigehois (one uocatalogued specimen5, MBS 5819, FMNH 51683, 17142,17143); V. cranthe
rnotirus microstictvs (MCZ 47369, 47375); J7 gilleni (BMNH 1910.5.28.13); V. geuldi (FMNH
31338,31340, 51706, MCZ 33916); J7 prisms (MBS 5819*, BMNH 1974.2481, 1974.2482; FMNH
3] 308, 51705); V. kosnodeeesis (BMNH 1934.9.2.1., FM1H 22200); V. iodicus (MBS 11099*);
V. vehnlesns (BMNH 1931.1.12.1); T7. vileticus (MBS. 3729*, ]3453*, ]9642, SMF 33251, 33252,

Fig. 1. The application of


the quadric crank chain
model (FRAzzETTA 1962) to
skull of Varonus se/veto,.
the
Abbreviations: a, angular; ar,
articular; bs, basisphenoid;
c, eoronoid; d, dentary; ep,
epipterygoid; f, frontal; hy.j, hypokinetic
joint; ju, jugal;
I, lacrimal; mes.j, mesokinetie
joint; mx maxifla; a, nasal;
p, parietal; pmx, premaxihla;
p0, postorhitofro. pot,
prootie; prf, prefrontal; ps,
parasphenoid; pt, pterygoid;
q, quadrate; Sm, septomaxilla; sq,
squamos; st, supratemporal
(From Riuppan. 1978,
by permission of the Birkhauser
Verlag, Basci)

F5]H 17145, 22084, MCZ 1066); T. eec/lotus (FMNH 22354); V. presinus (MBS 10 504*); V. re
dice//is (MBS 18874, FMNH 98947, 131538); J7 so/voter (MBS 3713*, 3736*, 6138*, 10755*,
osteol. 5028, 5878); V. stone (MBS 17891*); V. venus (AMNH 28698, 73361, FMNH 1492).
Abbreviations: AMNH. American Museum of Natural History, New York; BMNH British Muscum of Natural History, London; FMNH, Field Museum of Natural History, Chicago; MBS,
Natural History Museum,Basel, osteol, = osteological collection; MCZ Museum of Comparatie
Zoology, Cambridge, Mass.

C. The mechanical nnits of the sknll of Varanus

The decomposition of the kinetic lizard skull into various functional components
goes back to VER5LUY5 (1912). He subdivided the lizard skull into the oeeipfl
segment and into the maxillary segment which represent the braincase and the
dermatocranium respectively. The braincase is movably suspended within the des
matocranium on its paroccipital processes posterodorsally and on its basipterygotd
processes anteroventrally (Fig. 1). As the maxillary segment moves up and down
around the braincase, the cartilaginous processus ascendens which projects from the
anteromesial tip of the supraoccipital bone slides back and forth in a trough on tIlE
caudal edge of the parietal. This is the metakinetic joint.
The metakinetic joint is regularly fused up through the development of a sutil&
contact between the front edge of the supraoccipital and the hind edge of the par1e

bI in large (adult) speciauens


of Varanus exanthematicus, T.
niloticus and V. korno
doen sir. The same apparently
occurred in the extinct giant varanid
Megalanja from
the Pleistocene of Australia
(HEcHr, 1975). The obliteration
of
the
metakinetic
appears to be size dependent,
joint
but the functional reason of its
obliteration is not
Understood
Vnsys (1912) recognised
a further joint within the
derluatocranium of Varanus
located between the frontal
and the parietal bones, which he
called the mesokinetic
oit This system has
been more fully explored by
FRAzzETTA
(1962) who showed
mesk5 to eharacterise
the great majority of Modern
lizards.
The lack of meso
kinesis seems to be due to
secondary fusion or interlocking
of
the bones of the der
atocraniuiu As FuAzzErrA (1962)
realised, true iuesokinesis not
only involves the
POssibility of flexion between
frontal and parietal bones, but
also involves a hypo
kinetic joint within the
palate (Fig. 1, hy.j.). The
combination of the mesokinetic
and hypokinetic joints
allows the luuzzle unit to rotate
dorsally and ventrally relative
to the rest of
the dermatocranimu
The muzzle unit or the snout
complex of Varanus consists of the
mesial premaxilla
and the fused nasals
and of the paired maxillae, septomaxillae,
jugals, lacrimals,
Prefrontals, vomers and
palatines (Fig. 2). These bones move as a
unit
relative to the
st of the dermatoeraniusu
The dorsal hinge is located at the
frontoparietal
The latter represents
sutuse.
a more or less straight transverse line with
slight interdigitation
y (Fig. 3). The sutural plane
is oblique, in that the lower edge of the
hind end of

800

0.

thErruL:

A functional Interpretation of the Varenid Dentition

0. RIErPEL: A Fnnetional Interprotation of the


Varanid Dentition

801
pterygoids. Caudally the quadrates (quadrate
units) link the parietal unit to the
basal units. The quadrates receive the caudal
tips of the supratemporals and of the
squamosals in two collateral, shallow ai1ic
ular facets on the dorsal surfaces
of their
cephalic condyles. The epipterygoid bone is
the last component of the maxillary
segment of VER5LUY5 (1912). The foot of the
epipterygoid rests in the pterygoid or
coluuxellar fossa on the dorsal surface of the
pterygoid. The dorsal head of the epi
pterygoid is bound to the alar process of
the prootie and to the deseensus parie
talis
by fibrous ligaments.
B. The mandibular joint

The mandibular joint of laranns is a com


plex structure (Fig. 4) in order to allow
rotational movements both in the vertical
and in the transverse plane. The glen
oid
surface of the articular bone of the lower
jaw is concave from front to back,
but
it
is convex from mesially to laterally. It show
s a saddle-shaped surface. The mandibu
lar
condyle of the quadrate bears a mesial and
a lateral convex head with a longitudinally
oriented cavity between the two heads to
match the convexity of the glenoid surfa
ce
of the artieular. The mesial side of the
glenoid surface receiving the ouesial
man
di
bular coudyle of the quadrate lies deeper
than the lateral part of the glenoid surfa
ce
receiving the lateral mandibular eomlyle.

ju

Fig. 2. The snout complex of T7eraans bciiyeleiisis (MBS. 2410). Key to abbr,viatioflh
as in Fig. 1. Scale equals 10 mm

par.c

gI.s

Fig. 3. The mosokinetic joint of Uoron?8 solcotor (MBS. 5028)


Key to abbreviationS as in Fig. 1. Scale equc Is 10 mm

the
the frontal lies below the upper edge of the front end of the parietal. Towards
art
lateral edges of the suture, the parietal forms distinct anterior lappets which
eulate in lateral notches on the hind edge of the frontal. The ventral hinge, the hypo
lieS
kinetic joint, consists of a lateral and a mesial component. The mesial cmupoildhlt
the
een
betu
between the palatine and the pterygoid. The lateral component lies
the
ectop
Of
teryg
oid
maxifia and jugal of the muzzle unit on the one hand and the
basal unit (see below) on the other hand.
(1962)
The maxillary segment is broken down into five components by FnazZETTk
unit
The muzzle unit described above links the parietal unit dorsally to the basal
11toit0
ventrally. The parietal unit is made up from the parietal, supratemPorals
ecto
frontals and squaiuosals. The basal units are made up from the ptcrygoids and

I mc

S.

o
n
a
r
t
m
C

Fig. 4. The mandibular joint of Varo


ans solcotor (MBS. 3713). A. The hind end of the
right lower jaw in dorsal view. B. The land
end of the right lower jaw in mesial view.
C. The mandibular condyle of the quadrate in vent
ral view. Abbreviations: elit, cliorda
tvmparn forainen ; gl.s, glenoid surface
eular; line, lateral mandibular cond
yle
mine, mesial mandibular condyle; pare, prcartieular cend
yle ; rap, retrearticular pro
cess. Scale equals 5
0Jrpli. lb. 125/0

0. Riurran: A Functional Interpretation of the Varanid Dentition

802

The structure of the mandibular joint results in a lateral rotation of the lower jaw
around its long axis as it is depressed and in a mesial rotation around the long axis
as the lower jaw is elevated. Such a rotation of the lower was proposed by BRADLEY
(1903) on the basis of similar arguments.
When the jaws are fully adducted the dentary (as well as the maxillary) teeth pohit
slightly inwards. As the jaw is depressed relative to the long axis of the quadrate it
rotates around the dceper mesial side of the glenoid surface: the teeth are rtated
outward. They thus assume a fully erect position so that they hit the prey with their
tips instead of with the lateral surfaces of their crowns. Convcrsly, as the jaw is ele
vated relative to the iong axis of the quadrate, it rotates aronnd the higher lateral
side of the glenoid surface. Consequently the teeth rotate back inward again and
thus produce a clamping effect on the prey held between the jaws.
The second feature of the mandibular joint which has to he emphasised is the strong
anterior elevation of the glcnoid surface on the articular bone. Anteromesial to the
glenoid surface the surangular is also elevated and forms a prearticular condyle. Thc
anterior elevation of the glenoid surface provides an increase of area on which the
lower jaw can he rotated during depression (TmwcK3ionToN, 1976). But it also
locks the lower jaw firmly against the mandibular condyle of the quadrate whea
retractive forces are transmitted from the lower jaw through the mandibular joint
to the quadrate. As it will be shown below, the depression of the mnzzle unit is func
tionally important during the swallowing process in Varanus. Depression of the
muzzle unit results from retraction of the basal unit which can be achieved by back
w-ard rotation of the quadrate around its dorsal suspensioii. Jaw adductor muscles
which insert obliquely into the lower jaw will exert a force which can be broken down
and into a
into an adduetive component working at a right angle to the lower
jaw.
retractive component which works along the long axis of the lower
jaw

F. The Quadric Crank Chain Model

0. Rizp: A Functjonaj interpretation


of the Varanid Dent itjon

803
and the quadrate. The small amount of
translational movement possible results
from
the elasticity of the ligaments of the
joint capsule.
F. The Analysis of Feeding Movements in
Varanus bengaknsjs
The present account deals with Varanus
Sen galensis of which four live specimens
were available for study. The animals
were regularly fed on mice. Five
feeding se
quences were filmed with an ordinary
super-S movie cassette at a speed of 40
frames
per second. Two further feeding
sequences were recorded on a 16 mm
black and
I white film at the speed of 64 frames
per second. An eighth feeding sequence
was
filmed with an X-ray camera at a
speed of 48 frames per second. Cursorial
obser
vations were also made on one specimen
of Varanus exantheniat ices albigularis
and
on one adult specimen of Varanus komodoensis.
When Farce us bengaiensi.s locates prey, it
immediately starts to pursue it, seizing
it wherever possible without any
preference for any particular part of the
body. The
pi.ey is vigorously shaken and
banged against the substrate, rocks,
logs,
etc. until
it is immobilized or dead. The
JTaran mar then usually drops the prey and
starts
searcli
iag for the preys head. It has been
shown by Loop (1974) that varanids
feeding on
small mammals show a signifieamrt
preference to swallow- the prey head first.
Once
the Varaijus has started to swallow- its
prey, kinetic inertial feeding (GAN5,
1961,
1969) is observed. thuS (1961:
218-219) describes kinetic inertial feeding
as fol
lows: The jaw-s release the prey
quite suddenly and the entire head shifts
forward
and laterally, then bites in a newposition. The inertia of the prey reduces
prey
shifting. The head continues to sluft
from side to side in a w-alking sequence
that
graduajly produces ingestion of
the prey
The sequence results in a pushing of
the head over the prey rather
than drawring the prey into the month.
GANs (1961)
i5 FRAZZEYn who noted that
the lateral excursion of the forward
stroke of the
head does not occur in all lizards
during kinetic inertial feeding. In Varanus
benga
hens the lateral excursion of the
head occurs only occasionally It may also
be
noted
that Jaranns bee gaiensjs combines
the kinetic inertial feeding method witli
the
pos
Sibihty to draw- or rather to push the
prey back into the throat through jaw
mobility
(see belowr)
-

(1962) introduced the model of a kinematic chain which was further


occur
elaborated by IORDANSKY (1966, 1970) to describe the movements which
The
feeding.
within the maxillary segment of an amphikinetie lizard skull during
kine
A
laeertilian dermatoeranium would represent a quadrie crank chain (Fig. 1).
Fig. 5 shows outline tracings of
one kinetic inertial feeding cycle from the
link
one
if
that
way
such
in
a
X-ray
together
links
jointed
of
assembly
is
chain
an
matic
ifim of Vat-anus bengale,i.sjs and
w-ill
serve as basis for the description of the
(FRAzZErT&
way
predictable
head
in
a
moves
chain
the
whole
is fixed and one is moved,
movements as they occur during one
cycle of jaw opening and closure (one
together
jointed
orbit).
links
four
of
1966; ALEXANDER, 1968). A quadrie crank chain consists
1o frame 1, the animal
stands on its erect fore-limbs and holds the prey
betw-een its
Jaws off the ground. Mouth
by hinge joints the axes of which are parallel to each other.
opening lasts until frame 40. The head is rotated dorsally
un1
mozzle
around the oeeipitoatlantal
In the skull of Varanus, the four links would be represented by the
joint, and simultaneously the muzzle unit is
link)
(eaudal
elevated.
unit
quadrate
the
link),
by
(dorsal
unit
parietal
the
(anterior link), by
can be seen from frames 3 to 9, the
it
muzzle starts to he elevated before any rota
chain
crank
quadrie
the
of
function
the
on of the head around the
and by the basal unit (ventral link). For
occipital condyle takes place. On frame 9, the maxillary
tran
beth are cleared off the prey.
is critical that all the four joints involved be hinge joints, which do not allow
Therefore, muzzle elevation alone w-itliout any moveEx
I %t
lational movements. The critical point here is the quadrate-pterygoid contact.
of
the
head
or
of
the
mandible
is sufficient to free the maxillary teeth from the
WCTh Ic
Prey during the initial
amination of dissected heads of Varanus where all tendons and ligaments
phases of the orbit. The muzzle is further elevated during
pterygd
the
i ntaet showed that very little translational movement occurs between the
1Pward rotation of the head around the occipital eondyle,
and it reaches its most
$2
FRAZZETTA

0. Rixrzxi: A Functional Interpretation of the

804

Vara.nid

Dontition

0. RiEpp: A Functional Interpretation of the Varanid Dentition

805

48

N--

..-:

A
16

7/

Fig. 6. The cervical vertebrae (atlas and axis) of A: Betoderma s.


81594) and B: Iaranus griseus (MBS. 5819). Scale ecjuals mm

50

--

8Uspectwrn

(MB$.

20

N
A

N
-1
/7A

Fig. 5. Outline tracings of a feeding sequence of Verne us bengatensis recorded by cine


radiography at a speed of 48 frames a second

extremely elevated position on frame 16, that is before the mouth is maximally
opened. Maximal mouth opening is achieved on frame 40. It is essential to note that
the
opening of the mouth is brought about by upward rotation of the head around
prey.
the
occipito-atlantal joint. The mandible remains stationary and supports
As the mouth is maximally opened, the forward stroke of the head characteTiSt1
an
of the kinetic inertial feeding follows. Initially, the head moves very rapidly in
across
anterovesitral direction. The gape is held constant and the head now shifts
phase
directed
a
is
there
horizontally
the prey (Fig. 5 B). Following this movement
oCCI
the
around
of the forward stroke of the head. The head now rotates downward
gape
the
pita.l conthle, and simultaneously the snout is depressed. This closure of
the
of
brings the maxillary teeth into contact with the prey again. The final phase
snoUt
forward stroke of the head is directed anteroventrally again. The tip of the
around
now closely approaches the substrate. The head is strollgly rotated downward
strong
This
the occipito-atlantal joint and the muzzle unit is being strongly depressed.
and
mouth
and final depression of the muzzle unit pushes the prey further into the
down the throat.

99
\
\
\

\
\

Fig. 7. The application of the quadrie crank chain model to illustrate the movements
as they occur in the head of Varanus ben galensis during feeding

806

0.

RtmrEL: A Functional Interpretation of the Varanid Dentition

Varanus bengalensis does not perform a kinetic inertial feeding cycle with each
down the throat is shown. Tho
orbit. Often a different method of pushing the
animal presses the prey against the substrate. It opens the mouth, pushes hard against
the prey and closes the jaws tightly again, thereby depressing the muzzle unit.
During the kinetic inertial cycle, it is advantageous for the animal not only to
maximally accelerate the head but also the maximize the excursion of the head
during the forward stroke. The longer the excursion is, the more is the head likely to
shift across the prey. Varanid lizards, which are adapted to relatively large prey,
show an elongated neck which results from an increase in number of cervical verte
brae and from an increase in length of the individual cervical vertebrae (HOFE5TETTER
and GAse. 1969. and Fig. 6).
Fig. 7 shows the applieaioii of FRAzzETTAs (1962) quadrie crank chain model to
the analysis of another kinetic inertial feeding sequence of Varanus bengalensis as
it is observed on the X-ray film. The model as it is drawn in Fig. 7 does not correspond
to the actual movements of the head which are generated in the neck region. Rather
it is used in order to allow measurement of degrees of rotation. Joint X is considered
to be the fixed reference point within the quadrie crank chain (FAZzETTA, 1962),
and for the sake of easy comparison, the parietal unit is always drawn in a horizontal
position. The analysis shows that starting from the rest position (Fig. 7 A: frame 1)
the muzzle unit is elevated for 9 while the mouth is maximally opened. The qua
drate is protra etc ci for 210. The angle between the basal unit of the skull and the
dentary is 48 at the stage of maximal jaw opening.
Following the forward stroke of the head, the skull is rotated downward around
the oeeipito-atlantal joint while the muzzle unit is depressed. At the stage of maximal
150
depression of the muzzle unit (Fig. 7 D: frame 99) the latter has rotated around
compared to the stage of maximal muzzle elevation: thus the muzzle is 6 lower than
it is at the rest position. Correspondingly, the quadrate has rotated backward around
27 compared with the maximally protracted stage. This is 6 further backward than
it is at rest position. The analysis shows that during the final stages of each orbit,
the snout is depressed below its rest position, and that this final muzzle depression

0. Risppn: A Functional Interpietation of the


Varanid Dentition

807

prey

pushes the prey further down the throat.


G. The Functional Interpretation of Varanid Dentition

The preeeeding description revealed some implications of cranial kinesis with


respect to feeding (jaw movements as related to food intake). The following sections
will be clealmg with the adaptive significance which cranial kinesis may have in rela
tion to the structure anti function of the clentition.
Jcu-anus is adapted to relatively large prey. The functional significance of tooth
shape has been discussed in detail by FEAzzETTA (1966). What is critical in sueeeSs
lts
fully ;eizing and handling prey is not only the absolute size of the tooth but also
stronglY
degree of reeurvature (Fig. 8). The two parameters may be compensatory. A
ab
reeurved tooth will have the same or even the better holding effect even if it is
a
0f
solutely shorter than a straight tooth. Or, to put it in other words, reeurvature

/--------Fig. 8. Prey penetrated by a straight


(solid) and a recorved tdotted) tooth. To free
itself, the prey must deform its surfhco
or rupture it to the depth h. The straight
and the recurved tooth penetrate to the
same depth Ii, although the recurvedi tooth
is
absolutely shorter by ci than the straight
tooth,

tooth allows the reduction of the latters


length while maintaining the same holding
effect. At a given maximal gape, the shorter
tooth will interfere less with the uptake
of relatively large prey.
Reeurvature of a tooth may be subject to some
compromise, however. The tooth
hits the prey most effectively if it hits the
prey with its tip. Force is then transmitted
through a minimal area which results in maximal
stress on the preys surface. Pene
tration of the preys skin is then most likely
to occur. But to erect a strongly recurved
tooth enough so that it hits the prey with
its tip wUl require a kinetic skull or a very
wide gape.

I II. The Ontogeny of the Varanid Dentition


The tooth development in Taranus has
been described in great detail by BULLET
(1942). The replaeemeiit tooth grows in
an interdental position and migrates antero
hterallv when the preceeding functional
tooth is shed. During the ontogenetie
development the pulp cavity of the tooth
beeemes subdivided by dentine infolding.
The development of such plieidentine
is characteristic of T7aranus and of all
other
Platynota lizards. In the adult tooth,
plieidentine results in the subdivision of the
Pulp cavity through a complex system
of nnastomosg dentine lamellae (Fig.
9, A,
I E). As the replacement
tooth moves into its functional site, the
surface
of
the
tooth
bearing hone forms a system of irregulat
anastomosing ridges. To these ridges the
bases of the dentine lamellae of the
plieidentine become ankylosed through the
bone of attachment (PooLE. 1967). When
the functional tooth is to be shed, the
bone of attachment and the dentine just
above it are removed through resorption.
In Varann.s resorption has to occur sirnultaneousl3,
all along the base of each dentine
amelIa0 of the plieidentine. Only towards final
stages of tooth replacement do large
lranuloeytes attack the surface of the dentine
mantle low down on the tooth base
T aranus. But as the pulp cavity is subdivided
by dentine lamellae. no resorption
Pits develop comparable to those of
other lizards. In most lizards, the replacement
th move into resorption pits antI come up
from below the preeceding functional

I-

808

0. Rrurnn: A Functional Interpretation of the Varanid Dentition


.

I;

A Functional Interpretation of the


Varanid Dentition

809

/1/

4!
I

/
/

4
/

L
I
4.

Fig. 10. Succossive growth stage


s of replacement teeth of the 0th
tooth family on the
right maxilla of Veranus selm
tor (MBS. 0138). The functional tooth
is to the heft. The
scale equals 1 mm

a convex surface. The tip of the tooth


is concave mesially: it is bent in a lingu
al di
reetion.
In lateral view, the tooth is recurved
. Ontogenetic growth series (Fig.
10) show
that the replacement teeth are ahuo
st symmetrical during the earliest stage
s of cal
cification. The recurvature of the tip
of the tooth is more pronounced durin
g later
stages of ontogeny than it will be in
the functional tooth, however. App
aren
tly,
the
postrior cutting edge grows faster
than the anterior cutting edge shortly
before the
tooth moves into its functional site.
This developmental pattern leads
to the for
mation of a biconvcx shaft on whic
h the reeurved tip is observed.
As it was noted by MERTEN5 (194
2) already, the degree of reeurvature
changes
along the tooth row. To obtain som
e objective statements on the degree
of tooth
I
reeurvature, the circumferential
line was constructed which represents
the axis of
the tooth in lateral projection. Then
the tangential line was constructed
on this axis
through the tip of the tooth. Thus
the angle could be measured at whic
1
h
this
tangen
tial hile intersects the long axis
of the tooth bearing bone. The resul
t
of
a
serie
s of
maxillae and dentaries of T7aranus salva
tor representing various growth stage
s are
1 given in table 1 and 2. Recurvature increases towards the rear of the tooth row both
in the upper and lower quadrants (Fig
. 11). Reeurvature increases relatively
1 in the maxillary
more
tooth row, and the maxillary teeth
are absolutely more strongly
reeru-ved than the dentary teeth.

Fig. 9. Transverse ground section through a dentary tooth of Verenus selceber. A:


x20;B: x00

tooth. But due to the development of plicidentine, the replacement teeth of platY
notans develop in an interdental position. This results in the discontinuity of func
tion at any given site in the jaws of Varanws or other platynotans, a functiofl
disadvantage which must be compensated by a functional advantage of plieidentiTle
Varanus is a pleurodont lizard. The tooth becomes ankylosed to an obliquely sloping dental shelf on the tooth bearing bone. The teeth of Varanus show widely flared
bases which in some cases may be overgrown by bone (adult specimens of Var
niloticus) what gives the teeth further support. Striations on the bases of the teeth
result from the formation of plicidentine.
The crorn of the adult tooth of Varanus is a blade like structure, laterally C01fl
pressed with a recurved tip and an anterior and posterior serrated cuthng edge. It
is curved in two directions. The transverse section through a dentary tooth of Vart
saleator (Fig. 9) shows the lingual wall of the tooth base to buldge mesially, produdhh1

I. The Posterior Curvature of the Teeth

Recurvature of the teeth is of adaptive


significance in successfully seizing and
handling prey (Fig. 8). Selection will tend
to produce a maximum of recurvature.
However, the more reeurved a tooth is, the
more difficult is it to bring the tooth

810

0. Rizrrrr4: A Functional Interpretation of the Varanid Dentition

0. Rixnxn: A Functional Interpretation of the Varanid Dentition

811

Table 1
Same values as in Table 2, but for the dentaries of two specimens of Ttcronus solvotor
with complete dentitior.. L: left, R: right
MBSspecimen
numbers

Tooth positions

10755 H

74

73.5

10755 L

76

72

6 138 H

77

78

71.2

6138L

82

80

77

58
62

10

60

59.8

59

60

48.5

46.5

60

55

66.2

66.8

63

73

64

63

73.8

60

67

71

63

67

72

55

57

Table 2
Degrees of reeorvation of the teeth along the tooth row in seven specimens (loft maxillaries) of
Voronus solvotor. The angle is measured between the tangential line indicating tho direction into
which points the tip of the tooth and a horizontal line representing the lower edge of the maxillary
bone (Fig. 23). An increase of the recurvation results in the decrease of the numerical value of the
angle
MBS-specimen
numbers

Tooth positions
1

3713

69.5

78.8

60.8

55.5

3834

79

74

53.2

67

10 755

78.8

5028

85.5

3 835

74.5

79

85.4

69.5

77

76

74.5

77.8

69

59.5

5901

68.5

63

67.2

55

50.5

10

65

11

60.3

61

57.2

55

Fig. 11. Recurvation of the teeth along the tooth row in, top: maxilla of Varenns .901voter (MBS. 5878), bottom: dentary of Forming indicus (MBS. 11099)

43

59

51

12

45.8
59

67.5
53.2

9
56.9

58
76

74

5878

66.5

65

54.2
62

53.5
53

56

meso

62

into a position so that it may hit the prey with its tip. Yet it is critical for the tooth
to hit the prey with its tip. In doing so, maximal stress is exerted on the preys
surface and its penetration is more likely to be accomplished. A second critical
factor of tooth function is the axial loading of the tooth. This can only be achieved
if the tooth hits the prey with its tip. Axial loading reduces the bending moments
to which the tooth is subjected during the strike.
To bring a revurved tooth into an erect position it is most economical to rotate
the tooth along the circumferential line which represents its longitudinal axis. The
stronger the tooth is recurved, the closer will the center of rotation approach the
posterior cutting edge of the tooth. The lower jaw is hinged on the mandibular joint
which lies far posterior to the tooth row. Yet, the posterior dentary teeth are closer
to the mandibular joint than are the anterior teeth, and hence the posterior tEeth
show an increase in degree of recurvature which compensates a decrease in absolute

meta

occ

hy

Fig. 12. Loft maxillary dentition of Voronns solvator (MBS. 10755) rotated around the
]iypokinetie joint only (by; solid line) or around the occipito.atlantal joint only (0cc;
line). The rotation around the hypokinetic joint matchos the roeurvation of
the
teeth. Abbreviations: hy, liypokinetic joint; moso, mesokinetic joint; motn, rnota
dasliod
kinotik joint; oec, occipital condylo. Scab equals 10mm

size.

812

0.

RIErrEL: A Functional Interpretation of the Varanid Dentition

0.

The maxilla is hinged on the occipital condyle far back behind the functional
tooth row, but it is also hinged on the mesokinetic axis. Rotation of the muzzle unit
is made possible through the combination of the hypokinetic and the mesokinetic
joints (Fig. 12).
As the hypokinetic joint lies immediately behind the maxillary bone, a greater
degree of reeurvaftre of the maxillary teeth can be expected in comparison with the
dentary teeth. This is indeed the case, and again the more posterior teeth approach
ing the hypokinetie joint more closely show a relative increase in recurvature but a
decrease in absolute size in comparison with the anterior teeth. The increase in degree
of recurvature may be critical for the function of the maxillary teeth since it is the
upper tooth row which is more active during the feeding process.
The combined rotation of the upper jaw around the hypokinetie and mesokinetie joints
and around the occipito-atlantal joint not only allows an increase in degree of recurva
ture, but it also alters the line of action of a given tooth (Fig. 13). The line along which
a tooth tip of the upper tooth row will travel was studied with a simple mechanical model
of a TTaranus head, constructed according to the quadric crank chain model of FRAz
ZETTA (1962). If the maxilla rotates around the occipito-atlantal joint only, eery
maxillary tooth tip will describe the upper left quadrant of a circle line (with the
snout pointing to the left, Fig. 13 A). The gape required to bring the tooth into a
fully erect position is wide, and the tooth points backward at a very early stage of
jaw closure already. Thus the tip of the tooth may point backward as it hits the prey.
It may hit the prey with its anterior cutting edge. Moreover, the line of action of the
tooth contains a forward component of the strike ST which will affect the direction
of the resultant force produced by the tooth as it hits the prey. Thus, if a maxilla

Rririxe: A Fnnctional Interpretation


of the Varanid Dentition

813

which bears recurved teeth is rotated around the


occipito-atlantal joint only, the
recurved teeth are more likely to hit the prey with their
anterior cutting edges and
hence the teeth will not be loaded axially. Also, the resulta
nt force produced by the
teeth is likely to point in an anterovenfral direction
and the prey is then likely to
slip off the anterior cutting edge (I0RDANSKY, 1966).
If the maxilla is rotated sinaultaneously around the
occipito-atlantal joint and
around the hypokinetic and mesokinetic joint (Fig. 13 B),
the line of action of the teeth
is changed. The gape required to bring the teeth into a fully erect
position is smaller. The
teeth are more likely to hit the prey with their tips which
results in axial loading of the
teeth. The anterior force component of the strike ST is reduce
d and combined with
a posterior component which results from the simultaneous
snout depression during
jaw closure. The resultant force produced by the teeth is
now directed posteroven
trally, driving the teeth into the prey and the prey further
down the throat (Ion
nxNsuv, 1966).
K. Tooth attachment in

J7eranus

The dentary teeth of Varanus are not only curved in a posteri


or direction but also
in a lhgual direction as it is shown by the transverse section
(Fig. 9). Axial loading
is not only critical in the sagittal plane but also in the
transverse plane. In order to
be axially loaded in the transverse plane it is necess
ary for the teeth to be rotated
labially before they hit the prey. A labial rotation of
the dentary during mouth
opening is easily understood on the basis of the structure
of the mandibular joint
described above.
When Varanus is seizing and handling large prey, a heavy load
is imposed on its
teeth and on their attachment to the tooth hearing
hones. The flared bases are one
of the characteristics of the varanid teeth. Broad
bases reduce the forces working
per
I
unit area. Yet, tooth attachment is on an obliquely sloping
bony surface (pleuro
donty) and upon loading of the tooth a couple will build
up which produces tension
on the one and eouipression on the other side
of the tooth base. Resulting shear
stresses would tend to break the tooth off the bone,
especially since mineralized hard
tissues have a weak resistance to shear stresse
s (WAINwRmHT, Bmos, CURRY and
GOSLINE, 1976).
The transverse section through a varanid tooth (Fig.
9) shows that the labial wall
of the tooth base rests on the obliquely sloping
shelf of the tooth bearing bone. The
mesial rim of the tooth bearing shelf is horizontal
and was termed Basalpicue by
LESSMANN (1952). On it rests the lingual
wall of the tooth base. The mesial edge of
the tooth bearing shelf may slant upward and was
called Theka by LES5MANN (1952).
The dentine lamellae which subdivide the pulp cavity
as a result of the formation
of plicideutine appear as dentine trabeeulae
in the transverse section (Fig. 9). The
dentine trabeeulae run in an axial direction and meet
the tooth bearing bone at right
angles in any part of the tooth base (Fig. 9 B). The
dentine trabeeulae ate connected
With each other by weak transverse anastomoses.
Thus the base of the tooth be
Comes filled by a network of dentine lamellae in way
a
very similar to spongy bone.

Fig. 13. A: Maxillary tooth of Vera, us rotated around the oeeipito.atlantal jont only.
B: Maxillary tooth of Vereneis rotated simlIltaneoIsly around tIle oeeipitoatlt
joint and around the Itypokinetie and mesokinetie joint. ST: Forward componen f the
strike

I.

0. Rsarrna: A Functional Interpretation


of the Varanid Dentition

0. Riurrar,: A Functional Interpretation of the Varanid Dentition

814

This will certainly increase the stability of the tooth base. The formation of pliciden.
tine in platynotan lizards may indeed be correlated with the large. size of their teeth
(PEnn. 1968).
To study the stress distribution within the tooth base, plate models were cut from
plates of Araldite B to the size 35 times the size ofthe original tooth. Their shape
represented transverse sections at various levels through the functional tooth. The
loaded and inspected with light passing through crossed polaroids
tooth models
between which the models were mounted in an iron frame. Isoclinics appear as dark
lines on the model connecting these parts of the model where tbe directions of the
principal stresses are either parallel or perpendicular to the optical axis of the pola
roids. If the polaroids are rotated stepwise around 90, a series of sets of isoelinics
will be observed wandering across the model. Each set of isoclinics corresponds to
each step of rotation of the optic axis of the polaroids. The observed sets of isoclinics
can be superimposed on an outline drawing of the model. and with the indication
of the correspondiug optical axis of the polaroids the direction of the principal stresses
all across the model can he found. The isoelinics by themselves do not differentiate
between tensile and compressive stresses, but these can be inferred by pressing ones
nail into the edge of the model. If the isoelinies are attracted toward this
finger
stressing edge, tensile stress is indicated. If the isoelinics withdraw from the stressing
edge, compressive stresses are indicated (Nageiprobe: FPPL and MNdH, 1959). Note
that for any point of the model the direction of one principal stress (tension or corn
pression) is by definition perpendicular to the direction of the other principal stress
(centrifugal stress resulting from the deformation of the model).
Models of transverse sections through the functional varanid tooth were loaded
at different angles which resulted in changes of the directions of the resulting prin
cipal stresses (Fig. 14). The results indicate that upon axial loading (Fig. 14 A), ten
sile stresses result on the lingual side of the tooth base while compressive stresses
build up on the labial side of it. Moreover, upon axial loading the one direction of
the principal stresses (tension and compression) exactly parallels the direction of the
dentine lamellae of the plicidentine (Fig. 9 B) and meets the tooth bearing bony 5Wface at right angles anywhere within the tooth base. The other principal stress (cen
trifugal stress resulting from deformation) by definition runs parallel to the surface
of the tooth bearing bone. Therefore, no shear stresses will occur along the site of
ankylosis, and the dentine lamellae will be loaded in pure tension and compression

815

were

in5.

th
bp

Fig. 14. The direction of tensile


and compressive stresses within the
tooth base of
Vuranus. The tooth

is presented in a transverse
section under varying loads. For
further explanations see text.
Abbreviations: th, Theka; bp, Basalplattc; ling,
lingually;
lab, labially

as long as axial loading is maintained.


The same stress distribution is observed in the model if the tooth is loaded verti
cally. The loading force is then no more oriented axially through the tip of the tooth,
but it still passes through the surface of ankylosis. Hence, upon axial or vertical
surface
loading of the teeth of Varanus, shear stresses are absent or minimal on the
of ankylosis.
iiiore
If. however, the tooth is loaded abaxially so that the loading force does 110
stres
pass through the surface of ankylosis (Fig. 14 C), the directions of the prineivil
direction
the
ses will no more match neither the direction of the dentine lamellae nor

of the surface of ankylosis. As a


consequence, shear stresses ivihl build up along
the
site of ankylosis, which might cause the
tooth to break off the supporting bone.
The formation of plieidentine and the
degree of the slope of ankylosis may represent
adaptations of varanid lizards to use their
teeth to handle large prey imposing heavy
loads; The deee of pleurodonty is
modelled such that no shear stresses occur
along
the surface of ankylosis if the teeth
are loaded axially or vertically so that the
loading
force passes through the tooth base.
The dentine lamellae take up the compressive
or tensile stresses along their long
axes.
Acknowledgcmenis
I thank PD Dr. B. 0. San,
University of Basel, and Prof. Dr. II. L.
McGill Univer.
Sity, Montieal, under whose
supervision this work was carried out.
Prof. Dr. C. GANS, University of
Michigan, Ann Arbor, and Prof. Dr.
G. Ganuax, Institute
of Dental Medicine,
Univcrity of Basel, took a lively interest
in tho problems of mandibular
mechanics in loran us and engaged
in many stimulating
discussions. Dr. A. Kaamcuaxoaa

Vided the ground sections


through the teeth of

pro

and Prof. Dr. M. EaTCE made the X-ray


possible at the Kantonsspital, Basel. Dr.
IV. Ercssaaoan provided the pos
Sibility to carrying out tlio
experiments with phutoclastic tooth models
at the Ciba-Geigy AG,
Base1, and Dr. V. Essamean,
Eidgenossische Materialprufungsanstalt
Dbendorf, provided the
of a critical discussion of these
experiments.
Many Curators of Museum collections
provided generous access to their skeletal and
alcoholoic
Collections They are Dr. K.
Ki4anran, Seckcnberg Museum, Frankfurt
a. M., Dr. E. N. An
50t0 and A. C.
British Muserum (Natural History), London,
Dr. B. 0. ZWEIFEt,
Amcrican Museum of Natural History,
New York, Dr. F. F. WIIAMs, Museum of
Comparative
Zoology, Cambridge Mass.. Dr.
B. A. NussBAnr and Dr. A. 0. Ktuea, The
Museum of Zoology,
toM, Ann Arbor Mi., and Dr. H. M.&ax, Field
Museum of Natural History, Chicago.
lumens,

0. Rtnn4: A Functional interpretation of the Varanid Dentition

816
Summary

0. Rmna: A Functional Interpretation of the


Varaniti Dentition

817

Comments on inertial feeding. Copein, 855 (1969).


GosrEs, N. SI. B., J. P. Gssc: Etude
biomchanique du movement do ia formeturo do
Ia mandi
hub choz Oph/se-eras apodns (Sauna.
Anguitlao). Pap. Avulsos Zoo). 27 (1973) 125.
HECHT, M. K.: The morphology and
relationships of the largest known tern striai
lizard, Megoloaja
prism Owen, frem the Pleistocene of Australia.
R. Sec. Victoria Proc. 87 (1975) 239250.
ROPER, H.: Vergleichicnde Untersuchiungen
am Schatiel von Tupiaem&js und Coronas,
mit be
sonderer Berucksichtigung direr Kinctik,
Morph. Jb. 100 (1960) 706746.
Hol-misTEflEji H., J. P. Case: Vertebrae anti ribs,
In: Biology of the Reptilia (Gas, C., T.
S.
Pamisoets, eds.) 1 (1969) 201310.
lswv, 0.: Functional aspects of cranial
kiactism in the Lacertihia. Unpubi. Phi. D. thesis,
Univ.
Oxford (1967).
IoRnANsKv, N. N.: Cranial kinotism in lizards;
centnihution to the problem of the adaptive signi
J
ficance of the skull kinetism, Zoel. Zht. 45
(1966) 13981410 (In Russian, with English
summary).

A cinradiographic analysis of the feeding movements iii l7m-enus bengolenais produced the
following results. The mouth is opened by raising the head (upper jaw) rather than by lowering
the lower jaw. Starting from the resting position, the muzzle unit is elevated around 9 reintive
to the rest of the skull during jaw opening; the quadrate swings anteriorly around 21. During
jaw closure, the snout is depressed around 15 relative to the rest of the skull, hence 6 beyond
the resting position. The quadrate swings backwards around 27.
Amphikinesis is interpreted as allowing a stronger posterior recurvature of the maxillary teeth
in Verensis. This increases the holding effect of the teeth without increasing their length, an adap
tation of Vorcoius to capture relatively large prey.
The formation of plicidentine (dentino infolding) in the teeth of Vorenus increases the surface
of attachment of the teeth on the supporting bone. Moreover, the dentine lamellne take up ten
sile anti compressive stresses along their long axes upon axial or vertical loading of the teeth.
Tho slope of plourotlonty is modelled so as to minimalize shear stress on the surface of anky
losis upon axial or vertical loading of the teeth.
Zusammenfassuag
\4tiilirentl ties Frehoktes dffnet Voronus benqeien.sis das Maul rioreli Hebung dos Oberkiefers
und nicht dureh Senkong ties Unterkiefers. Wiihrontl ties Offnens tier Kieter wird the Sc loiauze
urn 9 relat iv zum Rest ties Sclikdels angeliohen; tlas Quadratum drelit sick tiahei um 21 nach
verne. Wkhrenti tier ScldieBung tier Kiefer wird die Sehnauze urn 15 reiativ ztim Rest des Sehd
dels gesenkt, tI. h. sic senkt sich urn 6 fiber die Ridielage hinaus. Das Quadrnttim dreht sich dabei
um 27 nach hinten.
Die ampliikinetiscbeii Bewegungen im Sehddel von Coronet eriatthten eine stdrkore Rdekwilrts
krfiminung tier Zkhne. Dabei wird tier Halteeffokt tier Ziilme vergrd6ert oline deren abseiute Lange
zo steigern.
(lie
Die Bildung von Pheidentin erhdht the Atifingefidehe cmos Waranzahnes. Zndem aehmen
eatngen
Druekspannu
untl
Ztigthe
Ziihne
tier
Dentinfalten bei axialer otler vertikaler Belastung
lang ihrer Liingsaehse en f.
Zahnc
Der Pleurotientiewinkel ist so eingeriehtet, tiall bci axialer other vertikaler Belastung der
t
werden.
Seherkrafte entiang der Atifiagefiiiche minimahsier
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