BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, SCIENCES DE LA TERRE, 62: 171-194, 1992

BULLETIN VAN H ET KONINKLIJK BELGISCH INSTITUT VOOR NATUURWETENSCH APPEN, AARDWETENSCHAPPEN, 62: 171-194, 1992

The Tylosaurine Mosasaurs (Reptilia, Mosasauridae) from the Upper

Cretaceous of Europe and Africa

by THEAGARTEN LINGHAM-SOLIAR

Abstract La biomécanique du crâne des tylosauriens est examinée principale-

ment suite à l'hypothèse selon laquelle le grand rostre servait à atta-
quer les proies (RUSSELL, 1967). Divers autres aspects du comporte-
This study represents the first relatively extensive description of the ment prédateur des tylosauriens sont aussi mentionnés.
PUUM Hainosaurus D o n o , 1885. The description of H. bernardi
DOLLO, 1885 is based on the holotype and on a previously undescri- Mots-clefs: Hainosaurus, Tylosaurus, Leiodon, prédation, plongée.
bed specimen. A new specimen of H. gaudryi (THEVENIN, 1896) from
France is also described. Mosasaurus iembeensis TELLES-ANTUNES,
1964, from the Turonian of Angola is reassigned to the genus Tylosau-
rus MARSH, 1872. The only other tylosaurine species from Africa, T. Introduction
capensis (BROOM, 1912) is also briefly described.
The biomechanics of the skull of tylosaurines is examined primarily The gigantic tylosaurine Hainosaurus bernardi DOLLO,
because of the hypothesis that the large rostrum was used in ramming
1885 was the first mosasaur to be discovered in Belgium.
prey (RUSSELL, 1967). Various other aspects of tylosaurine predatory
behaviour are also mentioned. Almost the entire skeleton was found intact, although
considerably abraded, in the Ciply Phosphatic Chalk in
Key-words: Hainosaurus, Tylosaurus, Leiodon, ramming, ambush- a region known as "La Malogne" (Fig.l). It was subse-
predation, diving quently described in several papers by D O L L O (1885a, c;
1889; see L I N G H A M - S O L I A R & N O L F , 1989 for the geo-
logical setting).
Résumé
Considerable confusion surrounds the tylosaurine
mosasaurs and the genus Leiodon. It originated with
Cette étude constitue la première description relativement approfondie
du genre Hainosaurus DOLLO, 1885. La description d'H. bernardi COPE'S (1869-1870, p. 200) misappropriation of the
DOLIO, 1885 se base sur l'holotype et sur un spécimen non décrit name Leiodon (and corruption to Liodori) to the genus
jusqu'à ce jour. Un nouvel exemplaire d'H. gaudryi (THEVENIN, Tylosaurus. C O P E (1870) realized that he had made a
1896) provenant de France, est également décrit. Mosasaurus iem- mistake in naming a mosasaur from the White Rotten
beensis TELLES.ANTUNBS, 1964, du Turonien d'Angola, est attribué
Limestone of Alabama, as Mosasaurus brumbyi. He
au genre Tylosaurus MARSH, 1872. Les seul autre tylosaurien africain,
T. capensis (BROOM, 1912) est brièvement décrit. was satisfied of the error by the presence of free haemal

Urnon
<?
quaternaire
o
IT)

E Sable landénien

Craie phosphatée

Fig. 1 — Discovery of Hainosaurus bernardi IRSNB R23 in La Malogne, near Mesvin in Belgium (after LECLERCQ & BOUKO,
1985)
 172 Theagarten LINGHAM-SOLIAR

Fig. 2 - Reconstruction of Tylosaurus incorrectly assigned to Leiodon by Owen 1879, (modified from Owen, 1880)

arches (fused in Mosasaurus) and concluded that the tically identifiable teeth for Hainosaurus at the time
specimen had to belong to Leiodon ( C O P E ' S "Liodon") (identifications on Leiodon are essentially based on its
for which there were no vertebral descriptions. O W E N unique tooth morphology). Exceedingly poor preserva-
(1879, 1880) himself was guilty of a similar error and tion of the few available fragmentary teeth in the holo-
added to the confusion that surrounds Leiodon to the type of Hainosaurus bernardi and IRSNB 3672 makes
present day. For instance he ( O W E N , 1879) also assi- this kind of judgement dubious. The conditions that are
gned postcranial remains of Tylosaurus ( = 1Macrosau- discernible are, an absence of ribbing on either side of
rus) to Leiodon. Indeed his reconstruction of Leiodon the carina and lack of lateral compression in the poste-
( O W E N , 1879; Fig.2) is evidently the same as Tylosaurus rior teeth - presence of either condition is highly diag-
(note also the massive premaxillary rostrum). These nostic for Leiodon. Recent evidence from a new species
errors were based on a simple premise. The postcranial Hainosaurus gaudryi, discussed in the taxonomic sec-
skeleton of Tylosaurus resembled neither that of Mosa- tion, conclusively indicates that Leiodon anceps and
saurus hoffmanni nor M. missouriensis, and both C O P E Hainosaurus are generically distinct.
(1870) and O W E N (1879, 1880) mistakenly concluded
that it therefore had to belong to the only other large
recognizable taxon at the time, Leiodon. Nor was the
genus Hainosaurus exempt from this confusion. D E P E - Abbreviations & Addresses
RET & Russo (1925, p.340) and P E R S S O N (1959, p.465)
have suggested that the tylosaurine Hainosaurus BMNH British Museum (Natural History).
D O L L O , might prove to be congeneric with Leiodon Department of Palaeontology. Cromwell
anceps. RUSSELL (1967, p. 142) on the other hand was Road. London S.W.7 5BD. U.K.
in no doubt that Leiodon compressidens and L. mosa-
sauroides are generically distinct from Hainosaurus but IRSNB Institut Royal des Sciences Naturelles de
he was uncertain about L. anceps. It is clear though that Belgique.
if the latter species was found to be identical to Haino- Rue Vautier 29. B-1040 Brussels. Belgium
saurus then because of Leiodon's priority ( O W E N , MNHN Museum National d'Histoire Naturelle
1840-1845) the name Hainosaurus would have to be (Dept. of Palaeontology). 43 Rue Cuvier.
abandoned and all the remaining Leiodon species placed 75231 Paris. VI. France
in a new genus. Fortunately this is not the case, for the
most curious aspect of D E P E R E T & Russo's (1925) and MMMN Manitoba Museum of Man and Nature,
PERSSON's (1959) suggestions that these two forms may Winnipeg, Manitoba, Canada
be congeneric lies in the fact that there were no diagnos- SGMA Servicos de Geologia e Minas de Angola
 The Tylosaurine Mosasaurs from Europe and Africa 173

USTL BUG (coll. Lab. Paléo) Bugarach atlas synapophysis small and flattened or rudimentary,
hypapophyseal peduncle located posteriorly on ventral
SL Saint-Louis surface of cervical centra, articulation for hypapophysis
JOU La Jouane. circular with central excavation, five hypapophyses-
bearing cervicals, two or three more with rudimentary
peduncles, transverse process of pygal vertebrae relati-
Systematic palaeontology vely short, neural spines of caudal, longest and vertical
on postsacrals 38-40.
Order S Q U A M A T A O P P E L , 1811 Scapula relatively the smallest in the Mosasauridae;
Family M O S A S A U R I D A E G E R V A I S , 1853 much smaller than coracoid. Superior border of scapula
Subfamily T Y L O S A U R I N A E W I L L I S T O N , 1895 strongly convex. Coracoid does not expand medially to
WlLLISTON 1897 point behind glenoid articulation. Distal and proximal
ends of slender humerus only slightly expanded, distal
Tylosauridae M A R S H , 1876: 59, nomen nudum. end more expanded than proximal; internal trochanter
"Mosasaurines megarhynques" D O L L O , 1890: 163. of average proportions and located medially from head;
Tylosauridae W I L L I S T O N , 1895: 169. radial process absent, facets for articulation with other
Tylosaurinae W I L L I S T O N , 1897: 177. elements and sites of muscle attachment not well diffe-
rentiated. Radius very elongate, proximal end very
DIAGNOSIS
slightly expanded, shaft narrow, distal end very slightly
(see R U S S E L L , 1967). expanded. Ulnare and fourth carpal, present, lack arti-
culating surfaces. Metacarpal one equal to metacarpal
two in length.
Hainosaurus D O L L O , 1885 Vertebral formula: 40 presacral vertebrae, 9 + pygals,
30-35 intermediate caudals, terminals 33 + (See DOLLO
Hainosaure D O L L O , 1885a: 285. 1885a, c).
Hainosaurus D O L L O , 1885a: 288.
Hainosaurus P O M P E C K J , 1910: 125.
Hainosaurus D O L L O , 1913: 612. Hainosaurus bernardi D O L L O , 1885
ILeiodon D E P E R E T R U S S O , 1925: 340. Figs. 1, 3, 4, 5, 6, 7, 9, 10, 12; Pis. 1, 2, 3, 4, 5, 6
ILeiodon P E R S S O N , 1959: 465.
Hainosaurus bernardi D O L L O , 1885a: 288.
Generic type Hainosaurus bernardi D O L L O , 1885b: 31.
Hainosaurus bernardi D O L L O , 1889: pis 9, 10.
Hainosaurus bernardi D O L L O , 1885 Hainosaurus bernardi D O L L O , 1904: 207.
Hainosaurus bernardi D O L L O , 1909: 103.
DIAGNOSIS Hainosaurius bernardi [sic!] Pompeckj, 1910: 139.
Double buttressed premaxillary suture. Twelve to thir- Hainosaurus bernardi D O L L O , 1924: 172.
teen teeth in maxilla. Supraorbital wing of prefrontal ILeiodon anceps D E P E R E T and R u s s o , 1925: 340.
covered dorsally by frontal; prefrontal forms part of ILeiodon anceps P E R S S O N , 1959: 465.
posterolateral margin of external nares. Prominent Hainosaurus bernardi L I N G H A M - S O L I A R , 1991C: 174¬
median dorsal ridge on frontal; frontal not emarginate 175, fig. 5.
above orbits. Margins of parietal straight as far as pos-
terior diverging suspensorial rami, forming rectangular HOLOTYPE

field medially on parietal. Squamosal wing to parietal IRSNB R23 (old n o . R1564), almost complete poorly
moderately developed. Large otosphenoidal crest on preserved cranial and postcranial remains.
prootic covers exits for cranial nerves VII and IX. Broad
projection on dentary anterior to first dentary tooth. H O R I Z O N AND LOCALITY

Ten to eleven teeth on pterygoid. Ciply Phosphatic Chalk, Upper Maastrichtian, in "La
Articulating surfaces of cervical and anterior dorsal Malogne", near the town of Mesvin, Belgium.
vertebrae nearly circular (except for the atlas which is
elliptical); synapophysis located in centre of lateral sur- REFERRED MATERIAL

face of cervical centra, occupies anterodorsal portion of IRSNB 3672, almost complete skull and large number of
lateral surface of dorsal vertebra. Ventral border of vertebrae. From the Maastrichtian Phosphatic Chalk of
anteroventral extension of synapophysis not strongly Baudoir, Belgium.
developed on cervicals and anterior dorsals, does not
reach level of undersurface of centrum; anterior zyga- DIAGNOSIS
pophyses of cervicals and dorsals connected by sharp Premaxilla with long ventral process that extends poste-
ram-rod straight crest posteroventrally to synapophysis, riorly to approximately the second maxillary tooth.
zygosphene-zygantrum rudimentary. Anterior base of External nares large - 28-31% of skull length. Parietal
atlas neural arch arises directly above condylar facet,
 174 Theagarten LINGHAM-SOLIAR

foramen small, located on the fronto-parietal suture. nae although in the holotype the dorsal outline is more
Ventromedial process of postorbitofrontal forms shal- rectangular than in IRSNB 3672 and the fragmentary
low excavation to receive distal process of the vertical specimen of Hainosaurus "lonzeensis" (no specimen
arm of the jugal, probably ligamentous; POF wing to number; D O L L O , 1904). Ventrally a long process on
parietal deeply invades posterior frontal border either either side of the premaxilla extends posteriorly to the
side of parietal foramen. Ventroposterior process on second maxillary tooth (Fig.4). The premaxillary suture
jugal absent. Tympanic ala of quadrate thin. Stapedial in IRSNB 3672 is highly unusual forming a double poin-
pit rectangular in form. Dentary long, thirteen teeth pre- ted buttress with the maxilla (Figs. 3A, 4A, PI. 2A). In
sent. Angular widely separated medially from coronoid. the holotype it is just distinguishable on the right side
Retroarticular process of articular posterodorsally roun- despite poor preservation. The suture then rises gently
ded, ventrally straight. Teeth - anterior and posterior from this point to the posterior margin of the external
carina extend full length of crown; internal and external nares and instead of descending as in Tylosaurus ( R U S -
striae fairly well developed. SELL, 1967, p.177) continues in a gentler gradient to the
dorsal termination. Unfortunately the posterior sutural
DESCRIPTIONS AND COMPARISONS contact with the prefrontal is not preserved in either the
SKULL holotype or IRSNB 3672.
The skull is massive with a prominent anterior rostrum The internarial bar extends deep into the frontal to
as in other members of the Tylosaurinae (Fig.3; Pis. 1 approximately a third its length (Fig.3B) very much as
& 2). Comparisons are made with the North American in Tylosaurus proriger ( R U S S E L L , 1967, p. 172, fig.92).
species Hainosaurus pembinensis (as described by In Hainosaurus pembinensis MMMN V95, however, it
N I C H O L L S , 1988) and the genus Tylosaurus proriger extends to well over half the length of the frontal (obser-
and T. napaeo/icus (as described by R U S S E L L , 1967). ved from photographs, Elizabeth N I C H O L L S , pers.
comm.)
PREMAXILLA
The premaxilla in the holotype specimen is poorly pre- DISCUSSION
served with fine detail very much obscured. The large D O L L O (1904, p.213) erected the new species H. lon-
premaxillary rostrum is characteristic of the Tylosauri- zeensis on the basis of the differences in shape between

Fig. 3 — Hainosaurus bernardi (IRSNB 3672). Restored skull. A, lateral view; B, dorsal view.
 The Tylosaurine Mosasaurs from Europe and Africa 175

species and I have therefore assigned it as Hainosaurus

sp.
The ventral projections in the premaxilla of Haino-
saurus bernardi I R S N B R 2 3 have not previously been
described in the Tylosaurinae. They are apparently a
more robust counterpart of slender extensions that I
have seen on the ventral surface of the premaxilla of
Mosasaurus hoffmanni.
The diagnostic importance of the double buttressed
premaxillary suture in Hainosaurus can not be oversta-
ted as the condition is certainly not found in Tylosaurus
(cf. RUSSELL, 1967, p.177, fig.95). For instance the
large premaxillary rostrum in "Mosasaurus" gaudryi
( T H E V E N I N , 1 8 9 6 ) makes the material clearly referable
to the Tylosaurinae but the presence of the double but-
tressed premaxillary suture enables a more specific
assignment to Hainosaurus (functional significance dis-
cussed further on).

MAXILLA
Preservation of the maxilla of specimen I R S N B R 2 3 is
poor but fortunately it is much better in I R S N B 3 6 7 2
( F i g . 5 A ) . The maxilla is stout and generally consistent
with the element in other taxa in the Tylosaurinae. The
right maxilla has just two fragmentary tooth crowns
preserved and one in the left maxilla. Tooth bases indi-
cate that there were 1 2 teeth in life.

FRONTAL
The frontal in Hainosaurus bernardi IRSNB R23
(Figs.3B, 6A,B) is quite poorly preserved although there
are several notable characters. The fronto-parietal
suture is narrower in the holotype compared to the con-
dition in H. pembinensis ( N I C H O L L S , 1988) and Tylo-
Fig. 4 Premaxillae of three mosasaur taxa showing marked
saurus ( R U S S E L L , 1967, fig.92). The posterior boundary
variation. A, Hainosaurus bernardi demonstrates a
large prow and double buttresses premaxillary/maxil- with the postorbitofrontal is also somewhat more con-
lary suture; B, Goronyosaurus nigeriensis - the second cave. Lateral margins above the orbits are straight to
pair of premaxillary teeth is, uniquely in mosasaurs, slightly convex which is consistent with the condition in
the longest in the dental batery; C , Prognathodon sol- H. pembinensis (photographs, N I C H O L L S , pers.
vayi with prognathous premaxillary teeth (not to scale). comm.) but differs from the somewhat concave orbital
margin of Tylosaurus proriger ( R U S S E L L , 1967, p.172).
The prefrontal and postorbitofrontal bones exclude the
frontal from the orbital border. A pronounced crest
extends along three quarters of the length of the bone
along the midline although this is a variable condition in
the premaxillary rostrum in the holotype of Hainosau- Tylosaurus ( R U S S E L L , 1967, p. 171).
rus bernardi and that of the premaxillary fragment from The ventral surface of the frontal, despite poor pre-
the Glauconie de Lonzee "...par son rostre plus conique servation, presents several interesting features. The pre-
et a face superieure plus arrondie, et par ses dimensions frontals underlie the frontals, extending almost to the
moindres". However, the apparent difference is almost fronto-parietal suture and they are tightly sutured to the
certainly the result of poor preservation in the holotype frontal in a relatively deep excavation (Fig.6, P1.3B).
material. N I C H O L L S (1988, p.1566, fig.3A) also descri- The excavation for the olfactory lobes are apparently
bed a pronounced rectangular premaxillary rostrum of shallow. The postorbitofrontal is quite broad and the
H. pembinensis. Perhaps the highly gypsiferous nature prefrontal wing extends to the midpoint of the lateral
of the specimen ( N I C H O L L S , pers. comm.) could margin of the frontal. Dorsally a narrow process
account for the apparent rectangular configuration. extends from the postorbitofrontal and is sutured into a
Certainly in Hainosaurus "lonzeensis" the apparent dif- deep narrow excavation in the prefrontal. The descrip-
ference in the shape of the rostrum with that of H. ber- tion of the frontal is very similar in the Paris specimen
nardi is insufficient to warrant the erection of a new of Hainosaurus (described later).
 176 Theagarten LINGHAM-SOLIAR

100mm

Fig. 5 — Hainosaurus bernardi (IRSNB 3672). Right maxilla and dentary.

LACHRYMAL Tylosaurus, a narrow tongue of the prefrontal forms a

The lachrymal is well preserved in the holotype of Hai- part of the narial border.
nosaurus bernardi IRSNB 1 5 6 4 . It is quite broad but is
characteristically an arrow-head shaped bone as in other POSTORBITOFRONTAL
mosasaurs ( P 1 . 1 A ) . The postorbitofrontal of IRSNB 3672, although frag-
mentary, was clearly a rather robust bone (P1.3A). The
PREFRONTAL supratemporal wing is long extending to the posterolate-
The prefrontal is very poorly preserved in both the ral corner of the squamosal. The wing to the jugal is
holotype and IRSNB 3 6 7 2 . It is nevertheless apparent substantial, although incompletely preserved. The wings
from the available fragment and from the borders with to the parietal and prefrontal are absent. The large ven-
the frontals and maxilla that it is quite similar to the ele- tral wing to the frontal has, however, already been des-
ment in Tylosaurus proriger ( R U S S E L L , 1 9 6 7 , p. 1 7 2 ) cribed in the holotype specimen.
and Hainosaurus pembinensis. However, unlike in
JUGAL
The jugal in the holotype, although incomplete, is evi-
dently similar to that of Tylosaurus proriger ( R U S S E L L ,
1967, p.177, fig.95) with a fairly broad ala-like process
at the junction of the vertical and horizontal axes. There
is no evidence of a postero-ventral process.

SQUAMOSAL
The squamosal is poorly preserved in the holotype and
in IRSNB 3672. Anteriorly the squamosal wing extends
as far as the jugal wing of the postorbitofrontal.

NASALS
Quite unusually the nasals are preserved in IRSNB 3672
(P1.2B), fused to the internarial bar. They are known in
the literature in three other specimens, the type speci-
men of Plotosaurus bennisoni ( C A M P , 1942, pp. 27-28,
Fig. 6 — Ventral view of the frontal of Hainosaurus bernardi fig. 14.), in Tylosaurus ( H U E N E , 1910, p.303, fig.5) and
(IRSNB R23) showing prefrontal process deeply in Clidastes Sternberg! ( W I M A N , 1920, p. 15, fig.4). I
sutured close to the fronto-parietal suture. have observed well preserved nasals in a Tylosaurus sp.
Fig. 7 - Hainosaurus bernardi (IRSNB R23) Frontal. A, dorsal view; B, ventral view.

BMNH 3 6 2 5 (Fig.8) in which they are clearly paired and red ( E S T E S et al., 1 9 8 8 , p. 1 4 3 ) . Among squamates,
were either free or lightly sutured to the premaxilla. An Lanthanotus and Varanus have fused nasals and the
apparently tighter fusion of the nasals to the premaxilla condition also occurs in some chamaeleodontids, some
in IRSNB 3 6 7 2 may be the result of preservation. gekkonids and pygopodids, some scolecophidians and
some Leptotyphlops ( E S T E S et al. 1 9 8 8 , p. 1 4 3 ) . E S T E S
DISCUSSION et al. ( 1 9 8 8 , p. 1 4 3 ) regarded nasal fusion as synapomor-
In lepidosauromorphs, nasals are almost uniformly pai- phies within various taxa. I have done likewise and
 178 Theagarten LINGHAM-SOLIAR

RUSSELL (1967, p. 172, fig.92) indicates that the parietal

foramen in T. proriger is situated just posterior to the
fronto-parietal suture whereas in Tylosaurus nepaeoli-
cus it is set well into the parietal.
Descending wings of the parietal are preserved only in
IRSNB 3672 where they are rather short.

O C C I P I T A L UNIT
The occipital is very poorly preserved in the holotype
but somewhat better preserved in IRSNB 3672 (P1.4).
The passage for cranial nerve VII is obscured by the
large otosphenoidal crest. Unusually, for the Tylosauri-
nae, there appear to be two foramina for cranial nerves
X, XI and XII. The basioccipital condyle is moderately
large although both the basal tuber and basisphenoid
are poorly developed.

PTERYGOID
The pterygoid in the holotype is poorly preserved but
almost complete, forming approximately 44% of the
skull length. The tooth row in the holotype which is pro-
Fig. 8 — Tylosaurus (BMNH R3625) showing rarely obser- bably complete supports nine to ten teeth. Despite the
ved nasals. fragmentary nature of the pterygoids in IRSNB 3673
(PI. 3E) the tooth counts confirm those of the holotype.

regard the presence of free nasals in the Tylosaurinae as QUADRATE

a plesiomorphy of the subfamily and possibly of the The quadrate in the holotype is poorly preserved and
Mosasauridae. The condition of paired nasals in at least distorted. The suprastapedial and infrastapedial proces-
the Tylosaurinae (unknown in other mosasaurs) and ses are very small (Fig.9A, B, C). Medially the stapedial
fused nasals in Lanthanotus and Varanus also serves to pit is a rather pinched vertical ellipse. In IRSNB 3672
add to the dubiousness of M C D O W E L L & B O G E R T ' S
(Fig.9D, E, F) the suprastapedial process is somewhat
(1954, p.57) suggestion that the "precise affinities" of better preserved and despite absence of an apparently
Lanthanotus are with "the aigialosaur-dolichosaur- small fragment confirms the rather small size of the
mosasaur complex rather than with the varanids or
holotype. A fairly delicate tympanic ala indicates that
Heloderma". This view was also previously questioned
the quadratic tympanum was rather thin. The basal
on a number of other points in a powerful critique by
condyle is convex and expanded from side to side.
U N D E R W O O D (1957).

DISCUSSION
VOMERS The quadrate is essentially consistent with descriptions
Relatively well preserved vomers are present in IRSNB of this element in other members of the Tylosaurinae
3672 ( P I . 3B, C). Distinct ligamentous surfaces, although the suprastapedial process appears somewhat
medially along most of the length of the vomers makes larger in Hainosaurus pembinensis and Tylosaurus pro-
it clear that they were sutured to each at the cranial mid- riger ( R U S S E L L , 1967). Generally in H. pembinensis
line and free only at their anterior terminations. The Gudging from photographs, N I C H O L L S , pers. comm.) it
Jacobson's organs extend from approximately the mid- seems that the quadrate is more robust compared to the
point of the fourth to just past the fifth maxillary tooth. element in H. bernardi.
In Tylosaurus they lie opposite the fourth maxillary
tooth (RUSSELL, 1967, p. 26). Judging from the shallow DENTARY
groove on the vomer the estimated size of the Jacob-
The dentaries in the holotype of Hainosaurus bernardi
son's organ is approximately 76mm by 14mm. The late-
are very poorly preserved ( P I . 1A). The tooth row com-
ral surface of the vomers reveals a large somewhat
prises 14 teeth either very poorly preserved or consisting
concave sutural surface anteriorly and a convex sutural
solely of tooth bases. The dentaries in IRSNB 3672 (Fig.
surface posteriorly.
5B) although rather fragmentary, have much better sur-
face preservation. They are almost identical in shape to
PARIETAL those of Tylosaurus nepaeolicus ( R U S S E L L , 1967,
The parietal is characteristically tylosaurine. The parie- p. 177, fig.95). The edentulous process anteriorly is as
tal foramen is small and of similar proportions to that large with a similar, somewhat pinched crest almost at
of Tylosaurus proriger, situated on the fronto-parietal the dorsal termination of the bone. Ventral and dorsal
suture with perhaps a small part within the frontal. margins of the bone are relatively straight in IRSNB
 The Tylosaurine Mosasaurs from Europe and Africa 179

3672. In the holotype the dorsal margin is apparently In IRSNB 3672 the right and left dentarles held 13
more concave (campylorhynchus of W I L L I S T O N , 1 8 9 8 ) teeth in each ramus although the left dentary gives the
a condition that may be accounted for by the larger size impression of 14 because of a large replacement tooth.
of this specimen. Poor preservation makes a description There are just three moderately preserved teeth altoge-
of the ventral border inconclusive. The mandibular ther, the fifth in both dentaries and the 13th in the right
foramina are large in IRSNB 3672 and extend along the dentary. Each of the two anterior teeth have a single,
length of the dentary at about the vertical midpoint of sharp anterior carina. The posterior tooth posseses both
the bone. anterior and posterior carinae. All the teeth show faint
180 Theagarten LINGHAM-SOLIAR

facets. The best preserved teeth however, are in the CORONOID

French Hainosaurus, specimen M N H N 1896-15 (PI. The coronoid is poorly preserved in the holotype. The
3/51) in which the striae are clearly developed in both right coronoid in I R S N B 3672 on the other hand is
the buccal and lingual surfaces (described later). moderately well preserved. Compared with Tylosaurus
nepaeolicus, for example, the coronoid eminence rises
DISCUSSION more sharply and the lateral wing is more substantially
It is clear that there are no significant differences in the developed.
dentaries of Hainosaurus and Tylosaurus. The somew- The coronoid is, however, relatively smaller than in
hat stouter appearance of the dentary of H. pembinen- most other mosasaurs with the exception of Plioplate-
sis, ( N I C H O L L S , pers. comm.) may be the result of gyp- carpus ( D O L L O , 1889).
sum swelling.
ARTICULAR
SPLENIAL
The articular is a very robust, laterally compressed bone
Fragments of the splenial are preserved in the holotype
in the holotype of Hainosaurus bernardi and forms part
and IRSNB 3672 (P1.3D). The articulation contacts are
of the excavation for the quadratic condyle.
moderately deep excavations on the splenial and slightly
tuberous on the angular.
V E R T E B R A E A N D RIBS

SURANGULAR
Descriptions are essentially as for Tylosaurus (RUS-
The surangular is a rather expansive bone, characteristi- SELL, 1967, p. 171 and herein Pis. 5 & 6)
cally cone shaped with the point posteriorly directed. It
does not extend past the quadratic cotyle. Immediately SCAPULA AND CORACOID
in front of the glenoid articulation, a large foramen is The scapula and coracoid are as in Tylosaurus ( R U S -
present in IRNSB 3672. In the holotype, in contrast, this SELL, 1967) and are relatively the smallest in the Mosa-
foramen is absent. The surangular fits into a recess in sauridae (PI. 6; also see Fig. 13).
the articular, the latter bone is missing in IRSNB 3672.
Deep striae on the anterior wall of the surangular and PADDLES
posterolateral wall of the dentary suggests the presence The paddles in Hainosaurus are characteristically tylo-
of a powerful ligamentous sheet attaching the two moei- saurine showing the most reduced condition in the
ties of the jaw. Mosasauridae (Fig. 10).
 The Tylosaurine Mosasaurs from Europe and Africa 181

Hainosaurus gaudryi ( T H E V E N I N , 1896) (1896, p. 903) had mistakenly considered that the
(Fig. 11, PI. 7) slightly smaller (in tylosaurine terms) premaxillary ros-
trum was intermediate between the megarhynchus and
Mosasaurus gaudryi T H E V E N I N , 1896: 900. mesorhynchus types of rostrum of D O L L O (1890, p. 163)
Mosasaurus gaudryi S U Z U K I , 1985: 51. and consequently assigned the specimen to a new species
Hainosaurus bernardi B A R D E T , 1990: 752. of Mosasaurus. At the University of Paris VI, I also
assigned to Hainosaurus? sp. vertebral material (USTL
BUG, 2-28, SL, 1-10, JOU 5-12) on loan to Miss N.
Generic type
B A R D E T , which she presented to me for comment. This
Hainosaurus bernardi, DOLLO material from Sougraigne, in the department of Aude in
France, had previously been assigned to Platecarpus
HOLOTYPE ictericus ( C O R R O Y , 1927, SÉNESSE, 1936). Both deter-
minations have subsequently appeared in print (BAR-
M N H N 1896-15. Partial skull, premaxilla, dentaries,
DET, 1991). However, regarding specimen M N H N
maxillaries, frontal and partial parietal.
1896- 15, two important diagnostic characters indicate
H O R I Z O N AND LOCALITY
Phosphatic Chalk, Upper Santonian. Eclusier-Vaux that it warrants recognition on the specific level - the
fairly straight fronto- parietal suture and the location of
near Péronne (Somme), France.
the parietal foramen some distance from the fronto-
parietal suture. In striking contrast in H. bernardi the
DIAGNOSIS
Relatively short premaxillary rostrum for the subfamily. postorbitofrontal wings on either side of the parietal
Straight fronto-parietal suture. Parietal foramen situa- foramen form rather deep invasions of the frontal for-
ted some distance from the fronto-parietal suture. ming an irregular fronto parietal suture. Secondly the
parietal foramen is situated on the fronto-parietal
suture. The plesiomorphic states of a straight fronto-
DISCUSSION
In June, 1990,1 reassigned Mosasaurus gaudryi M N H N parietal suture and location of the parietal foramen well
1896-15 to Hainosaurus ?bernardi on the basis of a cha- within the parietal were discussed previously by L I N G -
racter I had previously established for the genus, based H A M - S O L I A R & N O L F (1989). They also coincide with
on IRSNB R23 and 3672, the unique double buttressing the early geological age of H. gaudryi M N H N 1896-15
of the premaxillary suture (evident in M N H N 1896-15 when compared with H. bernardi IRSNB R23 and
despite considerable abrasion; Fig. 11). T H É V E N I N 3672.
 182 Theagarten LINGHAM-SOLIAR

In the American forms of Tylosaurus the location of HOLOTYPE

the parietal foramen shows a similar distinction in the South African Museum SAMK5625. Consists of an
two species. In the earlier occurring Tylosaurus nepaeo- almost complete frontal, anterior portion of parietals
licus the parietal foramen is situated some distance from and portions of postorbitofrontals. Found in the Upper
the fronto-parietal suture while in the later form, T. Cretaceous chalk of the Transkei ( = Pondoland).
proriger it is situated on the fronto-parietal suture
( R U S S E L L , 1967, p.174). This character seems to be DIAGNOSIS
highly consistent in earlier forms of mosasaurs (e.g. in Large frontal lacking dorso-median crest, posterior
the earliest member of the genus Platecarpus bocagei, margins relatively convex, small parietal foramen, some
L I N G H A M - S O L I A R , in press b) and in itself is sufficient- distance from fronto-parietal suture.
ly diagnostic for specific recognition.
DISCUSSION
DENTITION
The distance of the parietal foramen from the fronto-
For the first time, specimen M N H N 1896-15 makes parietal suture suggests that the specimen is probably
available relatively well preserved teeth for the genus closer related to Tylosaurus nepaeolicus than to T.
Hainosaurus. The teeth are clearly robust as in Tylosau- proriger.
rus, cutting edges are not particularly pronounced, there
is no ribbing of the carina, cross-sections are almost cir-
cular and the tooth crowns are deeply striated ( P I . 7A). Tylosaurus iembeensis ( T E L L E S - A N T U N E S , 1964)

DISCUSSION
Mosasaurus iembeensis T E L L E S - A N T U N E S , 1964, p.165,
M E R R I A M (1894, p. 14) demonstrated that the teeth of pls.23-25.
Leiodon are smooth, resembling those of Clidastes, in
contrast to striated labial crowns in Tylosaurus. Despite HOLOTYPE
this, as I have already mentioned, D E P E R E T & Russo S.G.M.A. specimen (no number, Universidade nova de
(1925) and P E R S S O N (1959) have both suggested such Lisboa, Lisbon, Portugal). Poorly preserved, incom-
close similarities in the teeth of these two forms as to plete skull material consisting of portions of dentaries,
make them congeneric. Clearly the above description of posterior jaws, premaxilla, maxilla, pterygoid, quadrate
the teeth in Hainosaurus gaudryi indicates that there is and basioccipital.
no resemblance between the teeth of Hainosaurus and
Leiodon anceps whatsoever. Indeed a well preserved
H O R I Z O N A N D LOCALITY
tooth in H. pembinensis ( N I C H O L L S , 1988 pers. comm.)
Upper Cretaceous, Upper Turonian Chalk, "Camadas
also demonstrates that it is quite unlike anything seen in
do Tadi" near the town of Iembe in Angola.
Leiodon and more consistent with those of the Tylo-
saurinae.
DIAGNOSIS
Because of the fragmentary nature of the tylosaurine Distinct premaxillary rostrum although relatively smal-
material described below, it is difficult to distinguish ler than in other members of the Tylosaurinae. Supras-
between Tylosaurus and Hainosaurus and the specimens tapedial process of quadrate very short, infrastapedial
are therefore tentatively referred to as Tylosaurus. process reduced, tympanum shallow, pinched stapedial
pit. Robust dentary, rostrum present. Marginal teeth
Tylosaurus M A R S H , 1872 large, striated (not prismatic), subcircular cross-secti-
ons; twelve maxillary teeth, 13 dentary. Pterygoid teeth
Leiodon in part, C O P E , 1869-1870: 200 moderately large. Zygosphenes absent on cervical ver-
Rhinosaurus M A R S H , 1872a: 461 (preoccupied, F I S H E R tebrae.
von W A L D H E I M , 1847).
Rhamphosaurus C O P E , 1872: 141 (preoccupied, FlTZlN- DISCUSSION
GER,1843) The large size of the specimen combined with the cha-
racters in the above diagnosis indicate that the material
Tylosaurus M A R S H , 1872b: 147.
is referable to Tylosaurus. These characters are inconsis-
tent with those of Mosasaurus hoffmanni the only other
Generic type
known mosasaur of equal proportions. Its Upper Turo-
nian age makes it the oldest member of the subfamily.
Tylosaurus proriger ( C O P E , 1869).
The previously oldest Tylosaurus specimen came from
the ?Coniacian of the U.S.A. It is quite probable that
DIAGNOSIS
the relatively smaller rostrum may reflect an earlier evo-
(see R U S S E L L , 1967, pp.171-173)
lutionary stage of this structure. Indeed the Santonian
Tylosaurus capensis B R O O M , 1912 Hainosaurus gaudryi shows a relative size increase of
the rostrum intermediate between T. iembeensis and H.
Tylosaurus capensis B R O O M , 1912: 332-333, pl.22. bernardi.
 The Tylosaurine Mosasaurs from Europe and Africa 183

Biomechanical implications of the skull structure and the initial shock waves to the head during ramming. A
the lifestyle of hainosaurs and tylosaurs. delicate bar sutured between the external nares, as in vir-
tually all other mosasaurs, would simply have fractured
RUSSELL previously suggested that the massive rostrum or dislocated. Furthermore, location of the mosasaur
in members of the Tylosaurinae was probably used as a brain in a robust bony case that formed a sliding joint
ram in defence, or to stun prey (1967, p.69), comparable with the skull roof at the posteriormost part of the
to the ramming behaviour in certain dolphins ( W A T - exceedingly long skull was probably sufficient to cus-
SON, 1988; M A R T I N & R O T H S C H I L D 1989). However, hion the brain from any severe shock. Further support
such a prospect has not been examined further. While for this hypothesis lies in the unique condition of a dou-
superficially the comparison with dolphins may be a ble locking suture between the premaxilla and maxilla in
good one, the problem involves two highly different Hainosaurus (Fig. 12A) that presumably provided stabi-
types of skulls: a more or less solidly fused mammalian lity of the rostrum by a lock and key arrangement with
skull, and a reptile skull composed of numerous delicate the maxilla. This was enhanced by the large ventral pro-
and lightly sutured bones. Of prime consideration must jections from the premaxilla that acted as a strut bet-
be the fact that the impact on the skull during ramming ween the maxilla and premaxilla.
would clearly have been enormous. The key structure I The number of biomechanical changes in tylosaurine
therefore looked at in tylosaurs and hainosaurs was the skulls tends to indicate rather convincingly the functio-
bone leading from the rostrum and premaxilla, the nal feasability of the use of the rostrum in ramming.
internarial bar, which is generally very slender and deli- However, the possibility that it was used purely in sexual
cate in mosasaurs and sutured to a narrow anterior behaviour i.e male-male combat as distinct from preda¬
extension of the frontals between the external nares. In tion also needs consideration. T H U R M O N D ' S (1969)
striking contrast, in the Tylosaurinae, it is unique in two study indicates that there was little variation in the size
ways: it is exceedingly robust and broad and arises from of the rostrum in Tylosaurus specimens which would
a wide rectangular base. Of further significance is the imply that they were of generally similar size in males
deeply interdigitated nature of the suture well within the and females. It would thus be curious if a rostrum capa-
frontal, that provides a large interfacial sheer area, thus ble of use in combat between large male sexual rivals
allowing the transfer of increased stress (Fig.l2B). was not used, by clearly a voracious predator (see
Hence there is greater resistance to forces such as ben- M A R T I N & B J O R K , 1987), in securing food (the situa-
ding, shearing and breaking. The significance of the tion is very different in e.g. deer rutting, primarily
highly modified internarial bar in tylosaurs and haino- because they are herbivores).
saurs is of fundamental importance for absorption of Tylosaurs and hainosaurs were probably not the fas-
test of swimmers (sustained) in the Mosasauridae (for
swimming in mosasaurs see L I N G H A M S O L I A R , 1991a
and in press a). This is indicated by, for instance, their
rather poorly developed tail fins. However, there
appears to have been a massive reduction in body weight
in these forms. For instance pectoral and pelvic girdles,
including the paddles, are highly reduced, relatively the
smallest in the mosasaurs (Fig. 13). A further striking
condition associated with weight reduction is indicated
by WILLISTON'S (1897) observation that the bones of
Tylosaurus are highly cancellous and were probably
impregnated in life, with fat, a condition that presu-
mably increased bouyancy. This cannot just be accoun-
ted for by the large size of tylosaurines and need for
enhanced bouyancy because an equally large or even lar-
ger mosasaur, Mosasaurus hoffmanni (paper in prep.),
lacks highly cancellous bones. Such an apparently enor-
mous weight loss suggests that tylosaurs and hainosaurs
were evidently much more conservative in their energy
requirements. Presumably this was useful for patrolling
by stealth rather than by speed, over perhaps a fairly
wide ranging area, moving from one ambush site to ano-
ther. Sharks probably provide the best modern day ana-
logue ( W E B B , 1984). Furthermore the lower density
probably assisted in rapid acceleration when prey was
sighted, a condition W E B B & SKADSEN (1979) described
Fig. 12 Anterior portion of the skull of Hainosaurus bernardi in pike and tiger musky. Again, the long powerful tail
(IRSNB 3672). A, lateral view; B, dorsal view. of the tylosaurines was an ideal adaptation for burst
Fig. 13 - Log graph of cranial and postcranial ratios in four mosasaur taxa, Mosasaurus lemonnieri (IRSNB 3120), Hainosau-
rus bernardi (IRSNB R23), Mosasaurus hoffmanni (IRSNB R12), Plioplatecarpus marshi (IRSNB R39 & R38).

speeds, as seen in crocodiles ( M A S S A R E , 1988; L I N G - SCHILD'S (1989) theory that certain necrotic vertebrae in
H A M - S O L I A R , 1991a). This in combination with a large for example Tylosaurus were caused by this phenomena.
solid rostrum provided the potential for a powerful While as they suggest, squid was probably part of the
blow that may have killed or stunned the prey. diet of the tylosaurs and hainosaurs it seems probable
I am not convinced that hainosaurs and tylosaurs that they were more pelagic forms. Large size of tylo-
were deep divers as postulated by M A R T I N & R O T H - saurs and an ambush form of predation would clearly
SCHILD (1989), certainly not in the way that many wha- favour a much more mixed diet ( L I N G H A M - S O L I A R ,
les are. There is no evidence in mosasaurs for the 1991c). This is in fact supported by M A R T I N & BJORK'S
enormous morphological and physiological changes (1987) record of the gastric contents of a Tylosaurus
connected with deep diving that are seen for instance in specimen in the South Dakota School of Mines that
sperm whales. On the other hand dives of up to approxi- shows a rather awesome and varied diet consisting of
mately 50 m seems more conceivable. Such depths are at part of a small mosasaur (confirmation of cannibalism
any rate sufficient to produce problems such as the first intimated by R U S S E L L , 1967), the marine teleost
bends and would not invalidate M A R T I N & R O T H - Bananogmius, a shark, and part of the Cretaceous
 The Tylosaurine Mosasaurs from Europe a n d Africa 185

diving bird Hesperornis. There is also record of turtle narial border. I feel it necessary t o add that a variation
bones bones in the gut cavity of Hainosaurus bernardi in presacral vertebral numbers particularly in the pygal
(DOLLO, 1891). region is not u n c o m m o n to the same species. I have
noted this in specimens of Mosasaurus lemonnieri (in
which there are a number of postcranial skeletons)
Concluding remarks where the pygal number m a y be anything between 12
and 22. Because of such differences it is important that
The main distinction between Hainosaurus pembinensis the large quantities of tylosaurine material in particu-
and Tylosaurus and similarity with H. bernardi appears larly in the U . S . A . , are examined closely.
to be in the larger number of precaudal vertebrae in the
genus Hainosaurus. I must point out though that H.
pembinensis lacks some of the diagnostic characters of Acknowledgements
the European Hainosaurus, for instance the double but
tressed premaxillary/maxillary suture ( N I C H O L L S , pers. I would like to express my thanks to M. Benton (Bristol) and D. Rus-
comm.) a n d prefrontal that forms part of the external sell (Ottawa) for reviewing this manuscript.

References

BARDET, N., 1990. Première mention du genre Hainosaurus DOLLO, L., 1891. La Vie au Sein des Mers. Paris, Librairie L¬
(Squamata, Mosasauridae) de France. Comptes Rendus de B. Baillière et fils. 304 pp.
l'Académie des sciences, Paris, 311, 2: 751-756. DOLLO, L., 1904. Les mosasauriens de la Belgique. Bulletin
BROOM, R., 1912. On a species of Tylosaurus from the Upper de la Société Belge de Géologie, de Paléontologie et d'Hydro-
Cretaceous of Pondoland. Annals of the South African logie, 18, mémoires: 207-216.
Museum, 1 : 332-333. DOLLO, L. 1913. Globidens fraasi, mosasaurien mylodonte
CAMP, C. L., 1942. California mosasaurs. Memoirs of the nouveau du Maestrichtien (Crétacé supérieur) du Limbourg, et
University of California, 13: 68pp. l'ethologie de la nutrition chez les mosasauriens. Archives de
Biologie, 28: 609-626.
COPE, E. D . , 1869-1870. Synopsis of the extinct Batrachia,
ESTES, R., D E QUEIROZ, K . & GAUTHIER, L , 1988. Phyloge-
Reptilia, and Aves of North America. Transactions of the
American Philosophical Society, n.s. (issued in parts), netic relations within Squamata. 15-98. In ESTES, R. & PRE-
1(1869), 1-105; 2(1870), 106-235; 3(1970), i-vii: 236-252. GILL, G . (eds.). Phylogenetic relationships of the lizard
families: essays commemorating CHARLES L. CAMP. Stanford
COPE, E . D . , 1870. On some Reptilia of the Cretaceous For- University Press, Stanford, California.
mation of the United States. Proceedings of the American Phi-
HUENE, F. von, 1910. Über einem Platecarpus in Tübingen.
losophical Society, 11: 271-274.
Neues Jahrbuch für Mineralogie, Geologie und Paläontologie,
COPE, E . D . 1872. (Remarks on discoveries recently made by 6: 48-50.
Prof. O. C. Marsh). Proceedings of the Philadelphia Academy
LECLERCQ, F. & BOUKO, P . , 1985. La Malogne. Faculté Poly-
of Natural Sciences, 24: 140-141.
technique Möns. 63 pp.
CORROY, G . , 1927. Sur la présence d'ossements de mosasaure
LINGHAM-SOLIAR, T., 1991a. Locomotion in Mosasaurs.
dans le Santonien de l'Aude. Bulletin de la Société géologique
Modern Geology, 16: 229-248.
de France, (4), 27: 135-137.
LINGHAM-SOLIAR, T., 1991b. Mosasaurs from the Upper Cre-
DEPERET, C , & Russo, P. 1925. Les phosphates de Melgou
taceous of the Republic of Niger. Palaeontology, 34 (3):
(Maroc) et leur faune de mosasauriens et de crocodiliens. Bul- 653-670.
letin de la Société géologique de France, (5), 4: 329-346.
LINGHAM-SOLIAR, T., 1991c. Prédation in mosasaurs - a
DOLLO, L., 1885a. Le Hainosaure. Revue des Questions functional approach. In Natural Structures, Principles, Strate-
Scientifiques, 18: 285-289. gies and Models in Architecture. Proceedings of SFB 230,
DOLLO, L., 1885b. Notes d'ostéologie herpétologique. Anna- Heft 6 (1), 169-177.
les de la Société Scientifique de Bruxelles, 9 (2): 309-338. LINGHAM-SOLIAR, T., In press a. A new mode of locomotion
DOLLO, L., 1885c. Première note sur le hainosaure. Bulletin in mosasaurs - subaqueous flight in Plioplatecarpus marshi.
du Musée d'Histoire naturelle de Belgique, 4: 25-35. Journal of Vertebrate Paleontology.
DOLLO, L., 1887. Le hainosaure et les nouveaux vertébrés fos- LINGHAM-SOLIAR, T. In press b. The mosasaur "Angolasau-
siles du Musée de Bruxelles. Revue des Questions Scientifi- rus" Antunes 1964, re-interpreted as the oldest member of the
ques. 1st séries, 22: 70-112. genus Platecarpus. Palaeontology.
DOLLO, L., 1889. Première note sur les mosasauriens de Mes¬ L I N G H A M - S O L I A R , T. & NOLF, D . 1989. The mosasaur
vin. Bulletin de la Société Belge de Géologie, de Paléontologie Prognathodon from the Upper Cretaceous of Belgium (Repti-
et d'Hydrologie, Mémoires 3: 271-304. lia, Mosasauridae). Bulletin de l'Institut Royal des Sciences
Naturelles de Belgique, 59: 137-190.
DOLLO, L. 1890. Première note sur les mosasauriens de Maes-
tricht. Bulletin de la Société Belge de Géologie, de Paléontolo- MARSH, O. C , 1872a. On the structure of the skull and limbs
gie et d'Hydrologie, 4: 151-169. in mosasauroid reptiles, with descriptions of new genera and
 186 Theagarten LINGHAM-SOLIAR

species. American Journal of Science, (3), 3 (18): 448-464. SUZUKI, S., 1985. A new species of Mosasaurus (Reptilia,
MARSH, O. C , 1872b. Note on Rhinosaurus. American Jour- Squamata) from the Upper Cretaceous Hakobuchi Group in
nal of Science. (3), 4 (20): 147. Central Hokkaido, Japan. Bulletin of Hobetsu Museum, 12
(30): 45-66.
MARSH, O. C , 1876. Recent discoveries of extinct animals.
American Journal of Science, (3), 12 (67): 59-61. TELLES-ANTUNES, M., 1964. O Neocretcico e o Cenozoico do
MARTIN, J . E. & BJORK, P. R., 1987. Gastric residues associa- litoral de Angola; 1 Estratigrafía; Reptéis. Junta de Investiga-
ted with the mosasaur from the Late Cretaceous (Campanian) cóes do Ultramar, Lisboa, 257pp.
Pierre Shale, South Dakota. Dakoterra 3: 68-70. THEVENIN, A., 1896. Mosasauriens de la Craie Grise de Vaux
MARTIN, L. D . & ROTHSCHILD, B . M., 1989. Paleopathology
Eclusier prés Peronne (Somme). Bulletin de la Société géologi¬
que de France, (3), 24: 900-916.
and diving mosasaurs. American Scientist, 77: 460-467.
MASSARE, J . A., 1988. Swimming capabilities of Mesozoic THURMOND, J.T., 1969. Notes on mosasaurs from Texas.
Marine reptiles: implications for method of predation. Paleo- Texas Journal of Science, 21 (1): 69-80.
biology, 14 (2): 187-205. UNDERWOOD, G., 1957. Lanthanotus and the anguinimor-
MCDOWELL, S. B . J R . & BOGERT, C. M., 1954. The systema-
phan lizards : A critical review. Copeta, 1: 20-30.
tic position of Lanthanotus and the affinities of the Anguino- WATSON, L., 1988, Whales of the world. Hutchinson. Lon-
morph lizards. Bulletin of the American Museum of Natural don, 302 pp.
History, 105(1): 1-142. WEBB, P. W., 1984. Form and function in fish swimming.
MERRIAM, J . C , 1894. Ueber die Pythonomorphen der Kan- Scientific American, 252: 72-82.
sas Kreide. Palaeontographica, 41: 1-39. WILLISTON, S. W., 1895. New or little known extinct verte-
NICHOLLS, E. L., 1988. The first record of the mosasaur Hai- brates. Kansas University Quarterly, 3: 165-176.
nosaurus (Reptilia, Lacertilia) from North America. Canadian WILLISTON, S. W., 1897. Range and distribution of the mosa-
Journal of Earth Sciences, 25: 1564-1570. saurs. Kansas University Quarterly, 6: 177-189.
OWEN, R., 1840-1845. Odontography. London, 2 volumes. WILLISTON, S. W. 1898. Mosasaurs. University of Kansas
OWEN, R., 1879. On the occurrence in North America of rare Geological Survey Bulletin, 4: 83-221.
extinct vertebrates found fragmentarily in England. Annals WILLISTON, S. W., 1925. The osteology of reptiles. Cam-
and Magazine of Natural History, (5), 4: 53-61. bridge. 300 pp.
OWEN, R., 1880. On the occurrence in North America of rare WIMAN, C , 1920. Some reptiles from the Niobrara group in
extinct vertebrates found fragmentarily in England. Annals Kansas. Bulletin of the Geological Institution of the University
and Magazine of Natural History (5), 5: 177-181. of Uppsala, 18: 9-18.
PERSSON, P. O., 1959. Reptiles from the Senonian of Scania.
Arkivför mineralogi och geologi, Stockholm, 2 (5): 431-480. T. LINGHAM-SOI.IAR
POMPECKJ, J . F. 1910. Über einem Fund von Mosasaurier¬ School of Animal and
Resten im Ober-Senon von Haldem. Jahresbericht Nieder- Microbial Sciences
sächsischen Geologischen Vereins. 3, 122-140. University of Reading
Whiteknights
RUSSELL, D . A., 1967. Systematics and Morphology of Ame- P.O. Box 228
rican Mosasaurs. Peabody Museum of Natural History. Yale Reading RG6 2AJ
University. Bulletin, 23: 237 pp. U.K.
SOLIAR, T., 1988. The mosasaur Goronyosaurus from the
Upper Cretaceous of Sokoto State, Nigeria. Palaeontology, 31 Typescript submitted: Sept. 15, 1991
(3): 747-762. Corrected typescript accepted: Dec. 20, 1991
 PLATE 1

Hainosaurus bernardi (IRSNB 1564). A, left lateral view; B, dorsal view.

 188 Theagarten LINGHAM-SOLIAR

PLATE 2

Hainosaurus bernardi (IRSNB 3672). A, right lateral view of skull; B, dorsal view.

PLATE 3

Hainosaurus bernardi (IRSNB 3672). A: ventral and dorsal views of P O F and portion of squamosal; B, dorsal view of vomers;
C, ventral view of vomers; D, medial and lateral views of splenio angular articulation; E, dorsal and ventral views of pterygoid!
 190 Theagarten LINGHAM-SOLIAR

PLATE 4

Hainosaurus bernardi (IRSNB 3672). Basioccipital. A, right lateral view; B, dorsal view.

PLATE 5

Hainosaurus bernardi (IRSNB 3672). Vertebrae. Caudals. A, B, posterior and lateral views. Cervicals C posterior and lateral
views. Posterior caudal. D, posterior and lateral views. Axis. E, posterior and lateral views. Pygal. F, posterior and anterior views.
 192 Theagarten LINGHAM-SOLIAR

PLATE 6

Cervical and thoracic region of Hainosaurus bernardi (IRSNB R23) showing vertebrae, ribs, scapula, coracoid and paddle.

PLATE 7

Hainosaurus bernardi (MNHN 1896-15) (= Mosasaurus gaudryi, Thevenin, 1896). A, detail of maxilla and dentary showing striae
on teeth; B, double buttressing of the premaxillary/maxillary suture evident despite erosion.
 194 Theagarten LINGHAM-SOLIAR

PLATE 8

Hainosaurus sp. (USTL BUG 2-28, SL 1-10, Jou 5-12). Vertebrae. Caudals.