GSP Marine Meat

D I A P S I DS
SMALL TO GIGANTIC TETRAPODS FROM THE

L AT E PA L E O Z O I C T O M O D E R N T I M E S , G L O B A L
ANATOMICAL CHARACTERISTICS Extremely variable. Two temporal

openings in the skull, often lost by partial opening or by closure. Ribs
usually swept backward.
HABITAT AND HABITS Extremely variable, marine to fully terrestrial to
highly aerial, highly herbivorous to archpredatory.
NOTES One of the three great tetrapod groups, the others being the
extinct anapsids, and the synapsids that include mammals.
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NEODIAPSIDS
SMALL TO GIGANTIC DIAPSIDS FROM THE LATE
PALEOZOIC TO MODERN TIMES, GLOBAL
HABITAT AND HABITS Extremely variable, marine to fully terrestrial

to highly aerial, from highly herbivorous to archpredatory.
NOTES The great bulk of diapsids.
ASKEPTOSAUROIDS
T H A L AT T O S A U R S
SMALL TO LARGE NEODIAPSIDS FROM SMALL TO MEDIUM-SIZED THALATTOSAURS
THE MIDDLE TO LATE TRIASSIC OF THE FROM THE MIDDLE TO LATE TRIASSIC OF
NORTHERN HEMISPHERE EURASIA
ANATOMICAL CHARACTERISTICS Fairly uniform. Skulls ANATOMICAL CHARACTERISTICS Necks moderate in

akinetic, heads small to moderate sized, nostrils small, set length.
just before orbits, eyes face at least somewhat upward,
scleral rings often present, lateral temporal opening at ENDENNASAURIDS
least partly open at bottom, coronoid projection on lower
jaw, teeth range from full set to absent, may be present on SMALL ASKEPTOSAUROIDS FROM
mouth roof. Necks very short to fairly long. Trunks THE LATE TRIASSIC OF EUROPE
moderately long, not very broad, gastralia present. Tails
exceptionally long, highly flattened side to side along ANATOMICAL CHARACTERISTICS Heads subtriangular,
most of length, straight and not expanded aft. Pelvis snouts pointed, toothless. Neck length moderate. Trunk
attached to vertebral column. Limbs short, modified into moderately broad. Shoulder and pelvic girdles fairly well
stiff-jointed paddles, fingers and toes present but not developed, limbs including fingers and toes not very
highly elongated, presumably webbed. Moderately short.
hydrodynamically streamlined, primarily axial undulators
of anguilliform grade, limbs primarily for stability and Endennasaurus acutirostris
maneuvering, pressed tight to body when hydrocruising. 1 m (3 ft) TL, 1.5 kg (3 lb)
HABITAT Coastal and brackish shorelines, lagoons, FOSSIL REMAINS Skull and majority of skeleton severely
reefs, estuaries, freshwaters. flattened top to bottom, partial skeleton.
HABITS Shallow-water ambush and pursuit fishers of ANATOMICAL CHARACTERISTICS Head medium sized.
small to medium-sized game. Possibly bred and probably Hand smaller than foot.
nested on beaches. AGE Late Triassic; Norian.
NOTES Relationships relative to other basal neodiapsids DISTRIBUTION AND FORMATIONS Italy; Zorzino
uncertain. Absence from Southern Hemisphere may Limestone.
reflect lack of sufficient sampling. HABITAT Island nearshore.
ASKEPTOSAURIDS
MEDIUM-SIZED ASKEPTOSAUROIDS FROM
THE MIDDLE TO LATE TRIASSIC OF EURASIA
ANATOMICAL CHARACTERISTICS Heads subtriangular,

not large, coronoid projection not large, teeth subconical.
Neck moderately long. Shoulder and pelvic girdles
somewhat reduced, same for arms and legs, with short
Askeptosaurus (Askeptosauroids) fingers and toes.
68

T H A L A T T O S A U R S
Askeptosaurus italicus AGE Middle Triassic; latest Anisian and earliest Ladinian.
3.9 m (13 ft) TL, 85 kg (180 lb) DISTRIBUTION AND FORMATIONS Switzerland-Italy
FOSSIL REMAINS Several skulls and skeletons. border; Besano.
ANATOMICAL CHARACTERISTICS Head somewhat HABITAT Island nearshore.
elongated, fairly narrow, teeth fairly sharp. Shoulder and NOTES Shared its habitat with Helveticosaurus,
pelvic girdles not strongly reduced, hand smaller than foot. Paraplacodus, Serpianosaurus, Wimanius, Besanosaurus,
unnamed genus buchseri, Mixosaurus cornalianus,
M.? kuhnschnyderi.
Askeptosaurus italicus
Askeptosaurus italicus
69

N E O D I A P S I D S
Anshunsaurus wushaensis
Anshunsaurus wushaensis DISTRIBUTION AND FORMATIONS Southeast China;

3.1 m (10 ft) TL, 35 kg (80 lb) upper Falang.
FOSSIL REMAINS Nearly complete skull and skeleton. HABITAT Continental nearshore.
ANATOMICAL CHARACTERISTICS Head rather small, HABITS May include A. huangnihensis. Shared its habitat
nostrils very small, teeth fairly sharp. Shoulder and pelvic with Yunguisaurus, Qianichthyosaurus zhoui,
girdles reduced, as are arm and leg. Hand and foot similar Guanlingsaurus, Guizhouichthyosaurus, unnamed genus
in size. orientalis, Miodentosaurus.
AGE Middle Triassic; late Ladinian.
DISTRIBUTION AND FORMATIONS Southeast China; Miodentosaurus brevis
middle Falang. 5 m (16 ft) TL, 190 kg (420 lb)
HABITAT Continental nearshore. FOSSIL REMAINS Two skulls and skeletons.
NOTES Shared its habitat with Xinpusaurus? xingyiensis. ANATOMICAL CHARACTERISTICS Head short, fairly
May be direct ancestor of A. huangguoshuensis. broad, snout rather short, nostrils very small, lower jaw
fairly deep, coronoid process shallow, modest upward
Anshunsaurus huangguoshuensis projection at front of lower jaw, teeth few and limited to
3.5 m (11 ft) TL, 50 kg (110 lb) front of jaws, rather blunt. Neck stout. Shoulder and
FOSSIL REMAINS Several skulls and skeletons, most pelvic girdles fairly well developed, same for arm and leg.
severely flattened top to bottom. Hand and foot about same size.
ANATOMICAL CHARACTERISTICS Nostrils very small, AGE Late Triassic; early Carnian.
teeth fairly sharp. Shoulder and pelvic girdles reduced, as DISTRIBUTION AND FORMATIONS Southeast China;
are arm and leg. Hand and foot similar in size. upper Falang.
AGE Late Triassic; early Carnian. HABITAT Continental nearshore.
Miodentosaurus brevis
70

T H A L A T T O S A U R S
THALATTOSAUROIDS XINPUSAURIDS
SMALL TO MEDIUM-SIZED THALATTOSAURS SMALL TO MEDIUM-SIZED THALATTOSAUROIDS
FROM THE MIDDLE TO LATE TRIASSIC OF FROM THE MIDDLE TO LATE TRIASSIC OF ASIA
NORTH AMERICA AND ASIA
ANATOMICAL CHARACTERISTICS Snout elongated into
ANATOMICAL CHARACTERISTICS Snouts strongly a spike, lower jaw much shorter than upper, teeth reduced.
modified, coronoid projections fairly large, teeth reduced. Arms and legs reduced, arms smaller than legs.
Necks very short. Shoulder and pelvic girdles strongly
reduced, same for arms and legs. Radius in lower arms Xinpusaurus (Xinpusaurids)
enlarged, projecting forward, fibula in lower leg enlarged,
projecting backward, hands smaller than feet.
NECTOSAURIDS
SMALL THALATTOSAUROIDS FROM THE LATE
TRIASSIC OF NORTH AMERICA
ANATOMICAL CHARACTERISTICS Heads short and Xinpusaurus? xingyiensis

deep, subrectangular because snouts are deep, snout tips 3 m (10 ft) TL, 40 kg (90 lb)
apparently sharply downturned, teeth few and limited to FOSSIL REMAINS Majority of skull and skeleton, severely
front of jaws, teeth medium sized, stout, subconical, and flattened top to bottom.
fairly sharp. ANATOMICAL CHARACTERISTICS Head subtriangular,
snout elongated into toothed slender spike, lower jaw
Nectosaurus halius much shorter than upper.
0.5 m (1.5 ft) TL AGE Middle Triassic; late Ladinian.
FOSSIL REMAINS Partial skulls and skeletons. DISTRIBUTION AND FORMATIONS Southeast China;
ANATOMICAL CHARACTERISTICS As for group. middle Falang.
AGE Late Triassic; late Carnian. HABITAT Continental nearshore.
DISTRIBUTION AND FORMATIONS Northern California; NOTES May not be the same genus and may be a direct
middle Hosselkus Limestone. ancestor of X. suni. Shared its habitat with Anshunsaurus
HABITAT Continental nearshore. wushaensis.
NOTES Not able to estimate mass because of insufficient
skeletal material. Shared its habitat with Thalattosaurus, Xinpusaurus suni
Shastasaurus, Toretocnemus? 1.3 m (4.3 ft) TL, 3.5 kg (8 lb)
FOSSIL REMAINS Several skulls and skeletons, juvenile
to adult.
ANATOMICAL CHARACTERISTICS Head large,
subtriangular, narrow, fairly shallow, most of snout
Xinpusaurus suni
71

Xinpusaurus suni
Concavispina biseridens
elongated into toothless slender spike, orbits large, lower HABITAT Continental nearshore.
jaw much shorter than upper jaw, slightly S curved, upper NOTES Tip of snout broken so length uncertain.
teeth limited to middle of upper jaw, lower teeth not at
tip of lower jaw, otherwise more numerous and along THALATTOSAURIDS
much of lower jaw length, teeth fairly large, subconical,
and fairly sharp. SMALL THALATTOSAUROIDS FROM THE
AGE Late Triassic; Carnian. EARLY AND/OR MIDDLE TO LATE TRIASSIC
DISTRIBUTION AND FORMATIONS Southern China; OF NORTH AMERICA
Xiaowa.
HABITAT Continental nearshore. ANATOMICAL CHARACTERISTICS Head subtriangular,
NOTES X. kohi and X. bamaolinensis are probably growth snout downturned.
stAGEs of this species. Shared its habitat with NOTES Not able to estimate masses because of insufficient
Concavispina. skeletal material.
Concavispina biseridens Agkistrognathus campbelli

3.7 m (12 ft) TL, 75 kg (179 lb) 1.5 m (5 ft) TL
FOSSIL REMAINS Nearly complete skull and skeleton. FOSSIL REMAINS Partial skull.
ANATOMICAL CHARACTERISTICS Head large, robustly ANATOMICAL CHARACTERISTICS Teeth robust.
constructed, snout enlarged into possible spike, lower jaw AGE Early or Middle Triassic.
shorter than upper, heavily built, teeth extremely reduced. DISTRIBUTION AND FORMATIONS British Columbia;
Limbs quite small. Sulphur Mountain.
AGE Late Triassic; Carnian. HABITAT Continental nearshore.
DISTRIBUTION AND FORMATIONS Southern China; NOTES Fossil found downslope of original layer, therefore
Xiaowa. uncertain. Shared its habitat with unnamed genus borealis,
Paralonectes.
Paralonectes merriami
1 m (3 ft) TL
FOSSIL REMAINS Three partial skulls.
ANATOMICAL CHARACTERISTICS Head fairly deep,
teeth modest in size.
Thalattosaurus (Thalattosaurids)
72

H E L V E T I C O S A U R S
AGE Early or Middle Triassic. Clarazia schinzi

DISTRIBUTION AND FORMATIONS British Columbia; 0.75 m (2.5 ft) TL, 0.6 kg (1.5 lb)
Sulphur Mountain. FOSSIL REMAINS Skull and majority of skeleton.
HABITAT Continental nearshore. ANATOMICAL CHARACTERISTICS Head not deep,
NOTES Fossils found downslope of original layer. moderately broad, snout tip downturned slightly, teeth
few in number, limited to jaw tips and middle of jaws,
Unnamed genus borealis front sets small, low, and blunt.
0.75 m (2.5 ft) TL AGE Middle Triassic; late Anisian and/or early Ladinian.
FOSSIL REMAINS Partial skull and minority of skeleton. DISTRIBUTION AND FORMATIONS Switzerland;
ANATOMICAL CHARACTERISTICS Snout fairly unnamed.
elongated, teeth robust. HABITAT Island nearshore.
AGE Early or Middle Triassic. NOTES Shared its habitat with Hescheleria.
DISTRIBUTION AND FORMATIONS British Columbia;
Sulphur Mountain.
HABITAT Continental nearshore.
NOTES Fossil found downslope of original layer. Is
unlikely to be same genus as much later Thalattosaurus.
Clarazia schinzi
Thalattosaurus alexandrae
2 m (6.5 ft) TL
FOSSIL REMAINS Partial skull and skeleton.
ANATOMICAL CHARACTERISTICS Head subtriangular,
narrow, tip of upper jaw bears serrations, upper teeth
limited to middle of upper jaw, low and blunt, lower front
teeth conical spikes, middle teeth low and blunt.
AGE Late Triassic; late Carnian. Hescheleria rubeli
DISTRIBUTION AND FORMATIONS Northern California; 1 m (3 ft) TL
middle Hosselkus Limestone. FOSSIL REMAINS Majority of skull.
HABITAT Continental nearshore. ANATOMICAL CHARACTERISTICS Head short and deep,
NOTES Shared its habitat with Nectosaurus, Shastasaurus, snout tip strongly curved downward, upper teeth limited
Toretocnemus? to tip of snout and middle of upper jaw, lower teeth to tip
of lower jaw astride a toothlike conical projection of the
jaw, upper front teeth small, middle upper teeth short and
blunt, lower front teeth small.
AGE Middle Triassic; late Anisian and/or early Ladinian.
Thalattosaurus alexandrae DISTRIBUTION AND FORMATIONS Switzerland;
unnamed.
HABITAT Island nearshore.
NOTES Not able to estimate masses because of insufficient
skeletal material.
CLARAZIIDS
H E LV E T I C O S A U R S
MEDIUM-SIZED NEODIAPSIDS FROM
SMALL THALATTOSAUROIDS FROM THE MIDDLE TRIASSIC OF EUROPE
THE MIDDLE TRIASSIC OF EUROPE
ANATOMICAL CHARACTERISTICS Skulls akinetic, heads
ANATOMICAL CHARACTERISTICS Heads subtriangular, small, short, and deep, nostrils near tip of snout, teeth large,
snouts not elongated, turned downward, nostrils set long spikes. Neck short, trunk fairly stout. Gastralia present.
halfway between snout tip and orbit, lower jaws stoutly Tails straight and not expanded aft. Pelvis attached to
built, teeth reduced in number. vertebral column, shoulder and pelvic girdles fairly well
NOTES Not able to estimate masses because of insufficient developed, same for arms and legs, which are modified into
skeletal material. stiff-jointed partial hydrofoils but are not full flippers,
73

fingers and toes present, not highly elongated but fairly primarily for stability and maneuvering, pressed tight to
long yet not ossified, presumably webbed. Moderately body when hydrocruising.
hydrodynamically streamlined, both undulatory and HABITAT Island coastal and brackish shorelines.
flipper swimmers. HABITS Either filter feeder on small invertebrates, or
HABITAT Island coastal and brackish shorelines. algae eater by scraping sea bottoms. Possibly bred and
HABITS Shallow-water ambush fishers of small game. probably nested on beaches.
Possibly bred and probably nested on beaches. NOTES Relationships highly uncertain. May be the earliest
NOTES Relationships relative to other basal neodiapsids herbivorous sea reptiles. Absence from at least some other
uncertain. Absence from at least some other seas may seas may reflect lack of sufficient sampling.
reflect lack of sufficient sampling.
Atopodentatus unicus
Helveticosaurus zollingeri 3 m (10 ft) TL, 50 kg (100 lb)
2 m (7 ft) TL, 15 kg (50 lb) FOSSIL REMAINS Several skulls and skeletons, severely
FOSSIL REMAINS Majority of skull and skeleton, badly flattened top to bottom.
damaged. ANATOMICAL CHARACTERISTICS Hand and foot about
ANATOMICAL CHARACTERISTICS As for group. same size.
AGE Middle Triassic; latest Anisian and earliest Ladinian. AGE Middle Triassic; middle Anisian.
DISTRIBUTION AND FORMATIONS Switzerland; Besano. DISTRIBUTION AND FORMATIONS Southern China;
NOTES Shared its habitat with Askeptosaurus, middle Guanling.
Paraplacodus, Serpianosaurus, Wimanius, Besanosaurus, HABITAT Continental nearshore. Shared its habitat with
unnamed genus buchseri, Mixosaurus cornalianus, Dianopachysaurus, Dinocephalosaurus.
M.? kuhnschnyderi.
SAUROPTERYGIANS
SMALL TO GIGANTIC NEODIAPSIDS FROM Atopodentatus unicus
THE MIDDLE TRIASSIC TO THE END OF
THE MESOZOIC, GLOBAL
ANATOMICAL CHARACTERISTICS Highly variable.

Skulls akinetic, scleral rings often present. Gastralia
present. Tails straight and aft vertebrae not expanded. PLACODONTIFORMES
Limbs at least partial stiff-jointed hydrofoils.
HABITAT AND HABITS Very variable, fully aquatic to SMALL TO MEDIUM-SIZED SAUROPTERYGIANS
fully terrestrial, highly herbivorous to archpredatory. FROM THE MIDDLE TO LATE TRIASSIC OF
EURASIA
ATOPODENTATIANS ANATOMICAL CHARACTERISTICS Highly variable.

Robustly built. Heads short and at least fairly broad,
MEDIUM-SIZED SAUROPTERYGIANS
FROM THE MIDDLE TRIASSIC OF ASIA Henodus (Placodontiformes)
ANATOMICAL CHARACTERISTICS Heads small, snout

forms a hammerhead much broader than main skull,
which is itself moderately broad, nostrils near tip of snout,
teeth very numerous, slender pegs along front rim of
hammerhead. Necks moderately long. Trunks fairly broad.
Pelvis attached to vertebral column, shoulder and pelvic
girdles fairly well developed, same for arms and legs,
which are modified into stiff-jointed paddles, fingers and
toes present but not highly elongated, presumably
webbed. Moderately hydrodynamically streamlined,
primarily axial undulators of anguilliform grade, limbs
74

S A U R O P T E R Y G I A N S
PLACODONTOIDS
SMALL TO MEDIUM-SIZED PLACODONTIFORMES
FROM THE MIDDLE TO LATE TRIASSIC OF
EUROPE
Paraplacodus muscle study

ANATOMICAL CHARACTERISTICS Heads small,
subtriangular, teeth at tips of jaws procumbent, large,
orbits face partly upward, coronoid process large. If present conical. Trunks not extremely broad, fairly deep. Armor if
at least some teeth form a flattened pavement, teeth often present limited to dorsal midline where it forms a serrated
present on mouth roof. Necks short. Trunks compact, ridge. Primarily axial undulators.
moderately to extremely broad. Tails moderately long.
Arms and legs rather short, narrow, modified into partial PARAPLACODONTIDS
hydrofoils but not full flippers, fingers and toes present
but not highly elongated, presumably webbed. Not highly SMALL PLACODONTOIDS FROM THE MIDDLE
hydrodynamically streamlined. TRIASSIC OF EUROPE
HABITAT Coastal and brackish shorelines, lagoons, reefs,
estuaries. ANATOMICAL CHARACTERISTICS Only mouth roof
HABITS Combination of tooth pavements and powerful teeth are flat plates.
jaw muscles anchored in large temporal region indicate
were primarily hard shellfish crushers, a few were filter Paraplacodus broilli
feeders. Some researchers argue that at least some were 1.25 m (4 ft) TL, 4 kg (12 lb)
manatee-like herbivores. Possibly bred and probably FOSSIL REMAINS Skull and minority of skeleton, partial
nested on beaches. skull and majority of skeleton.
NOTES Absence from at least some other seas may reflect ANATOMICAL CHARACTERISTICS Head medium sized,
lack of sufficient sampling. orbits large, teeth at tips of jaws very large, teeth at midjaws
Paraplacodus broilli
75

conical, very stout, blunt. Trunk robust. Tail very long. DISTRIBUTION AND FORMATIONS Netherlands; lower
Limbs large. Muschelkalk.
AGE Middle Triassic; late Anisian and early Ladinian. HABITAT Island nearshore.
DISTRIBUTION AND FORMATIONS Northern Italy; NOTES Palatodonta bleeker may be juvenile of this
Besano. species. Shared its habitat with Nothosaurus marchicus,
HABITAT Island nearshore. Lariosaurus vosseveldensis.
NOTES Shared its habitat with Askeptosaurus,
Helveticosaurus, Serpianosaurus, Wimanius, Placodus gigas
Besanosaurus, unnamed genus buchseri, Mixosaurus 3.2 m (10.5 ft) TL, 100 kg (220 lb)
cornalianus, M.? kuhnschnyderi. FOSSIL REMAINS Skulls and skeletons.
ANATOMICAL CHARACTERISTICS Head broad. Trunk
PLACODONTIDS robust. Tail length moderate. Shoulder and pelvic girdles
reduced, hand and foot same size.
SMALL TO MEDIUM-SIZED PLACODONTOIDS AGE Middle Triassic; late Anisian.
FROM THE MIDDLE TRIASSIC OF EUROPE DISTRIBUTION AND FORMATIONS Germany,
Netherlands, France; upper Muschelkalk.
ANATOMICAL CHARACTERISTICS Heads subtriangular, HABITAT Island nearshore.
very heavily built, nostrils halfway between snout tip and NOTES Shared its habitat with Nothosaurus mirabilis,
orbits, lower jaw deep, teeth at tips of jaws procumbent, N. giganteus, Pistosaurus, Phantomosaurus.
conical, large, teeth at midjaws flat, forming pavement.
Pararcus diepenbroeki
CYAMODONTOIDS
1.5 m (5 ft) TL, 10 kg (20 lb) SMALL TO MEDIUM-SIZED PLACODONTIFORMES
FOSSIL REMAINS Minority of skeleton, possible FROM THE MIDDLE TO LATE TRIASSIC OF
juvenile skull. EURASIA
ANATOMICAL CHARACTERISTICS Insufficient
information. ANATOMICAL CHARACTERISTICS Turtle-like, with
AGE Middle Triassic; early Anisian. flattened, very broad trunk. Hornlets adorn temporal
Placodus gigas
76

Placodus gigas
region of skull. Ribs not swept backward. Tails not very Main trunk and distinct pelvic carapaces present. Tail
long. Poorly hydrodynamically streamlined, propulsion armored by dorsal and lateral rows of hornlets, tail tip bare.
and maneuvering provided by tails and/or appendages.
NOTES Severe top-to-bottom flattening of all specimens Cyamodus tarnowitzensis
compromises mass estimates. Adult size uncertain
FOSSIL REMAINS Partial skull, possibly immature.
CYAMODONTIDS ANATOMICAL CHARACTERISTICS Insufficient
information.
SMALL CYAMODONTOIDS FROM THE MIDDLE AGE Middle Triassic; middle Anisian.
TO LATE TRIASSIC OF EURASIA DISTRIBUTION AND FORMATIONS Southern Germany;
lower Muschelkalk.
ANATOMICAL CHARACTERISTICS Heads large, HABITAT Island nearshore.
deep, very broad at back because temporal region is very NOTES Shared its habitat with Tholodus, Contectopalatus.
expanded, aft rim lined by hornlets, all teeth very blunted. May be direct ancestor of Cyamodus rostratus.
77

Cyamodus rostratus Cyamodus kuhnschnyderi

0.9 m (3 ft) TL, 5 kg (11 lb) 1.5 m (5 ft) TL, 25 kg (60 lb)
FOSSIL REMAINS Skulls and skeletons. FOSSIL REMAINS Skulls and skeletons.
ANATOMICAL CHARACTERISTICS 7 pairs of upper ANATOMICAL CHARACTERISTICS Front teeth further
teeth, 4 pairs of lower teeth. reduced, 5 pairs of upper teeth, 3 pairs of lower teeth.
AGE Middle Triassic; late Anisian. AGE Middle Triassic; middle Ladinian.
DISTRIBUTION AND FORMATIONS Southern DISTRIBUTION AND FORMATIONS Southern
Germany; middle Muschelkalk. Germany; upper Muschelkalk.
HABITAT Island nearshore. HABITAT Island nearshore.
NOTES May be direct ancestor of C. munsteri.
Unnamed genus? orientalis
Cyamodus munsteri 1.5 m (5 ft) TL, 20 kg (45 lb)
1.4 m (4.5 ft) TL, 22 kg (50 lb) FOSSIL REMAINS Skull and majority of skeleton.
FOSSIL REMAINS Several skulls and skeletons. ANATOMICAL CHARACTERISTICS Aft rim of temporal
ANATOMICAL CHARACTERISTICS 7 pairs of upper teeth, region not lined by hornlets, tooth rows short with
3 pairs of lower teeth. Main and pelvic carapace profiles just 4 pairs of upper teeth, 2 pairs of lower teeth.
rounded in top view, carapace pavements made of small Main carapace profile somewhat squarish in top view,
irregular platelets, rimmed by modest-sized hornlets consists of medium-sized irregular plates, pelvic carapace
forming a serrated edge. Hand somewhat smaller than foot. irregularly formed, neither lined by hornlets. Hand
AGE Middle Triassic; early Ladinian. and foot similar in size.
DISTRIBUTION AND FORMATIONS Southern Germany, AGE Late Triassic; early Carnian.
Switzerland, northern Italy?; upper Muschelkalk, lower DISTRIBUTION AND FORMATIONS
Meride Limestone, Besano? Southeastern China; upper Falang.
HABITAT Island nearshore. HABITAT Continental nearshore.
NOTES Probably includes C. hildegardis. Shared its NOTES Placement in earlier
habitat with Neusticosaurus, Ceresiosaurus. May be direct and European Cyamodus
ancestor of Cyamodus kuhnschnyderi. highly problematic.
Cyamodus
rostratus
Cyamodus munsteri
78

Shared its habitat with Anshunsaurus huangguoshuensis, PLACOCHELYIDS

Miodentosaurus, Yunguisaurus, Qianichthyosaurus zhoui,
Guanlingsaurus, Guizhouichthyosaurus. MEDIUM-SIZED CYAMODONTOIDS FROM THE
MIDDLE TO LATE TRIASSIC OF EUROPE
Macroplacus raeticus
1 m (3.3 ft) TL, 8 kg (20 lb) ANATOMICAL CHARACTERISTICS Heads not large,
FOSSIL REMAINS Partial skull. broad, somewhat flattened, snout square-tipped,
ANATOMICAL CHARACTERISTICS Insufficient toothless, elongated, and narrow, nostrils immediately
information. in front of orbits, temporal region expanded, all teeth
AGE Late Triassic; early Rhaetian. flat plates at middle of jaws. No hornlets rim carapaces.
DISTRIBUTION AND FORMATIONS Southern Germany; NOTES Although the skulls of the two known genera are
Koessen. very similar, the major differences in their skeletons may
HABITAT Island nearshore. warrant placement in different families.
Psephoderma alpinum
2.4 m (8 ft) TL, 95 kg (200 lb)
FOSSIL REMAINS Skulls and skeletons.
ANATOMICAL CHARACTERISTICS Head moderately
broad, nostrils very small, aft rim of temporal region
bears a hornlet, lower jaw rather shallow, 3 pairs of upper
and lower teeth. Main trunk and distinct pelvic carapaces
present, main carapace profile fairly rounded in top view,
pelvic carapace subrectangular in top view, platelets of
Cyamodus both modest in size, those of first fairly regular in pattern,
kuhnschnyderi three subtle fore-and-aft ridges present along midline,
neither carapace lined by hornlets. Tail moderately long,
adorned by dorsal and lateral rows of hornlet armor, tail
tip bare. Arms and hands smaller than legs and feet.
AGE Late Triassic; middle and/or late Norian.
DISTRIBUTION AND FORMATIONS Italy; Zorzino
Limestone.
Unnamed genus? orientalis
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Psephoderma alpinum
Placochelys placodonta
0.8 m (2.6 ft) TL
FOSSIL REMAINS Complete and partial skulls and
majority of skeleton.
ANATOMICAL CHARACTERISTICS Head very broad
aft because temporal region is very expanded, adorned
Placochelys
with numerous hornlets, lower jaw rather deep, 4 pairs
placodonta
of upper teeth. Only one united carapace present, profile
subrectangular in top view, pavement consists of small
irregular platelets surrounding larger, subconical
osteoderms. Tail short, unarmored. Appendages
are similar size fore and aft.
DISTRIBUTION AND FORMATIONS Hungary;
Physiocardia.
HABITAT Island nearshore. AGE Late Triassic; Carnian or Norian.
NOTES Lack of adequate modern description precludes DISTRIBUTION AND FORMATIONS Spain; unnamed.
accurate skeletal restoration or mass estimate at this time, HABITAT Island nearshore.
whether limbs were paddles or more flipper-like is not clear.
Henodus chelyops
1.4 m (4.5 ft) TL, 40 kg (90 lb)
HENODONTIDS
FOSSIL REMAINS Several skulls and skeletons.
SMALL TO MEDIUM-SIZED CYAMODONTOIDS ANATOMICAL CHARACTERISTICS Head rather small,
FROM THE LATE TRIASSIC OF EUROPE subrectangular in side and top views, moderately deep,
snout broad and square ended, nostrils very near snout
ANATOMICAL CHARACTERISTICS Upper temporal tip, orbits at front end of head face strongly upward,
openings very reduced or closed, teeth very reduced, upper temporal opening closed, lower jaw deep, one pair
main surfaces concave, may have had baleen-like soft of small teeth at front of upper jaw. Carapace extremely
tissues lining lower jaw. One united carapace, armored broad in dorsal view, side rims rounded in top view,
both dorsally and ventrally. platelets fairly well organized, form a pair of subtle fore-
HABITS Filter feeders on small invertebrates and/or algae. and-aft ridges, forward projection of carapace partly
shields neck, no hornlets rimming carapace. Arm and leg
Parahenodus atancensis very reduced, arm a little smaller than leg.
1 m (3.3 ft) TL, 15 kg (35 lb) AGE Late Triassic; early Carnian.
FOSSIL REMAINS Partial skull. DISTRIBUTION AND FORMATIONS Southern Germany;
ANATOMICAL CHARACTERISTICS Head subtriangular in lower Keuper.
top view, upper temporal opening small. HABITAT Coastal nearshore.
80

Henodus chelyops
HABITS Poor locomotion both in water and onshore.
81

Saurosphargis volzi
SAUROSPHARGIDS 0.6 m (2 ft) TL
SMALL TO MEDIUM-SIZED SAUROPTERYGIANS FOSSIL REMAINS Partial skeleton.
FROM THE MIDDLE TO LATE TRIASSIC OF ASIA ANATOMICAL CHARACTERISTICS Turtle-like trunk with
flattened and very broad carapace.
ANATOMICAL CHARACTERISTICS Heads moderate in AGE Middle Triassic; middle Anisian.
size, subtriangular, not deep, nostrils set well back, lower DISTRIBUTION AND FORMATIONS Poland; lower
jaw somewhat shorter than upper, teeth stout, subconical. Muschelkalk.
Necks rather short. Trunks broad, ribcages and gastralia HABITAT Island nearshore.
heavily built. NOTES Specimen destroyed in World War II. Shared its
NOTES Relationships to other basal neodiapsids habitat with Dactylosaurus, Germanosaurus.
uncertain, may be related to thalattosaurs and converged
with placodontiformes. Major differences in their anatomy
may warrant placement of the known genera in different EOSAUROPTERYGIANS
families.
SMALL TO GIGANTIC SAUROPTERYGIANS
Largocephalosaurus polycarpon FROM THE EARLY TRIASSIC TO THE END
2 m (6.5 ft) TL, 40 kg (90 lb) OF THE MESOZOIC, GLOBAL
FOSSIL REMAINS Several skulls and partial skeletons.
ANATOMICAL CHARACTERISTICS Head moderately ANATOMICAL CHARACTERISTICS Fairly variable.
broad, nostrils immediately in front of orbits. Trunk Heads very small to very large, generally subtriangular, not
moderately broad. Tail moderate in length. Arm and leg very broad, somewhat flattened with orbits facing partly
moderately large. Lightly armored. upward, nostrils not near tip of snout, lower jaw not deep,
AGE Middle Triassic; middle Anisian. full set of teeth always present, conical, not present on
DISTRIBUTION AND FORMATIONS Southern China; mouth roof, intermeshed if long. Neck, trunk, and front
upper Guanling. tail vertebrae fairly uniform in structure within a species.
HABITAT Coastal nearshore. Neck not short, sometimes extremely long. Trunks not
NOTES May include L. qianensis. Shared its habitat with very broad, gastralia well developed. Tails not long, straight
Panzhousaurus, Wumengosaurus, Nothosaurus and not expanded aft. Scapula well forward on elongated,
yangjuanensis, Xinminosaurus, Barracudasauroides, broad shoulder girdle complex, blade does not strongly
Sinosaurosphargis. overlap chest ribs. Moderately hydrodynamically
streamlined, propulsion provided by limbs modified into
hydrofoils, with aft limb sometimes somewhat less powerful
than fore, tail primarily for stability and maneuvering.
Largocephalosaurus HABITAT Freshwaters to deep oceans.
polycarpon HABITS Swimming performance mediocre to good.
Predaceous, most ambush and/or pursuit fishers and
hunters of small to big game, some crushers, some filter
feeders. Basal examples may have bred and nested on
beaches, those with full flippers reproduced aquatically.
Sinosaurosphargis yunguiensis
1 m (3.3 ft) TL, 10 kg (22 lb)
PACHYPLEUROSAURS
FOSSIL REMAINS Skull and partial skeletons, severely SMALL TO MEDIUM-SIZED EOSAUROPTERYGIANS
flattened top to bottom. FROM THE EARLY TO MIDDLE TRIASSIC OF
ANATOMICAL CHARACTERISTICS Head very broad aft. EURASIA
Turtle-like trunk with flattened and very broad carapace,
profile rounded in top view, pavement consists of very ANATOMICAL CHARACTERISTICS Temporal region and
large number of small ossicles, no hornlets along rim. upper temporal opening not enlarged, constricted in top
AGE Middle Triassic; middle Anisian. view. Tail base broad.
DISTRIBUTION AND FORMATIONS Southern China; HABITS Swimming performance modest. Ambush and
upper Guanling. pursuit predators of small to medium-sized game.
HABITAT Coastal nearshore. NOTES Appear to include smallest known sea reptiles.
82

Dianopachysaurus dingi
0.2 m (0.7 ft) TL, 0.02 kg (0.05 lb)
FOSSIL REMAINS Skull and majority of skeleton, severely
flattened top to bottom.
ANATOMICAL CHARACTERISTICS Skull fairly broad,
orbits set well forward, upper temporal opening small,
Keichousaurus (Pachypleurosaurs) teeth small. Arm smaller than foot.
AGE Middle Triassic; middle Anisian.
DISTRIBUTION AND FORMATIONS Southern China;
Majiashanosaurus discocoracoidis middle Guanling.
1 m (3 ft) TL, 2.5 kg (5.5 lb) HABITAT Continental nearshore.
FOSSIL REMAINS Majority of skeleton, severely flattened NOTES. Shared its habitat with Atopodentatus,
top to bottom. Dinocephalosaurus. Despite very small size appears to be
ANATOMICAL CHARACTERISTICS Hand as large as foot. an adult, in which case is the smallest known sea reptile
AGE Early Triassic; Olenekian. along with Dactylosaurus.
DISTRIBUTION AND FORMATIONS Eastern China;
upper Nanlinghu. Dactylosaurus gracilis
HABITAT Continental nearshore. 0.2 m (0.7 ft) TL, 0.02 kg (0.05 lb)
NOTES Shared its habitat with Sclerocormus, FOSSIL REMAINS Numerous skulls and skeletons,
Chaohusaurus geishanensis. severely flattened top to bottom.
ANATOMICAL CHARACTERISTICS Upper temporal
Hanosaurus hupehensis fenestra small.
0.7 m (2 ft) TL, 1 kg (2 lb) AGE Middle Triassic; middle Anisian.
FOSSIL REMAINS Partial skull and skeleton, severely DISTRIBUTION AND FORMATIONS Poland; lower
flattened top to bottom. Muschelkalk.
ANATOMICAL CHARACTERISTICS Upper temporal HABITAT Island nearshore.
opening small, slender teeth moderately long. NOTES Shared its habitat with Saurosphargis,
AGE Middle Triassic; Anisian. Germanosaurus.
lower Jialingjiang. Keichousaurus hui
HABITAT Continental nearshore. 2.7 m (9 ft) TL, 50 kg (110 lb)
FOSSIL REMAINS Very large number of skulls and
Panzhousaurus rotundirostris skeletons, adult and juvenile, severely flattened top to
0.7 m (2 ft) TL, 1 kg (2 lb) bottom.
FOSSIL REMAINS Skull and majority of skeleton, severely ANATOMICAL CHARACTERISTICS Head fairly small,
flattened top to bottom. moderately broad, upper temporal opening medium
ANATOMICAL CHARACTERISTICS Head small, snout sized, front teeth long, procumbent, irregularly
fairly broad and rounded, orbits set well forward, upper intermeshing, slender, gently curved spikes. Hand smaller
temporal opening small, teeth large, limited to front half than foot, ulna in lower arm greatly expanded.
of jaws, strongly curved, procumbent. Neck moderately AGE Middle Triassic; late Ladinian.
long. Trunk moderately broad. Hand as large as foot. DISTRIBUTION AND FORMATIONS Southeastern China;
AGE Middle Triassic; middle Anisian. lower Falang.
upper Guanling.
NOTES Shared its habitat with Largocephalosaurus,
Sinosaurosphargis, Wumengosaurus, Nothosaurus
yangjuanensis, Xinminosaurus, Barracudasauroides. Keichousaurus hui
83

HABITAT Continental nearshore. Neusticosaurus pusillus

HABITS Modest degree of sexual dimorphism appears to 0.5 m (1.6 ft) TL, 0.35 kg (0.75 lb)
be present. May have given live birth. FOSSIL REMAINS Numerous skulls and skeletons, most
NOTES The Mesozoic marine reptile perhaps known from severely flattened top to bottom.
the largest number of specimens. Shared its habitat with ANATOMICAL CHARACTERISTICS Head small, fairly
Nothosaurus youngi, Lariosaurus, Qianxisaurus, broad, upper temporal opening extremely small, teeth
Wangosaurus, Qianichthyosaurus. moderate in number, medium sized, stout. Hand smaller
than foot.
Anarosaurus heterodontus AGE Middle Triassic; early Ladinian.
0.5 m (1.6 ft) TL, 0.35 kg (0.75 lb) DISTRIBUTION AND FORMATIONS Switzerland; lower
FOSSIL REMAINS Skulls and skeletons, most severely Meride Limestone.
flattened top to bottom. HABITAT Island nearshore.
ANATOMICAL CHARACTERISTICS Head fairly broad, NOTES Shared its habitat with Ceresiosaurus calcagnii.
upper temporal opening very small, teeth numerous, May be the direct ancestor of N. peyeri.
small, blunt. Hand smaller than foot.
DISTRIBUTION AND FORMATIONS Central Germany;
middle Muschelkalk.
NOTES Shared its habitat with Serpianosaurus
germanicus, Nothosaurus juvenilis.
Anarosaurus
heterodontus
Neusticosaurus
pusillus
Neusticosaurus
peyeri
Neusticosaurus (or Pachypleurosaurus) edwardsi
84

Neusticosaurus peyeri ANATOMICAL CHARACTERISTICS Head small, fairly

0.5 m (1.6 ft) TL, 0.35 kg (0.75 lb) broad, teeth moderate in number, medium sized, stout.
FOSSIL REMAINS Numerous skulls and skeletons, most AGE Middle Triassic; late Ladinian and/or early Carnian.
severely flattened top to bottom. DISTRIBUTION AND FORMATIONS Switzerland;
ANATOMICAL CHARACTERISTICS Head small, fairly Partnach.
broad, upper temporal opening nearly closed, teeth HABITAT Island nearshore.
moderate in number, medium sized, stout. Hand
smaller than foot. Neusticosaurus (or Pachypleurosaurus)
AGE Middle Triassic; early Ladinian. edwardsi
DISTRIBUTION AND FORMATIONS Switzerland; lower 1.25 m (4 ft) TL, 5.5 kg (12 lb)
Meride Limestone. FOSSIL REMAINS Numerous skulls and skeletons, most
HABITAT Island nearshore. severely flattened top to bottom.
NOTES Shared its habitat with Ceresiosaurus lanzi. ANATOMICAL CHARACTERISTICS Head small, fairly
May be the direct ancestor of N. toeplitschi. broad, upper temporal opening very small, teeth moderate
in number, medium sized, stout. Arm longer and more
Neusticosaurus toeplitschi slender than leg, hand smaller than foot.
0.4 m (1.3 ft) TL, 0.18 kg (4 lb) AGE Middle Triassic; middle Ladinian.
FOSSIL REMAINS Numerous skulls and skeletons, DISTRIBUTION AND FORMATIONS Switzerland;
severely flattened top to bottom. middle Meride Limestone.
Neusticosaurus (or
Pachypleurosaurus) edwardsi
85

Serpianosaurus germanicus orbits large, upper temporal fenestra extremely small, teeth
2 m (6 ft) TL, 20 kg (45 lb) numerous and small. Hand almost as large as foot.
FOSSIL REMAINS Partial skeleton. AGE Middle Triassic; latest Anisian and earliest Ladinian.
ANATOMICAL CHARACTERISTICS Insufficient DISTRIBUTION AND FORMATIONS Switzerland; Besano.
information. HABITAT Island nearshore.
AGE Middle Triassic; middle Anisian. NOTES Shared its habitat with Askeptosaurus,
DISTRIBUTION AND FORMATIONS Central Germany; Helveticosaurus, Paraplacodus, Wimanius, Besanosaurus,
middle Muschelkalk. unnamed genus buchseri, Mixosaurus cornalianus,
HABITAT Island nearshore. M.? kuhnschnyderi.
NOTES Shared its habitat with Anarosaurus, Nothosaurus
juvenilis. May be direct ancestor of S. mirigiolensis. Wumengosaurus delicatomandibularis
1.3 m (4 ft) TL, 6 kg (12 lb)
Serpianosaurus mirigiolensis FOSSIL REMAINS Skull and two skeletons.
0.85 m (2.8 ft) TL, 1.7 kg (3.5 lb) ANATOMICAL CHARACTERISTICS Head small, narrow,
FOSSIL REMAINS Numerous skulls and skeletons. snout long, nostrils very small, teeth numerous, small,
ANATOMICAL CHARACTERISTICS Head fairly large, vertical. Arm and leg rather small, hand smaller than foot.
Serpianosaurus
mirigiolensis
Wumengosaurus
delicatomandibularis
86

AGE Middle Triassic; middle Anisian. SIMOSAURIDS

upper Guanling. MEDIUM-SIZED NOTHOSAURS FROM
HABITAT Continental nearshore. THE MIDDLE TRIASSIC OF EUROPE
NOTES Shared its habitat with Largocephalosaurus,
Sinosaurosphargis, Panzhousaurus, Nothosaurus ANATOMICAL CHARACTERISTICS Heads fairly broad,
yangjuanensis, Xinminosaurus, Barracudasauroides. numerous teeth short, blunt.
HABITS Small shellfish crushers.
NOTHOSAURS Simosaurus gaillardoti
SMALL TO LARGE EOSAUROPTERYGIANS FROM 4.2 m (14 ft), 200 kg (440 lb)
THE MIDDLE TRIASSIC OF EURASIA FOSSIL REMAINS Skulls and skeletons.
ANATOMICAL CHARACTERISTICS Head medium sized.
ANATOMICAL CHARACTERISTICS Heads medium sized, AGE Middle Triassic; middle Ladinian.
orbits placed well forward because temporal region DISTRIBUTION AND FORMATIONS France, southern
elongated, sometimes extremely so, nostrils not severely Germany; upper Muschelkalk.
reduced, fairly close to orbits, lower jaws shallow. Neck HABITAT Island archipelago shallows.
moderately long. Trunk moderately long, fairly narrow.
Tail moderately long. Pelvis attached to vertebral column, NOTHOSAURIDS
ventral elements not highly expanded, arms and legs not
full flippers. SMALL TO LARGE NOTHOSAURS FROM
HABITATS Coastal and brackish shorelines, lagoons, reefs, THE MIDDLE TRIASSIC OF EURASIA
estuaries, freshwaters.
HABITS Swimming performance modest. Most ambush ANATOMICAL CHARACTERISTICS Heads narrow,
and pursuit predators of small to medium-sized game, some temporal region and openings very to extremely
crushers. At least some examples probably gave live birth. elongated and constricted in top view, modest number of
NOTES Absence from at least some other seas may reflect teeth, front set long, procumbent, irregularly intermeshing,
lack of sufficient sampling. slender, gently curved spikes, back teeth small spikelets.
HABITS Ambush and pursuit predators of small to
medium-sized game.
Simosaurus gaillardoti
87

NOTES Not able to estimate masses because of insufficient DISTRIBUTION AND FORMATIONS Central Germany;
description of skeletal material. upper Muschelkalk.
HABITAT Island archipelago shallows.
Germanosaurus? schafferi NOTES Inadequate descriptions of remains of this well-
3 m (10 ft) TL known genus preclude accurate skeletal restorations at
FOSSIL REMAINS Partial skulls, severely flattened top to this time. Shared its habitat with Placodus, Pistosaurus,
bottom. Phantomosaurus, N. giganteus.
ANATOMICAL CHARACTERISTICS Temporal region very
elongated. Nothosaurus giganteus
AGE Middle Triassic; early Anisian. 7 m (23 ft) TL
DISTRIBUTION AND FORMATIONS Poland; lower FOSSIL REMAINS Several skulls and partial skeletons,
Muschelkalk. severely flattened top to bottom.
HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Temporal region
NOTES The validity of the generic title is problematic. extremely elongated.
Shared its habitat with Saurosphargis, Dactylosaurus. AGE Middle Triassic; late Anisian.
DISTRIBUTION AND FORMATIONS Central Germany;
Nothosaurus marchicus upper Muschelkalk.
1.2 m (4 ft) TL HABITAT Island archipelago shallows.
FOSSIL REMAINS Several skulls and partial skeletons,
severely flattened top to bottom. Nothosaurus haasi
ANATOMICAL CHARACTERISTICS Temporal region 1.3 m (4 ft) TL
extremely elongated. FOSSIL REMAINS Several skulls and partial skeletons,
AGE Middle Triassic; early and middle Anisian. severely flattened top to bottom.
DISTRIBUTION AND FORMATIONS Central Germany, ANATOMICAL CHARACTERISTICS Temporal region
Netherlands; lower Muschelkalk. extremely elongated.
HABITAT Island archipelago shallows. AGE Middle Triassic; late Anisian and/or early Ladinian.
NOTES Probably includes N. winterswijkensis. Shared its DISTRIBUTION AND FORMATIONS Israel; lower
habitat with Pararcus, Lariosaurus vosseveldensis. May Saharonium.
have been direct ancestor of N. juvenilis. HABITAT Continental shallows.
NOTES May include N. tchernovi. May be same species as
Nothosaurus juvenilis one of the European Nothosaurus of the same age.
2 m (6.5 ft) TL
FOSSIL REMAINS Majority of skull. Nothosaurus cymatosauroides
ANATOMICAL CHARACTERISTICS Temporal region 2.5 m (8 ft) TL
extremely elongated. FOSSIL REMAINS Skull, severely flattened top to bottom.
AGE Middle Triassic; middle and late Anisian. ANATOMICAL CHARACTERISTICS Temporal region
DISTRIBUTION AND FORMATIONS Central Germany; extremely elongated.
middle Muschelkalk. AGE Middle Triassic; late Ladinian.
HABITAT Island archipelago shallows. DISTRIBUTION AND FORMATIONS Northeastern Spain;
NOTES Shared its habitat with Anarosaurus, upper Muschelkalk.
Serpianosaurus germanicus. May have been direct ancestor HABITAT Island archipelago shallows.
of N. mirabilis and/or N. giganteus.
Nothosaurus mirabilis
3 m (10 ft) TL
Nothosaurus
FOSSIL REMAINS Several skulls and partial skeletons, mirabilis
severely flattened top to bottom.
ANATOMICAL CHARACTERISTICS Temporal region
extremely elongated.
AGE Middle Triassic; late Anisian.
88

Nothosaurus yangjuanensis
3.5 m (12 ft) TL
FOSSIL REMAINS A few skulls and skeletons, severely
flattened top to bottom.
ANATOMICAL CHARACTERISTICS Standard for group.
Lariosaurus balsami
upper Guanling.
HABITAT Continental shallows.
NOTES May include N. rostellatus and N. zhangi. May be
same species as one of the European Nothosaurus of the
same age. AGE Middle Triassic; middle to late Ladinian.
NOTES Shared its habitat with Largocephalosaurus, DISTRIBUTION AND FORMATIONS Northern Italy;
Sinosaurosphargis, Panzhousaurus, Wumengosaurus, Perledo-Varenna.
Xinminosaurus, Barracudasauroides. HABITAT Island archipelago shallows.
NOTES May include L. curionii and L. valcersii.
Nothosaurus youngi
2 m (6.5 ft) TL Lariosaurus xingyiensis
FOSSIL REMAINS Majority of two skulls and skeletons, 2.5 m (8 ft) TL
severely flattened top to bottom. FOSSIL REMAINS Skull and majority of skeleton, severely
ANATOMICAL CHARACTERISTICS Standard for genus. flattened top to bottom.
AGE Middle Triassic; late Ladinian. ANATOMICAL CHARACTERISTICS Temporal region very
DISTRIBUTION AND FORMATIONS Southeastern China; elongated.
lower Falang. AGE Middle Triassic; late Ladinian.
HABITAT Continental nearshore. DISTRIBUTION AND FORMATIONS Southeastern China;
NOTES May be same species as one of the western lower Falang.
Nothosaurus of the same age. Shared its habitat with HABITAT Continental nearshore.
Keichousaurus, Qianxisaurus, Wangosaurus,
Qianichthyosaurus, Lariosaurus. Ceresiosaurus calcagnii
3 m (10 ft) TL
Lariosaurus vosseveldensis FOSSIL REMAINS Skulls and skeletons, severely flattened.
5 m (1.5 ft) TL ANATOMICAL CHARACTERISTICS Head rather small.
FOSSIL REMAINS Skull, severely flattened top to bottom, Appendages more flipper shaped.
possibly juvenile. AGE Middle Triassic; early Ladinian.
ANATOMICAL CHARACTERISTICS Temporal region very DISTRIBUTION AND FORMATIONS Switzerland; lower
elongated. Meride Limestone.
AGE Middle Triassic; early Anisian. HABITAT Island archipelago shallows.
DISTRIBUTION AND FORMATIONS Netherlands; lower NOTES Shared its habitat with Neusticosaurus pusillus.
Muschelkalk.
HABITAT Island archipelago shallows. Ceresiosaurus lanzi
NOTES May include L. winkelhorsti. Shared its habitat 3 m (10 ft) TL
with Pararcus, Nothosaurus marchicus. May be a direct FOSSIL REMAINS Skulls and skeletons, flattened.
ancestor of L. balsami. ANATOMICAL CHARACTERISTICS Head fairly large.
Appendages more flipper shaped.
Lariosaurus balsami AGE Middle Triassic; early Ladinian.
1.5 m (5 ft) TL DISTRIBUTION AND FORMATIONS Switzerland; lower
FOSSIL REMAINS Skulls and skeletons, most severely Meride Limestone.
flattened top to bottom. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Temporal region very NOTES Shared its habitat with Neusticosaurus peyeri.
elongated.
89

PAXPLESIOSAURS DISTRIBUTION AND FORMATIONS Nevada; upper Favret.

SMALL TO GIGANTIC EOSAUROPTERYGIANS HABITS Fisher of small game.
FROM THE MIDDLE TRIASSIC TO THE END NOTES Not able to estimate mass because of inadequate
OF THE MESOZOIC, GLOBAL skeletal material. Shared its habitat with unnamed genus
and species, unnamed genus duelferi, unnamed genus
HABITATS Freshwaters to deep oceans. youngorum, Phalarodon, Thalattoarchon.
HABITS Swimming performance mediocre to good. Most
ambush and/or pursuit fishers and hunters of small to big COROSAURIDS
game, some crushers, some filter feeders.
NOTES Eosauropterygians including corosaurs and SMALL PAXPLESIOSAURS FROM THE EARLY
plesiosauriformes and their common ancestor. TRIASSIC OF NORTH AMERICA
PAXPLESIOSAUR MISCELLANEA ANATOMICAL CHARACTERISTICS Head medium sized,

broad, flattened, upper temporal opening large, teeth
short, procumbent spikes. Tail base very broad.
Qianxisaurus chajiangensis HABITATS Primarily coastal shallows.
1 m (10 ft) TL, 3 kg (6 lb) HABITS Swimming performance modest. Ambush and
FOSSIL REMAINS Skull and majority of skeleton, severely pursuit predators of small game.
flattened top to bottom. NOTES Absence from at least some other seas may reflect
ANATOMICAL CHARACTERISTICS Head fairly large and lack of sufficient sampling.
broad, snout fairly long, upper temporal fenestra nearly
closed, teeth numerous, short and stout. Hand smaller Corosaurus alcovensis
than foot. 1.5 m (5 ft) TL, 7 kg (15 lb)
AGE Middle Triassic; late Ladinian. FOSSIL REMAINS Skull and partial skeletons.
DISTRIBUTION AND FORMATIONS Southwestern China; ANATOMICAL CHARACTERISTICS Hand smaller than
lower Falang. foot.
HABITAT Continental nearshore. AGE Early Triassic; late Olenekian.
HABITS Small shellfish crusher. DISTRIBUTION AND FORMATIONS Wyoming; lower
NOTES Shared its habitat with Keichousaurus, Nothosaurus Chugwater.
youngi, Lariosaurus, Qianichthyosaurus, Wangosaurus. HABITAT Interior seaway near shore.
Wangosaurus brevirostris
3 m (10 ft) TL
FOSSIL REMAINS Majority of skull and skeleton.
ANATOMICAL CHARACTERISTICS Head fairly large,
narrow, snout very short because temporal region extremely
Corosaurus alcovensis
elongated. Neck fairly long. Hand smaller than foot.
DISTRIBUTION AND FORMATIONS Southeastern China;
lower Falang.
HABITS Small game fisher.
NOTES Not able to estimate masses because of inadequate
skeletal material. CYMATOSAURIDS
Augustasaurus hagdorni SMALL PAXPLESIOSAURS FROM THE MIDDLE
3 m (10 ft) TL TRIASSIC OF EUROPE
FOSSIL REMAINS Skull and minority of a skeleton,
severely flattened top to bottom. HABITS Swimming performance modest. Ambush and
ANATOMICAL CHARACTERISTICS Snout long, upper pursuit predators of small game.
temporal opening long, front teeth medium-sized spikes. NOTES Absence from at least some other seas may reflect
AGE Middle Triassic; late Anisian. lack of sufficient sampling.
90

Cymatosaurus friedericianus
2 m (6 ft) TL
PISTOSAURIDS
FOSSIL REMAINS Numerous skulls and skeletons, adult SMALL TO MEDIUM-SIZED PLESIOSAURIFORMES
and juvenile, partial and/or severely flattened top to FROM THE MIDDLE TO LATE TRIASSIC OF
bottom. EURASIA
ANATOMICAL CHARACTERISTICS Temporal region very
elongated. ANATOMICAL CHARACTERISTICS Heads not broad,
AGE Middle Triassic; lower Anisian. snout fairly long, nostrils very small, upper temporal
DISTRIBUTION AND FORMATIONS Eastern Germany, opening long, number of teeth modest, procumbent.
western Poland; lower Muschelkalk. NOTES Absence from at least some other seas may
HABITAT Island archipelago shallows. reflect lack of sufficient sampling.
NOTES May include C. latifrons, C. multidentatus, C. HABITAT Coastal shallows and continental shelves.
minor, C. erikae. HABITS Swimming performance modest. Able to dive
to deeper waters. Ambush and pursuit fishers of small
PLESIOSAURIFORMES to medium-sized game. May have given live birth.
SMALL TO GIGANTIC PAXPLESIOSAURS FROM Pistosaurus longaevus

THE MIDDLE TRIASSIC TO THE END OF THE 3 m (10 ft) TL, 100 kg (220 lb)
MESOZOIC, GLOBAL FOSSIL REMAINS Skulls and a skeleton.
ANATOMICAL CHARACTERISTICS Head fairly narrow,
ANATOMICAL CHARACTERISTICS Limbs are clawless all teeth fairly long and strongly built. Neck medium
full flippers with increased number of distal elements. length. Tail medium length. Foreflipper a little smaller
HABITATS Freshwaters to deep oceans. than aft.
HABITS Swimming performance mediocre to good. Most AGE Middle Triassic; late Anisian.
ambush and/or pursuit fishers and hunters of small to big DISTRIBUTION AND FORMATIONS Southern Germany;
game, some filter feeders. Little or no ability to come onto upper Muschelkalk.
land, so reproduced aquatically in most if not all cases. HABITAT Island archipelago shallows.
Thalassiodracon
(Plesiosauriformes)
Microcleidus muscle study
91

Pistosaurus longaevus
Pistosaurus longaevus
92

NOTES The one skeleton has no skull, so whether it AGE Early Jurassic; earliest Hettangian.
belongs to this species is not certain. Shared its habitat DISTRIBUTION AND FORMATIONS Southern England;
with Placodus, Nothosaurus mirabilis, N. giganteus, lower Blue Lias.
Phantomosaurus. HABITAT Island archipelago shallows.
NOTES Shared its habitat with Stratesaurus, Avalonnectes,
Yunguisaurus liae Eurycleidus, Atychodracon, Thalassiodracon,
1.7 m (5.5 ft) TL, 20 kg (45 lb) Protoichthyosaurus.
FOSSIL REMAINS Skull and majority of adult skeleton,
severely flattened top to bottom, juvenile skull and skeleton. PLIOSAUROIDS
ANATOMICAL CHARACTERISTICS Head small, modest
number of teeth, front set long, procumbent, irregularly SMALL TO GIGANTIC PLESIOSAURS FROM
intermeshing, slender, gently curved spikes. Neck and tail THE EARLY JURASSIC TO THE EARLY LATE
fairly long. CRETACEOUS, GLOBAL
AGE Late Triassic; early Carnian.
DISTRIBUTION AND FORMATIONS Southeastern China; ANATOMICAL CHARACTERISTICS Significantly variable.
upper Falang. Necks never extremely long.
HABITAT Continental nearshore. HABITATS Coastal shallows to deep oceans.
NOTES Shared its habitat with Anshunsaurus HABITS Swimming performance modest to good.
huangguoshuensis, Miodentosaurus, Qianichthyosaurus Ambush and/or pursuit fishers and hunters of small to
zhoui, Guanlingsaurus, Guizhouichthyosaurus, unnamed big game.
genus orientalis.
RHOMALEOSAURIDS
PLESIOSAURS
SMALL TO LARGE PLIOSAUROIDS FROM
SMALL TO GIGANTIC PLESIOSAURIFORMES THE EARLY TO MIDDLE JURASSIC OF THE
FROM THE EARLY JURASSIC TO THE END NORTHERN AND SOUTHERN HEMISPHERES
OF THE MESOZOIC, GLOBAL
ANATOMICAL CHARACTERISTICS Uniform. Heads
ANATOMICAL CHARACTERISTICS Fairly variable. Heads medium sized, temporal region expanded. Necks
very small to very large, nostrils small and immediately moderately long.
before orbits. Necks very short to extremely long. Trunk HABITATS Coastal shallows and continental shelves.
compact, gastralia reduced in number and increased in HABITS Swimming performance modest. Ambush and
complexity, robust, central shafts straight, lateral tips pursuit fishers of small to medium-sized game.
swept back. Tail never very long, tip bearing small, either
vertical or horizontal flukes in at least some examples. Anningasaura lymense
Lower pelvic elements enlarged into large, flat plates. Adult size uncertain
Flippers better developed, with shortened lower arm and FOSSIL REMAINS Skull and partial skeleton, juvenile.
increased distal complexes, about as long as trunk, ANATOMICAL CHARACTERISTICS Insufficient
foreflipper at least about as large as aft. information.
HABITATS Freshwaters to deep oceans. Probably all gave AGE Early Jurassic.
live birth. DISTRIBUTION AND FORMATIONS Southern England;
HABITS Swimming performance mediocre to good. Most lower Lias group.
ambush and/or pursuit fishers and hunters of small to big HABITAT Island archipelago shallows.
game, some filter feeders. NOTES Exact stratigraphic level unknown. May not be
within this family.
PLESIOSAUR MISCELLANEA Lindwurmia thiuda
2.5 m (8 ft) TL, 90 kg (200 lb)
Eoplesiosaurus antiquior FOSSIL REMAINS Partial skull and skeleton.
3 m (10 ft) TL, 150 kg (300 lb) ANATOMICAL CHARACTERISTICS Snout short and
FOSSIL REMAINS Nearly complete skeleton. broad, teeth large.
ANATOMICAL CHARACTERISTICS Neck long. Fore and AGE Early Jurassic; early Hettangian.
aft flippers similar in size.
93

DISTRIBUTION AND FORMATIONS Northwestern Macroplata tenuiceps

Germany; unnamed. 4.5 m (15 ft) TL, 550 kg (1,200 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS Partial skull and majority of skeleton.
NOTES May not be within this family. ANATOMICAL CHARACTERISTICS Neck fairly short.
Shoulder girdle well developed. Fore and aft flippers
Stratesaurus taylori similar in size.
1.2 m (4 ft) TL, 10 kg (20 lb) AGE Early Jurassic; Hettangian.
FOSSIL REMAINS Two skulls, partial skeleton. DISTRIBUTION AND FORMATIONS Southern England;
ANATOMICAL CHARACTERISTICS Head short, fairly Blue Lias.
deep, teeth large. HABITAT Island archipelago shallows.
AGE Early Jurassic; earliest Hettangian. HABITS Powerful swimmer.
DISTRIBUTION AND FORMATIONS Southern England;
lower Blue Lias. Meyerasaurus victor
HABITAT Island archipelago shallows. 3.4 m (11 ft) TL, 275 kg (600 lb)
HABITS Fishers of small to medium-sized game. FOSSIL REMAINS Complete skull and skeleton, severely
NOTES Placement in family uncertain. Shared its habitat flattened top to bottom.
with Eoplesiosaurus, Avalonnectes, Eurycleidus, ANATOMICAL CHARACTERISTICS Front teeth large and
Atychodracon, Thalassiodracon, Protoichthyosaurus. procumbent. Neck fairly short. Fore and aft flippers
similar in size.
Avalonnectes arturi AGE Early Jurassic; early Toarcian.
2 m (4 ft) TL, 50 kg (100 lb) DISTRIBUTION AND FORMATIONS Southern Germany;
FOSSIL REMAINS Partial skull and majority of skeleton, Posidonienschiefer.
partial skeleton. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Standard for group. HABITS Hunter of small and medium-sized game.
AGE Early Jurassic; earliest Hettangian. NOTES Has the best-preserved plesiosaur gastralia series.
DISTRIBUTION AND FORMATIONS Southern England; Shared its habitat with Hauffiosaurus, Seeleyosaurus,
lower Blue Lias. Hydrorion, Hauffiopteryx, unnamed genus trigonodon,
HABITAT Island archipelago shallows. Suevoleviathan, Eurhinosaurus, Stenopterygius,
Mystriosaurus, Platysuchus, Macrospondylus.
Eurycleidus arcuatus
2.5 m (8 ft) TL, 90 kg (200 lb) Maresaurus coccai
FOSSIL REMAINS Partial skull and skeleton. 6 m (20 ft) TL, 1.3 tonnes
ANATOMICAL CHARACTERISTICS Insufficient FOSSIL REMAINS Majority of skull and minority of
information. skeleton.
AGE Early Jurassic; earliest Hettangian. ANATOMICAL CHARACTERISTICS Standard for group.
DISTRIBUTION AND FORMATIONS Southern England; AGE Middle Jurassic; early Bajocian.
lower Blue Lias. DISTRIBUTION AND FORMATIONS Central Argentina;
HABITAT Island archipelago shallows. upper Los Molles.
Atychodracon megacephalus NOTES Shared its habitat with Chacaicosaurus and
4.9 m (16 ft) TL, 650 kg (1,400 lb) Mollesaurus.
FOSSIL REMAINS Skull and partial skeleton, severely
flattened top to bottom. Archaeonectrus rostratus
ANATOMICAL CHARACTERISTICS Insufficient 3.1 m (10 ft) TL, 190 kg (400 lb)
information. FOSSIL REMAINS Two skulls and skeletons, complete and
AGE Early Jurassic; earliest Hettangian. partial.
DISTRIBUTION AND FORMATIONS Southern England; ANATOMICAL CHARACTERISTICS Head somewhat large,
lower Blue Lias. temporal region subrectangular in top view, teeth medium
HABITAT Island archipelago shallows. sized and procumbent. Foreflipper a little smaller than aft
flipper.
94

Archaeonectrus
rostratus
AGE Early Jurassic; middle or late Sinemurian. Neck medium length. Flippers very large, similar in size.
DISTRIBUTION AND FORMATIONS Southeastern AGE Early Jurassic; early Toarcian.
England; lower or middle Charmouth Mudstone. DISTRIBUTION AND FORMATIONS Northeastern
HABITAT Island archipelago shallows. England; lower Whitby Mudstone.
Rhomaleosaurus cramptoni HABITS Powerful swimmer.
6.7 m (22 ft) TL, 2.1 tonnes NOTES May include R. zetlandicus, R. propinquus,
FOSSIL REMAINS Skulls and skeletons. and R. thorntoni. Shared its habitat with Hauffiosaurus,
ANATOMICAL CHARACTERISTICS Head somewhat Eurhinosaurus, Plagiophthalmosuchus, Mystriosaurus,
large, strongly triangular in top view, teeth medium sized. Macrospondylus.
Rhomaleosaurus
cramptoni
95

Rhomaleosaurus
cramptoni
Borealonectes russelli
3 m (10 ft) TL, 190 kg (400 lb)
PLIOSAURIDS
FOSSIL REMAINS Majority of skull and minority SMALL TO GIGANTIC PLIOSAUROIDS FROM
of skeleton. THE EARLY JURASSIC TO EARLY CRETACEOUS,
ANATOMICAL CHARACTERISTICS Standard for group. GLOBAL
AGE Middle Jurassic; late Callovian.
DISTRIBUTION AND FORMATIONS Northwest ANATOMICAL CHARACTERISTICS Significantly variable.
Territories; upper Hiccles Cove. Heads never very small. Necks never very long.
HABITAT Continental shallows, polar. HABITATS Coastal shallows to deep oceans.
HABITS Swimming performance modest to good. Ambush
and/or pursuit fishers and hunters of small to big game.
96

Cryonectes neustriacus
ATTENBOROSAURS 5 m (16 ft) TL, 450 kg (1,000 lb)
SMALL TO GIGANTIC PLIOSAURIDS FROM FOSSIL REMAINS Partial skull.
THE EARLY TO LATE JURASSIC OF EUROPE ANATOMICAL CHARACTERISTICS Teeth large.
AGE Early Jurassic; late Pliensbachian.
ANATOMICAL CHARACTERISTICS Uniform. Heads DISTRIBUTION AND FORMATIONS Northern
not large, necks moderately long. France; Calcaire à Bélemnites.
HABITATS Coastal shallows to deep oceans. HABITAT Island archipelago shallows.
HABITS Swimming performance modest. Ambush HABITS Fishers of medium-sized game.
and pursuit fishers of small to medium-sized game.
NOTES Absence from at least some other seas may reflect Hauffiosaurus longirostris
lack of sufficient sampling. 4.5 m (15 ft) TL, 400 kg (900 lb)
FOSSIL REMAINS Three skulls and skeletons,
Thalassiodracon hawkinsii severely flattened top to bottom.
2 m (6.5 ft) TL, 40 kg (90 lb) ANATOMICAL CHARACTERISTICS Head fairly
FOSSIL REMAINS Half a dozen skulls with at least partial long, narrow, snout long and narrow.
skeletons, latter severely flattened top to bottom. AGE Early Jurassic; early Toarcian.
ANATOMICAL CHARACTERISTICS Head short, deep aft, DISTRIBUTION AND FORMATIONS Southern
moderately broad, teeth medium sized. Fore and aft Germany, northern England; Posidonienschiefer,
flippers similar in size. lower Whitby Mudstone.
AGE Early Jurassic; earliest Hettangian. HABITAT Island archipelago shallows.
DISTRIBUTION AND FORMATIONS Southern England; NOTES May include H. zanoni and H. tomistomimus.
lower Blue Lias. Shared its habitat with Meyerasaurus, Rhomaleosaurus,
HABITAT Island archipelago shallows. Seeleyosaurus, Hydrorion, Hauffiopteryx, unnamed genus
NOTES Shared its habitat with Eoplesiosaurus, trigonodon, Suevoleviathan, Eurhinosaurus, Stenopterygius,
Stratesaurus, Avalonnectes, Eurycleidus, Atychodracon, Plagiophthalmosuchus, Mystriosaurus, Platysuchus,
Protoichthyosaurus. Macrospondylus.
Attenborosaurus conybeari
7.2 m (24 ft) TL, 1.7 tonnes
FOSSIL REMAINS Skull and majority of skeleton.
Thalassiodracon elongated, snout long and fairly robust, teeth large.
hawkinsii Flippers large, fore a little smaller than aft.
AGE Early Jurassic; middle Sinemurian.
lower Charmouth Mudstone.
97

HABITAT Island archipelago shallows. temporal regions greatly expanded. Necks short. Flippers
HABITS Fisher of medium-sized game. medium sized to large.
NOTES Original specimen destroyed by Axis bombing in HABITAT Coastal shallows to deep oceans.
World War II. HABITS Swimming performance mediocre to good. Pursuit
fishers and hunters of medium-sized and especially big
Marmornectes candrewi game.
10 m (30 ft) TL, 5 tonnes NOTES Some partial remains indicate largest known
FOSSIL REMAINS Partial skull and partial skeleton. examples exceeded 12 m (40 ft) in length and approached
ANATOMICAL CHARACTERISTICS Snout narrow. 20 tonnes, probably the most massive known sea reptiles.
AGE Middle Jurassic; early Callovian.
DISTRIBUTION AND FORMATIONS Southern England; Peloneustes philarchus
lower Oxford Clay. 3.8 m (8.4 ft) TL, 600 kg (1,300 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS Skull and skeleton(s).
NOTES Shared its habitat with Peloneustes, ANATOMICAL CHARACTERISTICS Snout long, teeth
Pachycostasaurus, Simolestes, Liopleurodon, Cryptoclidus, medium sized, stout. Flippers very large, similar in size.
Muraenosaurus, Tricleidus, Ophthalmosaurus, AGE Middle Jurassic; early Callovian.
Tyrannoneustes, Suchodus, Gracilineustes. DISTRIBUTION AND FORMATIONS Southern England;
lower Oxford Clay.
Anguanax zignoi HABITAT Island archipelago shallows.
3.5 m (12 ft) TL, 200 kg (450 lb) HABITS Powerful swimmer. Prey included shelled
FOSSIL REMAINS Partial skull and majority of ammonoids.
skeleton. NOTES Shared its habitat with Simolestes, Liopleurodon,
ANATOMICAL CHARACTERISTICS Insufficient Cryptoclidus, Muraenosaurus, Tricleidus,
information. Ophthalmosaurus, Tyrannoneustes, Suchodus,
AGE Late Jurassic; middle Oxfordian. Gracilineustes, Pachycostasaurus.
DISTRIBUTION AND FORMATIONS Northern Italy;
middle Rosso Ammonitico Veronese. Pachycostasaurus dawni
HABITAT Island archipelago shallows. 3 m (10 ft) TL, 300 kg (600 lb)
FOSSIL REMAINS Nearly complete skull and skeleton,
THALASSOPHONEANS severely flattened top to bottom.
ANATOMICAL CHARACTERISTICS Flippers medium
MEDIUM-SIZED TO GIGANTIC PLIOSAURIDS sized.
FROM THE MIDDLE JURASSIC TO EARLY AGE Middle Jurassic; early Callovian.
CRETACEOUS, GLOBAL DISTRIBUTION AND FORMATIONS Southern England;
lower Oxford Clay.
ANATOMICAL CHARACTERISTICS Uniform. Heads very HABITAT Island archipelago shallows.
large, elongated, snouts half or more head length, HABITS Mediocre swimmer.
98

Peloneustes philarchus
Gallardosaurus iturraldei Simolestes? keileni

Adult size uncertain 6 m (20 ft) TL, 2 tonnes
FOSSIL REMAINS Partial skull and minority of skeleton, FOSSIL REMAINS Partial remains.
immature. ANATOMICAL CHARACTERISTICS Insufficient
ANATOMICAL CHARACTERISTICS Insufficient information.
information. AGE Middle Jurassic; late Bajocian.
AGE Middle Jurassic; middle and/or late Oxfordian. DISTRIBUTION AND FORMATIONS Northeastern
DISTRIBUTION AND FORMATIONS Cuba; Jagua. France; La Gravclotte Marls.
HABITAT Continental shallows. HABITAT Island archipelago shallows.
NOTES At that time Cuba was attached to South America. NOTES Whether this species belongs in the much later
Shared its habitat with Vinialesaurus. genus Simolestes is problematic.
99

Peloneustes philarchus
Simolestes
vorax
Acostasaurus
pavachoquensis
Pliosaurus
brachydeirus
Simolestes vorax NOTES Shared its habitat with Peloneustes, Pachycostasaurus,

4.5 m (15 ft) TL, 1 tonne Cryptoclidus, Muraenosaurus, Tricleidus, Ophthalmosaurus,
FOSSIL REMAINS Skulls and skeletons, complete to Tyrannoneustes, Suchodus, Gracilineustes, Liopleurodon.
partial, severely flattened top to bottom.
ANATOMICAL CHARACTERISTICS Head moderately Acostasaurus pavachoquensis
broad, snout about half head length, front teeth large and 4 m (14 ft) TL, 1 tonne
robust. Flippers large. FOSSIL REMAINS Skull and minority of skeleton.
AGE Middle Jurassic; early Callovian. ANATOMICAL CHARACTERISTICS Snout stout, about
DISTRIBUTION AND FORMATIONS Southern England; half of head length, front teeth, especially upper, large and
lower Oxford Clay. robust, lower jaw robust.
HABITAT Island archipelago shallows. AGE Early Cretaceous; early Barremian.
HABITS Powerful swimmer. Prey included shelled DISTRIBUTION AND FORMATIONS Colombia; lower Paja.
ammonoids. HABITAT Continental shallows.
100

Liopleurodon ferox
Pliosaurus brachydeirus
Liopleurodon ferox 8 m (26 ft) TL, 5 tonnes
6.6 m (22 ft) TL, 3.3 tonnes FOSSIL REMAINS Skulls complete and partial, partial
FOSSIL REMAINS Skulls and skeletons. skeletons.
ANATOMICAL CHARACTERISTICS Snout long, front ANATOMICAL CHARACTERISTICS Snout long, teeth
teeth very large. Tail quite short. Flippers large, aft set a medium sized.
little larger. AGE Late Jurassic; early Kimmeridgian.
AGE Middle Jurassic; early Callovian. DISTRIBUTION AND FORMATIONS Southern England;
DISTRIBUTION AND FORMATIONS Southern England; lower Kimmeridge Clay.
lower Oxford Clay. HABITAT Island archipelago shallows.
HABITAT Island archipelago shallows. NOTES May include P. kevani. Shared its habitat with
HABITS Powerful swimmer. Colymbosaurus megadeirus, Bathysuchus, Torvoneustes,
NOTES Claims of this being 25 m (80 ft) long are greatly Plesiosuchus. May be the direct ancestor of Pliosaurus
exaggerated. westburyensis.
101

Pliosaurus westburyensis Pliosaurus? rossicus

8 m (26 ft) TL, 5 tonnes 10 m (33 ft) TL, 11 tonnes
FOSSIL REMAINS Crushed skulls and partial skeletal FOSSIL REMAINS A few partial skulls and skeletons.
remains. ANATOMICAL CHARACTERISTICS Insufficient
ANATOMICAL CHARACTERISTICS Insufficient information.
information. AGE Late Jurassic; early and/or middle Tithonian.
AGE Late Jurassic; middle Kimmeridgian. DISTRIBUTION AND FORMATIONS Southwestern
DISTRIBUTION AND FORMATIONS Southern England; Russia, Kazakhstan; unnamed.
middle Kimmeridge Clay. HABITAT Continental shallows.
HABITAT Island archipelago shallows. NOTES May not be within Pliosaurus.
NOTES May include P. carpenteri. Shared its habitat with
Nannopterygius enthekiodon, Grendelius mordax. May be Stenorhynchosaurus munozi
the direct ancestor of P. macromerus. 5.3 m (17 ft) TL, 1.7 tonnes
FOSSIL REMAINS Skull and minority of skeleton of adult,
Pliosaurus macromerus majority of skull and skeleton of immature.
8 m (26 ft) TL, 5 tonnes ANATOMICAL CHARACTERISTICS Snout very long,
FOSSIL REMAINS Partial skull and other remains. teeth very numerous and small. Flippers long, aft set a
ANATOMICAL CHARACTERISTICS Snout long, robustly little larger.
built. AGE Early Cretaceous; middle Barremian.
AGE Late Jurassic; late Kimmeridgian. DISTRIBUTION AND FORMATIONS Colombia; lower Paja.
DISTRIBUTION AND FORMATIONS Southern England; HABITAT Continental shallows.
upper Kimmeridge Clay. HABITS Powerful swimmer. Fisher of medium-sized game.
Pliosaurus? funkei Stenorhynchosaurus

10–12+ m (33–40+ ft) TL munozi
FOSSIL REMAINS Minority of two skulls and skeletons.
information.
AGE Late Jurassic; middle Tithonian. Sachicasaurus vitae
DISTRIBUTION AND FORMATIONS Svalbard; upper 10.8 (35 ft) TL, 13.5 tonnes
Agardhfjellet. FOSSIL REMAINS Majority of skull and skeleton.
HABITAT Open continental shelf, polar. ANATOMICAL CHARACTERISTICS Head very large,
NOTES Placement in earlier Pliosaurus problematic. massive, especially broad aft section, most teeth medium
Claims of this being 15 m (50 ft) long are exaggerated. sized except one pair of large teeth on upper and lower
Shared its habitat with Colymbosaurus? svalbardensis, jaws. Flippers fairly large.
Spitrasaurus, Keilhauia, Palvennia, Janusaurus, AGE Early Cretaceous; late Barremian.
Cryopterygius. DISTRIBUTION AND FORMATIONS Colombia; middle
Paja.
Pliosaurus? patagonicus HABITAT Continental shallows.
Adult size uncertain HABITS Good swimmer. Prey included shelled
FOSSIL REMAINS Partial skull, badly damaged, possibly ammonoids.
juvenile. NOTES Most massive well-preserved plesiosaur.
information. Monquirasaurus boyacensis
AGE Late Jurassic; middle Tithonian. 9.4 m (31 ft) TL, 9.1 tonnes
DISTRIBUTION AND FORMATIONS Western Argentina; FOSSIL REMAINS Majority of skull and skeleton.
lower Vaca Muerta. ANATOMICAL CHARACTERISTICS Head very large,
HABITAT Continental shallows. especially relative to body, particularly massive, especially
NOTES Placement in earlier Pliosaurus problematic. unusually broad aft section, front teeth large, rest medium
Shared its habitat with Caypullisaurus, Cricosaurus sized to small. Flippers not large, aft a little larger.
araucanensis. AGE Early Cretaceous; late Aptian.
102

Stenorhynchosaurus munozi
Sachicasaurus
vitae
DISTRIBUTION AND FORMATIONS Colombia; upper Paja.
HABITS Mediocre swimmer. Prey included shelled
ammonoids.
103

Monquirasaurus boyacensis
NOTES Originally placed in Kronosaurus. Shared its AGE Early Cretaceous; late Aptian.
habitat with Callawayasaurus, Kyhytysuka. The biggest- DISTRIBUTION AND FORMATIONS Northeastern
headed and most strongly built known plesiosaur, with Australia; Wallumbilla.
the most powerful bite force. HABITAT Interior seaway.
HABITS Good swimmer. Prey included shelled ammonoids.
Eiectus longmani NOTES Originally placed in later and more fragmentary
10 m (33 ft) TL, 11 tonnes Kronosaurus queenslandicus, the remains constitute an
FOSSIL REMAINS A few partial skulls and skeletons. iconic large-headed plesiosaur, albeit now with a new
ANATOMICAL CHARACTERISTICS Head long, somewhat name that may be challenged in the future. Remains are
shallow, moderately broad aft, snout long and rather not well preserved or described, and claims of this being
narrow, some front teeth large, others medium sized to 13 m (42 ft) long are exaggerated.
small. Flippers fairly large.
104

?Kronosaurus queenslandicus
Size uncertain
PLESIOSAUROIDS
FOSSIL REMAINS A few partial skull and skeletal remains. SMALL TO GIGANTIC PLESIOSAURS FROM
ANATOMICAL CHARACTERISTICS Insufficient THE EARLY JURASSIC TO THE END OF
information. THE MESOZOIC, GLOBAL
AGE Early Cretaceous; middle Albian.
DISTRIBUTION AND FORMATIONS Northeastern ANATOMICAL CHARACTERISTICS Fairly variable. Heads
Australia; Toolebuc. very small to very large. Necks very short to extremely long.
HABITAT Interior seaway. HABITATS Freshwaters to deep oceans.
HABITS Prey included shelled ammonoids. HABITS Swimming performance mediocre to good. Most
NOTES An iconic large-headed plesiosaur name, the ambush and/or pursuit fishers and hunters of small to big
best remains are currently placed in the new Eiectus game, some filter feeders.
longmani; the original K. queenslandicus specimen is
too fragmentary to base a species on, but an attempt to PLESIOSAURIDS
revive the classic name may occur by reapplying it to
Eiectus longmani fossils. Shared its habitat with SMALL TO GIGANTIC PLESIOSAUROIDS FROM
Eromangasaurus, Longirostra, Cratochelone, Notochelone, THE EARLY TO LATE JURASSIC OF EURASIA
Bouliachelys.
ANATOMICAL CHARACTERISTICS Uniform. Heads not
large, necks moderately long.
HABITATS Freshwaters, coastal shallows, continental
shelves.
Kronosuarus
HABITS Swimming performance modest to good.
queenslandicus
Ambush and pursuit fishers of small to medium-sized
game.
NOTES Absence from at least some other seas may reflect
lack of sufficient sampling.
Bishanopliosaurus youngi
4 m (13 ft) TL, 300 kg (650 lb)
Brachauchenius lucasi FOSSIL REMAINS Minority of skeleton.
7 m (23 ft) TL, 3.5 tonnes ANATOMICAL CHARACTERISTICS Insufficient
FOSSIL REMAINS Four skulls and minority of skeletons. information.
ANATOMICAL CHARACTERISTICS Snout very long, AGE Early Jurassic; Toarcian.
small crest atop back of skull, teeth medium sized, robust. DISTRIBUTION AND FORMATIONS Central China; upper
AGE Late Cretaceous; early to middle Turonian. Ziliujing.
DISTRIBUTION AND FORMATIONS Kansas; lower Carlile HABITAT Lakes, possibly rivers.
Shale. HABITS May have occasionally snapped up small
HABITAT Interior seaway. swimming land creatures.
HABITS Prey included shelled ammonoids. NOTES Whether this taxon was a permanent freshwater
NOTES Megacephalosaurus eulerti appears to be the adult plesiosaur or a transient is not certain.
of this species. Shared its habitat with Trinacromerum
bentonianum. Bishanopliosaurus? zigongensis
4 m (13 ft) TL, 300 kg (650 lb)
FOSSIL REMAINS Minority of skeleton.
Brachauchenius information.
lucasi AGE Middle Jurassic.
DISTRIBUTION AND FORMATIONS Central China; lower
Shaximiao.
HABITAT Lakes, possibly rivers.
HABITS May have occasionally snapped up small
swimming land creatures.
105

NOTES May not be within Bishanopliosaurus. Whether AGE Early Jurassic; early Sinemurian.
this taxon was a permanent freshwater plesiosaur or a DISTRIBUTION AND FORMATIONS Southern England;
transient is not certain. upper Blue Lias.
Plesiosaurus dolichodeirus
3.4 m (11 ft) TL, 185 kg (400 lb)
FOSSIL REMAINS A few skulls with skeletons and other
remains.
ANATOMICAL CHARACTERISTICS Head small, snout
short, teeth long spikes. Neck long. Flippers long, fore
a little larger than aft.
106

HABITS Swimming performance good. Lusonectes sauvagei

NOTES Shared its habitat with Excalibosaurus, 2 m (6.5 ft) TL, 35 kg (80 lb)
Eretmosaurus, Leptonectes tenuirostris, Temnodontosaurus, FOSSIL REMAINS Partial skull.
Ichthyosaurus communis, unnamed genus eurycephalus. ANATOMICAL CHARACTERISTICS Insufficient information.
AGE Early Jurassic; Toarcian.
MICROCLEIDIDS DISTRIBUTION AND FORMATIONS Portugal; Sao Giao.
SMALL TO MEDIUM-SIZED PLESIOSAUROIDS
FROM THE EARLY TO LATE JURASSIC OF Eretmosaurus rugosus
EURASIA 3.7 m (12 ft) TL, 225 kg (500 lb)
FOSSIL REMAINS Skeleton, severely flattened top to
ANATOMICAL CHARACTERISTICS Head small, snout bottom.
short. Neck fairly long. ANATOMICAL CHARACTERISTICS Neck long. Flippers
HABITAT Coastal shallows. large, similar in size.
HABITS Swimming performance good. Ambush and AGE Early Jurassic; middle Sinemurian.
pursuit fishers of small to medium-sized game. DISTRIBUTION AND FORMATIONS Southern England;
NOTES Absence from at least some other seas may reflect upper Blue Lias.
lack of sufficient sampling. HABITAT Island archipelago shallows.
107

NOTES Shared its habitat with Excalibosaurus, Seeleyosaurus guilelmiimperatoris

Plesiosaurus, Leptonectes tenuirostris, Temnodontosaurus, 3.8 m (12.5 ft) TL, 250 kg (550 lb)
Ichthyosaurus communis, unnamed genus eurycephalus. FOSSIL REMAINS Complete skull and skeleton.
ANATOMICAL CHARACTERISTICS Head very small, fairly
Westphaliasaurus simonsensii broad and subtriangular in top view, teeth numerous and
4.5 m (15 ft) TL, 400 kg (900 lb) small. Neck long. Tail moderately long. Flippers very large,
FOSSIL REMAINS Majority of skeleton. similar in size, strongly distally swept back.
ANATOMICAL CHARACTERISTICS Insufficient AGE Early Jurassic; early Toarcian.
information. DISTRIBUTION AND FORMATIONS Southern Germany;
AGE Early Jurassic; Pliensbachian. Posidonienschiefer.
DISTRIBUTION AND FORMATIONS Northwestern HABITAT Island archipelago shallows. Shared its habitat
Germany; unnamed. with Meyerasaurus, Hauffiosaurus, Hydrorion,
HABITAT Island archipelago shallows. Hauffiopteryx, unnamed genus trigonodon,
Suevoleviathan, Eurhinosaurus, Stenopterygius,
Plesiopharos moelensis Mystriosaurus, Platysuchus, Macrospondylus.
Adult size uncertain
FOSSIL REMAINS Partial skeleton, probably juvenile. Microcleidus melusinae
ANATOMICAL CHARACTERISTICS Insufficient 3 m (10 ft) TL, 120 kg (250 lb)
information. FOSSIL REMAINS Skull and partial skeleton, severely
AGE Early Jurassic; late Sinemurian. damaged.
DISTRIBUTION AND FORMATIONS Portugal; lower ANATOMICAL CHARACTERISTICS Head moderately
Coimbra. broad, subrectangular at back, front teeth large and
HABITAT Island archipelago shallows. procumbent.
Seelyosaurus guilelmiimperatoris
108

AGE Early Jurassic; early Toarcian. Microcleidus? tournemirensis

DISTRIBUTION AND FORMATIONS Luxembourg; 4 m (13 ft) TL, 300 kg (650 lb)
unnamed. FOSSIL REMAINS Skull and majority of skeleton.
HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Head moderately
NOTES May be the direct ancestor of M. homalospondylus. broad, widest at midpoint, teeth large and procumbent.
Neck long.
Microcleidus homalospondylus AGE Early Jurassic; late Toarcian.
5.1 m (17 ft) TL, 650 kg (1,400 lb) DISTRIBUTION AND FORMATIONS Southern France;
FOSSIL REMAINS Three nearly complete skulls and unnamed.
skeletons. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Head very small,
narrow, subrectangular at back, front teeth very large and Hydrorion brachypterygius
procumbent, intermeshing spikes. Neural spines over 3.2 m (10.5 ft) TL, 150 kg (330 lb)
trunk very tall. Neck very long. Flippers very large, similar FOSSIL REMAINS Complete skull and skeleton.
in size. ANATOMICAL CHARACTERISTICS Head small, fairly
AGE Early Jurassic; late early Toarcian. broad, widest at midpoint, teeth fairly long, slender,
DISTRIBUTION AND FORMATIONS Northeastern procumbent intermeshing spikes. Trunk vertebral spines
England; upper Whitby Mudstone. tall. Flippers large, fore a little smaller than aft.
HABITAT Island archipelago shallows. AGE Early Jurassic; early Toarcian.
HABITS Probably includes M. macropterus. Combination DISTRIBUTION AND FORMATIONS Southern Germany;
of tall spines of trunk vertebrae and large flippers indicates Posidonienschiefer.
very powerful propulsion. HABITAT Island archipelago shallows.
Microcleidus homalospondylus
109

Hydrorion brachypterygius
HABITS Combination of tall spines of trunk vertebrae and CRYPTOCLIDIDS

large flippers indicates very powerful propulsion.
NOTES Shared its habitat with Meyerasaurus, MEDIUM-SIZED TO LARGE CRYPTOCLIDIANS
Hauffiosaurus, Seeleyosaurus, Hauffiopteryx, unnamed FROM THE MIDDLE JURASSIC TO EARLY
genus trigonodon, Suevoleviathan, Eurhinosaurus, CRETACEOUS OF THE NORTHERN
Stenopterygius, Mystriosaurus, Platysuchus, Macrospondylus. HEMISPHERE.
CRYPTOCLIDIANS ANATOMICAL CHARACTERISTICS Necks moderately

long.
MEDIUM-SIZED TO GIGANTIC PLESIOSAUROIDS HABITAT Freshwaters, coastal shallows, continental shelves.
FROM THE MIDDLE JURASSIC TO THE END HABITS Swimming performance modest. Ambush and
OF THE MESOZOIC, GLOBAL pursuit fishers of small to medium-sized game.
NOTES Absence from Southern Hemisphere may reflect
ANATOMICAL CHARACTERISTICS Flippers more highly lack of sufficient sampling.
developed, with lower arm and leg elements shorter and
more blocklike. Abyssosaurus nataliae
HABITAT Freshwaters to deep oceans. 7 m (23 ft) TL, 1.8 tonnes
HABITS Swimming performance mediocre to good. Most FOSSIL REMAINS Partial skeleton.
ambush and/or pursuit fishers and hunters of small to big ANATOMICAL CHARACTERISTICS Insufficient
game, some filter feeders. information.
AGE Early Cretaceous; late Hauterivian.
DISTRIBUTION AND FORMATIONS Western Russia;
unnamed.
110

Colymbosaurus megadeirus HABITAT Open continental shelf, polar.

5 m (16 ft) TL, 700 kg (1,500 lb) NOTES Djupedalia engeri may be the juvenile of this
FOSSIL REMAINS Possible partial skull, partial skeletons. species. Placement in much earlier Colymbosaurus is
ANATOMICAL CHARACTERISTICS Insufficient uncertain. Shared its habitat with Pliosaurus? funkei,
information. Spitrasaurus, Keilhauia, Palvennia, Janusaurus,
AGE Late Jurassic; middle and/or late Kimmeridgian. Cryopterygius.
lower Kimmeridge Clay, Haddenham. Cryptoclidus eurymerus
HABITAT Island archipelago shallows. 5.5 m (18 ft) TL, 1 tonne
NOTES Kimmerosaurus langhami skull may belong to this FOSSIL REMAINS Skull and skeleton.
species. Shared its habitat with Pliosaurus brachydeirus, ANATOMICAL CHARACTERISTICS Front teeth large,
Bathysuchus, Torvoneustes, Plesiosuchus. procumbent, intermeshing slender spikes. Flippers similar
in size.
Colymbosaurus? svalbardensis AGE Middle Jurassic; early or middle Callovian.
5 m (16 ft) TL, 700 kg (1,500 lb) DISTRIBUTION AND FORMATIONS Central England;
FOSSIL REMAINS Four partial skeletons, one juvenile. lower Oxford Clay.
ANATOMICAL CHARACTERISTICS Insufficient HABITAT Island archipelago shallows.
information. NOTES Shared its habitat with Peloneustes,
AGE Late Jurassic; late Tithonian. Pachycostasaurus, Simolestes, Liopleurodon, Tricleidus,
DISTRIBUTION AND FORMATIONS Svalbard; upper Ophthalmosaurus, Tyrannoneustes, Suchodus,
Agardhfjellet. Gracilineustes, Muraenosaurus.
HABITAT Open continental shelf, polar.
Cryptoclidus eurymerus
111

Cryptoclidus eurymerus
Muraenosaurus leedsii Tatenectes laramiensis

5.9 m (19 ft) TL, 1 tonne 3 m (10 ft) TL, 450 kg (1,000 lb)
FOSSIL REMAINS Complete and partial skulls and FOSSIL REMAINS Two partial skeletons.
skeletons. ANATOMICAL CHARACTERISTICS Trunk may be
ANATOMICAL CHARACTERISTICS Head fairly broad, shallower than usual.
teeth medium sized. Trunk fairly long, hefty. Flippers AGE Late Jurassic; early Oxfordian.
moderate and similar in size. DISTRIBUTION AND FORMATIONS Wyoming; upper
AGE Middle Jurassic; middle Callovian. Sundance.
DISTRIBUTION AND FORMATIONS Southern England; HABITAT Interior seaway.
lower Oxford Clay.
HABITAT Island archipelago shallows. Spitrasaurus wensaasi
HABITS Swimming performance mediocre. Adult size uncertain
FOSSIL REMAINS Partial juvenile skeletons.
Pantosaurus striatus ANATOMICAL CHARACTERISTICS Insufficient
3.5 m (11 ft) TL, 250 kg (550 lb) information.
FOSSIL REMAINS Partial skeletons. AGE Late Jurassic; late Tithonian.
ANATOMICAL CHARACTERISTICS Foreflipper distally DISTRIBUTION AND FORMATIONS Svalbard; upper
swept back because radius is much larger than ulna. Agardhfjellet.
AGE Late Jurassic; early Oxfordian. HABITAT Open continental shelf, polar.
DISTRIBUTION AND FORMATIONS Wyoming; upper NOTES May include S. larseni. Shared its habitat with
Sundance. Pliosaurus? funkei, Colymbosaurus? svalbardensis,
HABITAT Interior seaway. Keilhauia, Palvennia, Janusaurus, Cryopterygius.
NOTES Shared its habitat with Tatenectes, Baptanodon.
112

Ophthalmothule cryostea
4.5 m (15 ft) TL, 500 kg (1,000 lb)
BASOELASMOSAURIDS
FOSSIL REMAINS Skull severely flattened top to bottom, MEDIUM-SIZED TO GIGANTIC ELASMOSAURIDS
majority of skeleton. FROM THE EARLY CRETACEOUS TO THE END
ANATOMICAL CHARACTERISTICS Neck fairly long, OF THE MESOZOIC, GLOBAL
foreflipper somewhat larger than aft.
AGE Late Jurassic or Early Cretaceous; latest Tithonian or HABITS Swimming performance modest.
early Berriasian.
DISTRIBUTION AND FORMATIONS Svalbard; uppermost Lagenanectes richterae
Agardhfjellet. 8 m (26 ft) TL, 1.3 tonnes
HABITAT Open continental shelf, polar. FOSSIL REMAINS Partial skull and minority of skeleton.
NOTES May extend group into the Early Cretaceous. ANATOMICAL CHARACTERISTICS Front teeth large.
AGE Early Cretaceous; late Hauterivian.
Tricleidus seeleyi DISTRIBUTION AND FORMATIONS Northern Germany;
5 m (16 ft) TL, 700 kg (1,500 lb) Stadthagen.
FOSSIL REMAINS Partial skull and skeleton. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Front teeth large. NOTES Shared its habitat with Enaliosuchus.
AGE Middle Jurassic; middle Callovian.
DISTRIBUTION AND FORMATIONS Southern England; Eromangasaurus australis
lower Oxford Clay. 5 m (17 ft) TL, 350 kg (800 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS Nearly complete skull, severely
NOTES Shared its habitat with Peloneustes, damaged.
Pachycostasaurus, Simolestes, Liopleurodon, Cryptoclidus, ANATOMICAL CHARACTERISTICS Teeth small.
Muraenosaurus, Ophthalmosaurus, Tyrannoneustes, AGE Early Cretaceous; middle Albian.
Suchodus, Gracilineustes. DISTRIBUTION AND FORMATIONS Northeastern
Australia; Toolebuc.
Vinialesaurus caroli HABITAT Interior seaway.
3.5 m (11 ft) TL, 250 kg (550 lb) HABITS Fisher of small game.
FOSSIL REMAINS Majority of skull and minority of NOTES Shared its habitat with ?Kronosaurus, Longirostra,
skeleton. Cratochelone, Notochelone, Bouliachelys.
ANATOMICAL CHARACTERISTICS Head broad, teeth
large spikes. Callawayasaurus colombiensis
AGE Late Jurassic; middle and/or late Oxfordian. 8 m (26 ft) TL, 1.3 tonnes
DISTRIBUTION AND FORMATIONS Cuba; Jagua. FOSSIL REMAINS Two nearly complete skulls and
HABITAT Continental shallows. skeletons.
NOTES At that time Cuba was attached to South America. ANATOMICAL CHARACTERISTICS Teeth large, robust.
Shared its habitat with Gallardosaurus. AGE Early Cretaceous; Aptian.
DISTRIBUTION AND FORMATIONS Colombia; upper Paja.
ELASMOSAURIDS HABITAT Continental coastal.
NOTES Shared its habitat with Monquirasaurus,
MEDIUM-SIZED TO GIGANTIC CRYPTOCLIDIANS Kyhytysuka.
FROM THE EARLY CRETACEOUS TO THE END
OF THE MESOZOIC, GLOBAL
Callawayasaurus
ANATOMICAL CHARACTERISTICS Heads small, ears colombiensis
absent. Necks very to extremely long.
HABITAT Mainly coastal shallows, a few in freshwaters.
HABITS Swimming performance mediocre to modest.
Ambush hunters of small and medium-sized game.
NOTES The placement of the very poorly preserved Late
Triassic Alexeyisaurus in the otherwise entirely Cretaceous
group is highly problematic.
113

Wapuskanectes betsynichollsae HABITAT Continental coastal.

10 m (33 ft) TL, 2 tonnes
FOSSIL REMAINS Partial skeleton. Fluvionectes sloanae
ANATOMICAL CHARACTERISTICS Insufficient 5 m (16 ft) TL, 500 kg (1,000 lb)
information. FOSSIL REMAINS Partial skeleton.
AGE Early Cretaceous; earliest Albian. ANATOMICAL CHARACTERISTICS Insufficient
DISTRIBUTION AND FORMATIONS Alberta; lower information.
Clearwater. AGE Late Cretaceous; late Campanian.
HABITAT Interior seaway, polar. DISTRIBUTION AND FORMATIONS Alberta; lower
NOTES Shared its habitat with Athabascasaurus, Dinosaur Park.
Nichollssaura. HABITAT Coastal rivers, possibly brackish waters.
HABITS May have occasionally snapped up small
Woolungasaurus glendowerensis swimming land creatures.
9 m (30 ft) TL, 1.8 tonnes NOTES With other remains from the region reaching 7 m
FOSSIL REMAINS Majority of skeleton, three partial (23 ft), these are the largest known freshwater plesiosaurs.
skeletons.
ANATOMICAL CHARACTERISTICS Insufficient Nakonanectes bradti
information. 5.5 m (18 ft) TL, 600 kg (1,300 lb)
AGE Early Cretaceous; late Aptian. FOSSIL REMAINS Complete skull and partial skeleton.
DISTRIBUTION AND FORMATIONS Northeastern ANATOMICAL CHARACTERISTICS Head deep at back,
Australia; Wallumbilla. most teeth large and stout. Neck moderately long.
HABITAT Interior seaway. AGE Late Cretaceous; early Maastrichtian.
DISTRIBUTION AND FORMATIONS Montana; upper
Libonectes morgani Bearpaw Shale.
9 m (30 ft) TL, 1.8 tonnes HABITAT Interior seaway was becoming very narrow.
FOSSIL REMAINS Partial skull, minority of skeleton. NOTES Shared its habitat with Terminonatator,
ANATOMICAL CHARACTERISTICS Teeth large, stout, Tylosaurus saskatchewanensis, Plioplatecarpus? primaevus.
procumbent.
AGE Late Cretaceous; late Turonian.
DISTRIBUTION AND FORMATIONS Texas; Britton.
HABITAT Continental coastal. Nakonanectes
NOTES Presence of taxon in Morocco is problematic. bradti
Futabasaurus suzukii
7 m (23 ft) TL, 700 kg (1,500 lb)
ANATOMICAL CHARACTERISTICS Teeth medium sized. Cardiocorax mukulu
Foreflipper larger than aft. 6 m (20 ft) TL
AGE Late Cretaceous; early Santonian. FOSSIL REMAINS Skull crushed and partial skeleton,
DISTRIBUTION AND FORMATIONS Japan; upper two partial skeletons.
Tamayama. ANATOMICAL CHARACTERISTICS Some teeth large.
HABITAT Continental coastal. AGE Late Cretaceous; early Maastrichtian.
NOTES At that time Japan was attached to Asia. DISTRIBUTION AND FORMATIONS Angola; Mocuio.
HABITAT Continental coastal.
Kawanectes lafquenianum
3.8 m (12 ft) TL, 140 kg (300 lb) Vegasaurus molyi
FOSSIL REMAINS A few partial skulls and skeletons. 7 m (23 ft) TL, 700 kg (1,500 lb)
ANATOMICAL CHARACTERISTICS Insufficient FOSSIL REMAINS Majority of skeleton.
information. ANATOMICAL CHARACTERISTICS Neck very long.
AGE Late Cretaceous; late Campanian and/or early AGE Late Cretaceous; early Maastrichtian.
Maastrichtian. DISTRIBUTION AND FORMATIONS Antarctic
DISTRIBUTION AND FORMATIONS Southern Argentina; Peninsula; Snow Hill Island.
middle Allen, lower La Colonia? HABITAT Continental coastal, polar.
114

Morenosaurus stocki HABITAT Continental coastal.

6 m (20 ft) TL, 450 kg (1,000 lb) NOTES Shared its habitat with Hydrotherosaurus
FOSSIL REMAINS Two partial skeletons, one juvenile. alexandrae.
information. Hydrotherosaurus alexandrae
AGE Late Cretaceous; late Maastrichtian. 8 m (26 ft) TL, 1.1 tonnes
DISTRIBUTION AND FORMATIONS Central California; FOSSIL REMAINS Complete skull and skeleton.
middle Moreno. ANATOMICAL CHARACTERISTICS Head moderately
HABITAT Continental coastal. broad, front upper teeth large, robust. Neck extremely
NOTES Fresnosaurus drescheri may be the juvenile of this long. Flippers very large, fore a little larger than aft.
species. AGE Late Cretaceous; late Maastrichtian.
DISTRIBUTION AND FORMATIONS Central California;
Aphrosaurus furlongi upper Moreno.
10 m (33 ft) TL, 1 tonne HABITAT Continental coastal.
FOSSIL REMAINS Minority of skeleton.
ANATOMICAL CHARACTERISTICS Flippers rather
slender, similar in size.
AGE Late Cretaceous; late Maastrichtian.
DISTRIBUTION AND FORMATIONS Central California;
upper Moreno.
Hydrotherosaurus alexandrae
115

Hydrotherosaurus alexandrae
Cimoliasaurus magnus Tuarangisaurus keyesi

7.5 m (25 ft) TL, 1 tonne 8 m (26 ft) TL, 1 tonne
FOSSIL REMAINS Partial skeleton. FOSSIL REMAINS Nearly complete, badly damaged skull
ANATOMICAL CHARACTERISTICS Insufficient and minority of skeleton, possible juvenile remains.
information. ANATOMICAL CHARACTERISTICS Insufficient
AGE Late Cretaceous; early Maastrichtian. information.
DISTRIBUTION AND FORMATIONS New Jersey; AGE Late Cretaceous; late Campanian and/or early
Navesink. Maastrichtian.
HABITAT Continental shelf. DISTRIBUTION AND FORMATIONS Southern New
NOTES Assignment of other remains from various Zealand; upper Tahora.
locations to this taxon is problematic. Shared its habitat HABITAT Island coastal.
with Mosasaurus? conodon.
Thalassomedon hanningtoni
Zarafasaura oceanis 10.1 m (33 ft) TL, 2.5 tonnes
3.5 m (11 ft) TL, 100 kg (220 lb) FOSSIL REMAINS Two skulls and several skeletons from
FOSSIL REMAINS Partial skull and skeleton. nearly complete to partial.
ANATOMICAL CHARACTERISTICS Teeth large and stout. ANATOMICAL CHARACTERISTICS Head moderately
Neck moderately long. broad, deep aft, teeth medium sized. Neck extremely
AGE Late Cretaceous; latest Maastrichtian. long. Tail short. Flippers large, similar in size.
DISTRIBUTION AND FORMATIONS Morocco; unnamed. AGE Late Cretaceous; early Cenomanian.
HABITAT Continental coastal. DISTRIBUTION AND FORMATIONS Colorado, Montana;
NOTES Shared its habitat with Ocepechelon. Graneros Shale, Belle Fourche.
HABITAT Interior seaway.
NOTES Shared its habitat with Plesiopleurodon.
116

Thalassomedon hanningtoni
ELASMOSAURINES ANATOMICAL CHARACTERISTICS Small bump atop

midline of back of snout, teeth modest in number, fairly
GIGANTIC ELASMOSAURIDS FROM THE LATE stout. Neck extremely long. Flippers long, fairly narrow,
CRETACEOUS OF NORTH AMERICA similar in size.
AGE Late Cretaceous; earliest Campanian.
ANATOMICAL CHARACTERISTICS Long, procumbent DISTRIBUTION AND FORMATIONS Kansas, South
spike teeth. Necks very to extremely long. Dakota; uppermost Niobrara.
HABITAT Interior seaway. HABITAT Interior seaway was continuing to become less
HABITS Swimming performance mediocre. Ambush broad and deep.
fishers of small and medium-sized game. NOTES Because seaway was so broad and deep, shallow
NOTES Absence from at least some other seas may reflect coastal waters that preserved elasmosaurine fossils in
lack of sufficient sampling. marine sediments were are not abundant from this time
and region. Shared its habitat with Polycotylus latipinnis,
Styxosaurus snowii Dolichorhynchops osborni, Platecarpus tympanicus,
10.5 m (34.5 ft) TL, 2.3 tonnes Eonatator sternbergii, Clidastes propython, Ctenochelys
FOSSIL REMAINS Two skulls and several skeletons from stenoporus, Protostega gigas.
nearly complete to partial.
Styxosaurus snowii
117

Styxosaurus snowii
Styxosaurus browni absolute and relative, with 76 vertebrae and 6.5 m

10.5 m (34.5 ft) TL, 2.3 tonnes (21.3 ft) long.
FOSSIL REMAINS Complete skull and partial skeleton.
ANATOMICAL CHARACTERISTICS Head not broad, Terminonatator ponteixensis
teeth modest in number, medium sized. Neck extremely 9 m (30 ft) TL, 1.6 tonnes
long. Flippers long, fairly narrow, similar in size. FOSSIL REMAINS Partial skull and skeleton.
AGE Late Cretaceous; early Campanian. ANATOMICAL CHARACTERISTICS Teeth modest in
DISTRIBUTION AND FORMATIONS Wyoming; lower number, large, quite robust, procumbent spikes.
Pierre Shale. AGE Late Cretaceous; latest Campanian.
HABITAT Interior seaway was continuing to become less DISTRIBUTION AND FORMATIONS Saskatchewan; upper
broad and deep. Bearpaw Shale.
NOTES May be a little larger than S. snowii. Shared its HABITAT Interior seaway was becoming very narrow.
habitat with Dolichorhynchops bonneri, Tylosaurus proriger, NOTES Shared its habitat with Nakonanectes, Tylosaurus
Latoplatecarpus, Globidens? dakotensis, Toxochelys saskatchewanensis, Plioplatecarpus? primaevus.
latiremus, Elasmosaurus.
Elasmosaurus platyurus
ARISTONECTIDS
10.3 (34 ft) TL, 2 tonnes LARGE TO GIGANTIC CRYPTOCLIDIANS FROM
FOSSIL REMAINS Minority of skull and partial skeleton. THE LATE CRETACEOUS OF THE SOUTHERN
ANATOMICAL CHARACTERISTICS Neck extremely long. HEMISPHERE
AGE Late Cretaceous; early Campanian.
DISTRIBUTION AND FORMATIONS Kansas; lower Pierre ANATOMICAL CHARACTERISTICS Skull short, broad,
Shale. tall aft, snout rounded, teeth numerous, small, delicate
HABITAT Interior seaway was continuing to become less needle spikes, forming interlocking but nonoccluding
broad and deep. dental combs. Necks moderately long.
NOTES Although an iconic plesiosaur, is known from only HABITS Swimming performance mediocre. Probably
one incomplete specimen. suction filter feeders comparable to baleen whales.
NOTES Placement of these highly specialized filter
Albertonectes vanderveldei feeders within elasmosaurids not logical. Absence
11 m (24 ft) TL, 2 tonnes from the Northern Hemisphere may reflect lack
FOSSIL REMAINS Complete skeleton without skull. wwof sufficient sampling.
ANATOMICAL CHARACTERISTICS Neck extremely long,
flippers similar in size. Kaiwhekea katiki
AGE Late Cretaceous; late Campanian. 7 m (23 ft) TL, 1.5 tonnes
DISTRIBUTION AND FORMATIONS Alberta; lower FOSSIL REMAINS Majority of skull and skeleton.
Bearpaw Shale. ANATOMICAL CHARACTERISTICS Skull very deep
HABITAT Interior seaway was becoming much narrower. aft, fairly narrow, small teeth numerous, somewhat
HABITS Extremely long neck combined with modest procumbent.
propulsive power indicates a rather slow ambush fisher. AGE Late Cretaceous; middle Maastrichtian.
NOTES Shared its habitat with Dolichorhynchops DISTRIBUTION AND FORMATIONS Southern New
herschelensis, Prognathodon? overtoni. Longest known Zealand; lower or middle Katiki.
plesiosaur, and longest known plesiosaur neck both HABITAT Island coastal.
118

LEPTOCLEIDIANS
MEDIUM-SIZED TO LARGE CRYPTOCLIDIANS
FROM THE EARLY CRETACEOUS TO THE
END OF THE MESOZOIC, GLOBAL
Kaiwhekea katiki
ANATOMICAL CHARACTERISTICS Heads at least
moderately long. Necks not long.
HABITAT Freshwaters to continental shelves.
HABITS Swimming performance good. Pursuit fishers and
hunters of medium-sized to big game.
Aristonectes parvidens
LEPTOCLEIDIDS
Adult size not certain MEDIUM-SIZED TO LARGE LEPTOCLEIDIANS
FOSSIL REMAINS Partial skull and skeleton, possibly FROM THE EARLY CRETACEOUS TO THE
immature. END OF THE MESOZOIC, GLOBAL
ANATOMICAL CHARACTERISTICS Very numerous, small
and slender teeth, upper teeth strongly procumbent, ANATOMICAL CHARACTERISTICS Heads not broad,
lower teeth directed sideways and a little downward. temporal region somewhat elongated. Necks moderate in
AGE Late Cretaceous; Maastrichtian. length.
DISTRIBUTION AND FORMATIONS Southern Argentina; HABITAT Freshwaters to continental shelves.
Paso del Sapo. HABITS Fishers of medium-sized game.
NOTES May be same as, or direct ancestor of, Nichollssaura borealis
A. quiriquinensis. 2.6 m (8.5 ft) TL, 80 kg (170 lb)
FOSSIL REMAINS Complete skull and skeleton, severely
Aristonectes quiriquinensis flattened top to bottom.
10 m (33 ft) TL, 4 tonnes ANATOMICAL CHARACTERISTICS Teeth medium sized.
FOSSIL REMAINS Partial skull and skeleton. Flippers medium sized, similar in size.
ANATOMICAL CHARACTERISTICS Very numerous, small AGE Early Cretaceous; early Albian.
and slender teeth, upper teeth strongly procumbent, DISTRIBUTION AND FORMATIONS Alberta; lower
lower teeth directed sideways and a little downward. Clearwater.
Flippers very large. HABITAT Interior seaway, polar.
AGE Late Cretaceous; late Maastrichtian. NOTES Shared its habitat with Athabascasaurus,
DISTRIBUTION AND FORMATIONS Central Chile; Wapuskanectes.
Quiriquina.
HABITAT Continental coastal. Hastanectes valdensis
Morturneria seymourensis FOSSIL REMAINS Two partial skeletons, possibly
Adult size uncertain immature.
FOSSIL REMAINS Partial skull and minority of skeleton, ANATOMICAL CHARACTERISTICS Insufficient
juvenile. information.
ANATOMICAL CHARACTERISTICS Very numerous, small AGE Early Cretaceous; Valanginian.
and slender teeth, upper teeth strongly procumbent, DISTRIBUTION AND FORMATIONS Southern England;
lower teeth directed sideways and a little downward. Wadhurst Clay.
AGE Late Cretaceous; late Maastrichtian. HABITAT Island estuary.
DISTRIBUTION AND FORMATIONS Antarctic Peninsula;
Lopez de Bertodano.
HABITAT Continental coastal, polar.
NOTES Shared its habitat with Kaikaifilu.
119

Brancasaurus brancai
Brancasaurus brancai AGE Early Cretaceous; Aptian and/or early Albian.

4.1 m (13 ft) TL, 350 kg (750 lb) DISTRIBUTION AND FORMATIONS South Australia;
FOSSIL REMAINS Majority of skull and skeleton, Bulldog Shale.
partial skull and skeleton. HABITAT Interior seaway, polar, lacking sharks.
ANATOMICAL CHARACTERISTICS Head fairly narrow, NOTES Shared its habitat with Opallionectes.
teeth medium sized, fairly stout. Tail short. Flippers very
large, similar in size, lower arm and leg very short, rest Opallionectes andamookaensis
of flippers elongated, strongly swept back. 5.5 m (19 ft) TL, 750 (1,600 lb)
AGE Early Cretaceous; middle to late Berriasian. FOSSIL REMAINS Teeth, partial skeleton.
DISTRIBUTION AND FORMATIONS Northwestern ANATOMICAL CHARACTERISTICS Teeth slender.
Germany; Bückeberg. AGE Early Cretaceous; Aptian and/or early Albian.
HABITAT Lakes, possibly rivers. DISTRIBUTION AND FORMATIONS South Australia;
HABITS Powerful swimmers. May have occasionally Bulldog Shale.
snapped up small swimming land creatures. HABITAT Interior seaway, polar, lacking sharks.
NOTES Whether this taxon was a permanent freshwater HABITS Consumed small prey.
plesiosaur or a transient is not certain. NOTES Phylogenetic position is not certain.
Umoonasaurus demoscyllus Leptocleidus capensis

2.5 m (8 ft) TL, 70 kg (150 lb) 4.5 m (15 ft) TL, 400 kg (1,300 lb)
FOSSIL REMAINS Partial skull and majority of skeleton. FOSSIL REMAINS Majority of skull, severely flattened
ANATOMICAL CHARACTERISTICS Teeth long and top to bottom, and minority of skeleton.
slender. Neck short. Last five tail vertebrae fused ANATOMICAL CHARACTERISTICS Insufficient
into a pygostyle. Flippers large. information.
AGE Early Cretaceous; Valanginian.
120

DISTRIBUTION AND FORMATIONS South Africa; DISTRIBUTION AND FORMATIONS South Dakota; lower
Sundays River. Greenhorn Limestone.
HABITAT Continental coastline. HABITAT Interior seaway rather narrow and shallow.
Leptocleidus superstes Edgarosaurus muddi

4 m (13 ft) TL, 300 kg (650 lb) 3.5 m (11 ft) TL, 250 kg (550 lb)
FOSSIL REMAINS Majority of skull and skeleton. FOSSIL REMAINS Majority of skull and minority of
ANATOMICAL CHARACTERISTICS Head fairly broad, skeleton.
snout rather short, teeth moderate in number, fairly large. ANATOMICAL CHARACTERISTICS Head elongated,
AGE Early Cretaceous; Barremian. snout long and narrow but robust, teeth large. Neck
DISTRIBUTION AND FORMATIONS Southern England; moderately long.
upper Weald Clay. AGE Early Cretaceous; middle Albian.
HABITAT Island archipelago shallows. DISTRIBUTION AND FORMATIONS Montana; middle
Thermopolis Shale.
Vectocleidus pastorum HABITAT Interior seaway narrow and shallow.
1.5 m (5 ft) TL, 15 kg (30 lb)
FOSSIL REMAINS Partial skeleton.
information. Edgarosaurus
AGE Early Cretaceous; latest Barremian. muddi
upper Vectis.
Plesiopleurodon wellesi
POLYCOTYLIDS 7 m (23 ft) TL, 2 tonnes
MEDIUM-SIZED TO LARGE LEPTOCLEIDIANS FOSSIL REMAINS Majority of skull and minority
FROM THE EARLY CRETACEOUS TO THE END of skeleton.
OF THE MESOZOIC, GLOBAL ANATOMICAL CHARACTERISTICS Teeth moderate
in size.
ANATOMICAL CHARACTERISTICS Necks fairly short. AGE Late Cretaceous; early Cenomanian.
HABITAT Coastal nearshore to deep oceans. DISTRIBUTION AND FORMATIONS Wyoming;
HABITS Swimming performance modest to good. Belle Fourche Shale.
Fishers and hunters of medium-sized to big game. HABITAT Interior seaway.
Mauriciosaurus fernandezi
FOSSIL REMAINS Skull and skeleton, juvenile, severely Plesiopleurodon
flattened top to bottom, soft tissues. wellesi
ANATOMICAL CHARACTERISTICS Head large,
elongated, snout long and narrow, teeth medium sized,
procumbent. Flippers large, similar in size.
AGE Late Cretaceous; early Turonian. Eopolycotylus rankini
DISTRIBUTION AND FORMATIONS Northeastern 4 m (13 ft) TL, 350 kg (700 lb)
Mexico; lower Agua Nueva. FOSSIL REMAINS Partial skull and skeleton.
HABITAT Continental shelf. ANATOMICAL CHARACTERISTICS Insufficient
information.
Pahasapasaurus haasi AGE Late Cretaceous; early Turonian.
Adult size uncertain DISTRIBUTION AND FORMATIONS Utah; middle
FOSSIL REMAINS Skull and skeleton, juvenile. Tropic Shale.
ANATOMICAL CHARACTERISTICS Head elongated, HABITAT Interior seaway.
snout long and narrow. NOTES Shared its habitat with Dolichorhynchops?
AGE Late Cretaceous; middle Cenomanian. tropicensis.
121

Polycotylus latipinnis DISTRIBUTION AND FORMATIONS Kansas; upper

5 m (16 ft) TL, 700 kg (1,500 lb) Greenhorn Limestone, lower Carlile Shale.
FOSSIL REMAINS Partial skeletons, including a probable HABITAT Interior seaway shifting from its greatest
fetus. maximum to less broad and deep.
ANATOMICAL CHARACTERISTICS Insufficient NOTES Shared its habitat with Brachauchenius. May have
information. been directly ancestral to Trinacromerum kirki.
AGE Late Cretaceous; earliest Campanian.
DISTRIBUTION AND FORMATIONS Kansas, Alabama; Trinacromerum kirki
uppermost Niobrara, lower Mooreville Chalk. 4.5 m (15 ft) TL, 500 kg (1,100 lb)
HABITAT Interior seaway was continuing to become less FOSSIL REMAINS Minority of skeleton.
broad and deep, continental shelf. ANATOMICAL CHARACTERISTICS Insufficient
NOTES The presence of a young skeleton within an adult information.
skeleton indicates live birth in the group, the calves being AGE Late Cretaceous; middle Turonian.
large in at least some plesiosaurs. Shared its habitat with DISTRIBUTION AND FORMATIONS Manitoba; upper
Styxosaurus snowii, Dolichorhynchops osborni, Selmasaurus Favel.
russelli, Platecarpus tympanicus, Eonatator sternbergii, HABITAT Interior seaway.
Clidastes propython, Ctenochelys stenoporus, Protostega
gigas. Manemergus anguirostris
2 m (6 ft) TL, 30 kg (60 lb)
Polycotylus? sopozkoi FOSSIL REMAINS Skull and majority of skeleton.
4 m (13 ft) TL, 350 kg (700 lb) ANATOMICAL CHARACTERISTICS Head elongated,
FOSSIL REMAINS Partial skull and skeleton. narrow, snout long, quite narrow and shallow, temporal
ANATOMICAL CHARACTERISTICS Fore and aft flippers region elongated and robustly constructed, teeth few,
similar in size. slender, and long.
AGE Late Cretaceous; early Campanian. AGE Late Cretaceous; late Turonian.
DISTRIBUTION AND FORMATIONS Central Russia; DISTRIBUTION AND FORMATIONS Morocco; Akrabou.
unnamed. HABITAT Continental coastal.
HABITAT Continental coastline. NOTES Shared its habitat with Thililua.
NOTES Placement in Polycotylus not certain.
Georgiasaurus penzensis
4 m (13 ft) TL, 450 kg (1,000 lb) Manemergus
FOSSIL REMAINS Partial skull and skeleton. anguirostris
information.
AGE Late Cretaceous; Santonian.
DISTRIBUTION AND FORMATIONS Western Russia;
unnamed.
HABITAT Continental coastline. Thililua longicollis
6 m (20 ft) TL, 1.3 tonnes
Trinacromerum bentonianum FOSSIL REMAINS Skull and minority of skeleton.
3 m (10 ft) TL, 150 kg (300 lb) ANATOMICAL CHARACTERISTICS Head elongated,
FOSSIL REMAINS Three skulls. low along entire length, robustly constructed, teeth
ANATOMICAL CHARACTERISTICS Head elongated, moderate in number and medium sized, fairly robust.
snout long and narrow, teeth rather small. Neck length moderate.
AGE Late Cretaceous; middle Turonian. AGE Late Cretaceous; late Turonian.
Trinacromerum
bentonianum Thililua longicollis
122

DISTRIBUTION AND FORMATIONS Morocco; Akrabou. Dolichorhynchops osborni

HABITAT Continental coastal. 5.2 m (17 ft) TL, 1 tonne
FOSSIL REMAINS Three skulls and skeletons.
Dolichorhynchops? tropicensis ANATOMICAL CHARACTERISTICS Head elongated and
3 m (10 ft) TL, 200 kg (450 lb) narrow, snout very long, modest crest along midline of
FOSSIL REMAINS Majority of skull and skeleton, minority temporal region, teeth fairly numerous, rather small, not
of skeleton. slender, procumbent, intermeshing. Neck moderately
ANATOMICAL CHARACTERISTICS Insufficient long. Trunk somewhat elongated. Tail short. Flippers
information. long, rather narrow, similar in size.
AGE Late Cretaceous; early Turonian. AGE Late Cretaceous; earliest Campanian.
DISTRIBUTION AND FORMATIONS Utah; middle Tropic DISTRIBUTION AND FORMATIONS Kansas; uppermost
Shale. Niobrara.
HABITAT Interior seaway. HABITAT Interior seaway was continuing to become less
NOTES Whether this species belongs to the same genus as broad and deep.
the much later D. osborni is problematic. Shared its NOTES Shared its habitat with Styxosaurus snowii,
habitat with Eopolycotylus. Polycotylus latipinnis, Platecarpus tympanicus, Eonatator
Dolichorhynchops osborni
123

sternbergii, Clidastes propython, Ctenochelys stenoporus, Sulcusuchus erraini

Protostega gigas. May be directly ancestral to D. bonneri. Adult size uncertain
FOSSIL REMAINS Two partial skulls, possibly immature.
Dolichorhynchops bonneri ANATOMICAL CHARACTERISTICS Head elongated,
4.8 m (15.5 ft) TL, 800 kg (1,750 lb) heavily constructed.
FOSSIL REMAINS Majority of skull and skeletons. AGE Late Cretaceous; late Campanian and/or
ANATOMICAL CHARACTERISTICS Head elongated and Maastrichtian.
narrow, snout very long, fairly large crest along midline of DISTRIBUTION AND FORMATIONS Southern Argentina;
temporal region, teeth fairly numerous, rather small, fairly Los Alamitos.
robust, intermeshing. Neck moderately long. HABITAT Continental coastal.
AGE Late Cretaceous; early Campanian. HABITS Fisher and hunter of big game, including hard-
DISTRIBUTION AND FORMATIONS Wyoming; lower shelled ammonoids.
Pierre Shale.
HABITAT Interior seaway was continuing to become less
broad and deep. ICHTHYOSAURO-
NOTES Shared its habitat with Styxosaurus browni,
Elasmosaurus, Tylosaurus proriger, Latoplatecarpus,
MORPHS
Globidens? dakotensis, Toxochelys latiremus. SMALL TO GIGANTIC NEODIAPSIDS FROM
THE EARLY TRIASSIC TO THE END OF
ANATOMICAL CHARACTERISTICS Highly variable.

Dolichorhynchops
Skulls akinetic, nostrils far behind snout tip and
bonneri
immediately in front of orbits, scleral rings often present,
lower temporal openings ventrally open, teeth neither
very large nor on mouth roof. Necks short. Tails flattened
side to side. Limbs modified into stiff-jointed partial
hydrofoils. Forelimb at least as long as aft. Primarily axial
Dolichorhynchops herschelensis undulators from anguilliform to thunniform, flippers
Adult size not certain primarily for stability and maneuvering.
FOSSIL REMAINS Majority of skull and partial skeleton, HABITAT Coastal nearshore to deep oceans.
possibly immature. HABITS Swimming performance mediocre to extremely
ANATOMICAL CHARACTERISTICS Head elongated and high. Predaceous, ambush and/or pursuit fishers and
narrow, snout very long, fairly large crest along midline of hunters of small to big game, some crushers.
temporal region, teeth fairly numerous.
AGE Late Cretaceous; late Campanian and/or early
Maastrichtian. HUPEHSUCHIANS
DISTRIBUTION AND FORMATIONS Saskatchewan; lower
Bearpaw Shale. SMALL ICHTHYOSAUROMORPHS FROM
HABITAT Interior seaway was becoming much narrower. THE EARLY TRIASSIC OF ASIA
NOTES Shared is habits with Albertonectes, Prognathodon?
overtoni. ANATOMICAL CHARACTERISTICS Skeletons very heavy
boned. Necks not very short. Trunks long, narrow, not
Palmulasaurus quadratus deep, neural spine/hornlet complexes elevated, gastralia
Adult size not certain massive. Tails long, straight, and unexpanded aft. Arms
FOSSIL REMAINS Partial skull and skeleton, possibly larger than legs, which are not full flippers, rigidly jointed,
immature. presumably webbed. Moderately hydrodynamically stream-
ANATOMICAL CHARACTERISTICS Snout very slender. lined, transitional anguilliform-carangiform swimmers.
Foreflipper smaller than aft. HABITAT Coastal and brackish shorelines, lagoons, reefs,
AGE Late Cretaceous; earliest Turonian. estuaries.
DISTRIBUTION AND FORMATIONS Utah; lower Tropic HABITS Swimming performance mediocre to modest.
Shale. Ambush and/or pursuit fishers of small to medium-sized
HABITAT Interior seaway. game.
124

I C H T H Y O S A U R O M O R P H S
NANCHANGOSAURIDS HUPEHSUCHIDS
SMALL HUPEHSUCHIANS FROM THE EARLY SMALL HUPEHSUCHIANS FROM THE EARLY
TRIASSIC OF ASIA TRIASSIC OF ASIA
ANATOMICAL CHARACTERISTICS Heads elongated,

narrow, snout a highly elongated, very shallow, fairly HUPEHSUCHINES
broad, toothless beak, upper beak longer than lower.
SMALL HUPEHSUCHIDS FROM THE EARLY
Nanchangosaurus suni TRIASSIC OF ASIA
1 m (3 ft) TL, 1.5 kg (3 lb)
FOSSIL REMAINS Two partial skulls and skeletons, ANATOMICAL CHARACTERISTICS Upper and lower
severely flattened top to bottom, probable juvenile skull beaks same length. Neural spines/armor hornlets
and majority of skeleton. complex, very tall. Arms large.
ANATOMICAL CHARACTERISTICS Neural spines/armor
hornlets complex, moderately tall. Hupehsuchus nanchangensis
AGE Early Triassic; late Olenekian. 0.95 m (3.1 ft) TL, 2.5 kg (5.5 lb)
DISTRIBUTION AND FORMATIONS Eastern China; FOSSIL REMAINS Two skulls and skeletons.
upper Jialingjiang. ANATOMICAL CHARACTERISTICS Standard for group.
HABITAT Continental nearshore. AGE Early Triassic; late Olenekian.
NOTES Eohupehsuchus brevicollis may be the juvenile of DISTRIBUTION AND FORMATIONS Eastern China;
this species. Shared its habitat with Eohupehsuchus, upper Jialingjiang.
Parahupehsuchus, Hupehsuchus, Eretmorhipis, HABITAT Continental nearshore.
Chaohusaurus zhangjiawanensis.
Hupehsuchus nanchangensis and Eretmorhipis (right)
125

Hupehsuchus nanchangensis
NOTES Shared its habitat with Nanchangosaurus, AGE Early Triassic; late Olenekian.
Eohupehsuchus, Parahupehsuchus, Eretmorhipis, DISTRIBUTION AND FORMATIONS Eastern China;
Chaohusaurus zhangjiawanensis. upper Jialingjiang.
PARAHUPEHSUCHINES HABITS Insufficient information.
NOTES Shared its habitat with Nanchangosaurus,
SMALL HUPEHSUCHIDS FROM THE EARLY Eohupehsuchus, Hupehsuchus, Eretmorhipis, Chaohusaurus
TRIASSIC OF ASIA zhangjiawanensis.
ANATOMICAL CHARACTERISTICS Trunks elongated, Eretmorhipis carrolldongi

very massively constructed. 0.93 m (3 ft) TL, 2 kg (4.5 lb)
NOTES Without the skull of Parahupehsuchus, is not clear FOSSIL REMAINS Skull and two skeletons.
whether it and Eretmorhipis belong in same subfamily. ANATOMICAL CHARACTERISTICS Head lightly built,
small, flattened, subrectangular in top view with duck-like
Parahupehsuchus longus beak that includes central floating bone between beak
1 m (3.3 ft) TL, 2 kg (4.5 lb) prongs, orbits small, mechanoreceptors apparently on
FOSSIL REMAINS Majority of skeleton. beak. Tail rather rigid. Forepaddle especially large.
ANATOMICAL CHARACTERISTICS Limbs not large. AGE Early Triassic; late Olenekian.
Eretmorhipis carrolldongi
126

DISTRIBUTION AND FORMATIONS Eastern China; HABITAT Coastal shallows.

upper Jialingjiang. HABITS Swimming performance good. Ambush and
HABITAT Continental nearshore. pursuit fishers of small and medium-sized game.
HABITS Limbs about as important for swimming as axial
undulations. Apparently used electrical field Omphalosaurus nevadanus
mechanoreceptors to detect small prey items, especially 2 m (6 ft) TL, 25 kg (50 lb)
arthropods, probably at night. FOSSIL REMAINS Minority of skull and skeleton.
NOTES Shared its habitat with Nanchangosaurus, ANATOMICAL CHARACTERISTICS Standard for group.
Eohupehsuchus, Hupehsuchus, Parahupehsuchus, AGE Middle Triassic; Anisian.
Chaohusaurus zhangjiawanensis. Head in particular DISTRIBUTION AND FORMATIONS Nevada; Prida.
remarkably convergent with that of the duck-billed HABITAT Continental shallows.
platypus. Shows that very sophisticated reptilian nocturnal NOTES The placement of other poorly preserved
aquatic hunter of small items evolved quickly after P/T specimens in the genus is uncertain.
extinction, but apparently failed to do so again during
Mesozoic. Tholodus schmidi
1.5 m (5 ft) TL, 10 kg (20 lb)
FOSSIL REMAINS Partial remains.
ICHTHYOSAURIFORMES ANATOMICAL CHARACTERISTICS Standard for group.
SMALL TO GIGANTIC ICHTHYOSAUROMORPHS DISTRIBUTION AND FORMATIONS Southern Germany;
FROM THE EARLY TRIASSIC TO THE END OF lower Muschelkalk.
THE MESOZOIC, GLOBAL HABITAT Island archipelago shallows.
NOTES Shared its habitat with Cyamodus tarnowitzensis,
ANATOMICAL CHARACTERISTICS Fairly uniform. All Contectopalatus.
streamlined, fish- or cetacean-like. Snouts almost always
narrow, scleral rings may be universal, teeth sometimes have NASOROSTRANS
bulbous roots. Neck very short. Trunks consist of a very
large number of vertebrae and ribs, trunks never broader SMALL ICHTHYOSAURIFORMES FROM
than deep and never extremely deep. Tail never very long, THE EARLY TRIASSIC OF ASIA
aft downward flexure supported a soft upper tail fin. Scapula
short, rounded. Hydrodynamically streamlined, transitional ANATOMICAL CHARACTERISTICS Heads short, broad,
anguilliform-carangiform to thunniform swimmers. Very very triangular, with a very short, pointed snout and large
small, fine scales formed a smooth skin texture. temporal region, lower jaw shallow, toothless. Trunk
HABITAT Coastal nearshore to deep oceans. Probably all moderately long, ribs and especially gastralia robust. Tall
gave live birth. vertebrae at modest downward flexure of aft tail
HABITS Swimming performance good to extremely high. supported a modest upper tail fin. Pelvis attached to
Ambush and/or pursuit fishers and hunters of small to vertebral column. Flipper elements from wrist and ankle
big game, some crushers. below very poorly ossified. Transitional anguilliform-
carangiform swimmers.
OMPHALOSAURIDS HABITAT Coastal shallows.
HABITS Swimming performance good. Appear to have
SMALL ICHTHYOSAURIFORMES FROM THE been suction feeders.
MIDDLE TRIASSIC OF NORTH AMERICA
AND ASIA Sclerocormus parviceps
1.5 m (5 ft) TL, 9 kg (20 lb)
ANATOMICAL CHARACTERISTICS Teeth irregularly FOSSIL REMAINS Skull and skeleton severely flattened
placed, small buttons. Trunk moderately long. Tall top to bottom, possible juvenile skull and majority of
vertebrae at modest downward flexure of aft tail skeleton.
supported a modest upper tail fin. Pelvis attached to ANATOMICAL CHARACTERISTICS Head very small.
vertebral column. Transitional anguilliform-carangiform Tail fairly long. Foreflipper very large and larger than aft.
swimmers. AGE Early Triassic; late Olenekian.
NOTES Absence from at least some other seas may reflect DISTRIBUTION AND FORMATIONS Eastern China;
lack of sufficient sampling. upper Nanlinghu.
127

Sclerocormus parviceps
HABITAT Continental shallows. UTATSUSAURIANS

NOTES Cartorhynchus lenticarpus is probably the juvenile
of this or a closely related species. Shared its habitat with SMALL ICHTHYOPTERYGIANS FROM THE
Majiashanosaurus, Chaohusaurus geishanensis. EARLY AND POSSIBLY MIDDLE TRIASSIC OF
NORTH AMERICA
ICHTHYOPTERYGIANS
ANATOMICAL CHARACTERISTICS Snouts at least
SMALL TO GIGANTIC ICHTHYOSAURIFORMES somewhat spike shaped. Trunks moderately long. Tall
FROM THE EARLY TRIASSIC TO THE END OF vertebrae at modest downward flexure of aft tail supported
THE MESOZOIC, GLOBAL a modest upper tail fin. Pelvis attached to vertebral column.
Transitional anguilliform-carangiform swimmers.
ANATOMICAL CHARACTERISTICS Snouts almost always HABITAT Coastal shallows.
narrow. Vertebral centra spool shaped. Limbs are clawless HABITS Swimming performance good. Ambush and
full flippers with increased number of distal elements. pursuit fishers of small and medium-sized game.
Transitional anguilliform-carangiform to thunniform
swimmers. Utatsusaurus hataii
HABITAT Coastal nearshore to deep oceans. 2.5 m (8 ft) TL, 45 kg (100 lb)
HABITS Swimming performance good to extremely high. FOSSIL REMAINS Skulls and skeletons.
Ambush and/or pursuit fishers and hunters of small to ANATOMICAL CHARACTERISTICS Snout long and
big game. Some crushers, some may have been suckers. somewhat spike shaped, teeth very numerous and small.
Flippers similar in size, distal flipper elements poorly
ossified.
AGE Early Triassic.
128

Utatsusaurus hataii
DISTRIBUTION AND FORMATIONS Central Japan; upper tail fin. Pelvis attached to vertebral column.
Osawa. Transitional anguilliform-carangiform swimmers.
HABITAT Continental shallows. HABITAT Coastal shallows.
HABITS Swimming performance good. Mollusk crushers,
Parvinatator wapitiensis also perhaps ambush and pursuit fishers of small and
1 m (3.3 ft) TL, 3 kg (6.5 lb) medium-sized game.
FOSSIL REMAINS Skull and minority of skeleton. NOTES Absence from at least some other seas may reflect
ANATOMICAL CHARACTERISTICS Snout spikelike, lack of sampling.
orbit very large, aft lower jaw massive, teeth moderate in
number and fairly large. Distal flipper elements ossified, Xinminosaurus catactes
foreflipper swept back distally. 2.3 m (7.5 ft) TL, 30 kg (65 lb)
AGE Late Early or Middle Triassic. FOSSIL REMAINS Disrupted skull and skeleton, severely
DISTRIBUTION AND FORMATIONS British Columbia; flattened top to bottom.
Sulphur Mountain. ANATOMICAL CHARACTERISTICS Anterior jaws may be
HABITAT Continental shallows. toothless. Distal paddle elements ossified, fore and aft
NOTES Exact stratigraphic level of taxon not certain. paddles similarly small.
upper Guanling.
Parvinatator NOTES Shared its habitat with Largocephalosaurus,
wapitiensis Sinosaurosphargis, Panzhousaurus, Wumengosaurus,
Nothosaurus yangjuanensis, Barracudasauroides.
EOICHTHYOSAURS
XINMINOSAURS SMALL TO GIGANTIC ICHTHYOSAURIFORMES
FROM THE EARLY TRIASSIC TO THE END OF
SMALL ICHTHYOPTERYGIANS FROM THE MESOZOIC, GLOBAL
THE MIDDLE TRIASSIC OF ASIA
HABITAT Coastal nearshore to deep oceans.
ANATOMICAL CHARACTERISTICS Midteeth form a HABITS Swimming performance good to extremely high.
flattened pavement. Trunks moderately long. Tall vertebrae Ambush and/or pursuit fishers and hunters of small to
at modest downward flexure of aft tail supported a modest big game, some crushers.
129

GRIPPIIDS AGE Early Triassic; late Olenekian.

DISTRIBUTION AND FORMATIONS Svalbard; lower
SMALL EOICHTHYOSAURS FROM THE MIDDLE Vikinghogda.
TRIASSIC OF THE NORTHERN HEMISPHERE HABITAT Open continental shelf, polar.
ANATOMICAL CHARACTERISTICS Heads rather short, Chaohusaurus zhangjiawanensis

snouts short, deep aft because orbits very large. Trunks 0.9 m (3 ft) TL, 1.8 kg (4 lb)
moderately long. Tall vertebrae at modest downward FOSSIL REMAINS Two partial skulls and complete
flexure of aft tail supported a modest upper tail fin. skeletons, severely flattened top to bottom, including
Transitional anguilliform-carangiform swimmers. probable fetuses.
HABITAT Coastal shallows. ANATOMICAL CHARACTERISTICS Foreflipper large,
HABITS Swimming performance good. Ambush and much larger than aft, distal elements in both poorly ossified.
pursuit fishers of small game. AGE Early Triassic; late Olenekian.
NOTES Grippiids show that truly marine swimmers DISTRIBUTION AND FORMATIONS Eastern China;
evolved just a few million years after the P/T extinction. upper Jialingjiang.
Gulosaurus helmi NOTES Presence of small apparent fetal remains in and
Adult size uncertain exiting adult skeleton indicates live birth, headfirst.
FOSSIL REMAINS Skull and partial skeleton, juvenile. Shared its habitat with Nanchangosaurus, Eohupehsuchus,
ANATOMICAL CHARACTERISTICS Insufficient Hupehsuchus, Parahupehsuchus, Eretmorhipis.
information.
AGE Early Triassic; Olenekian.
middle Sulphur Mountain.
Grippia longirostris Grippia longirostris

1.5 m (5 ft) TL, 8 kg (18 lb)
FOSSIL REMAINS Partial skulls and skeletons.
ANATOMICAL CHARACTERISTICS Snout not slender,
only a few small teeth near front of upper jaw.
Chaohusaurus geishanensis
130

Chaohusaurus geishanensis
0.9 m (3 ft) TL, 1.8 kg (4 lb)
CYMBOSPONDYLIDS
FOSSIL REMAINS A number of skulls and skeletons. LARGE TO GIGANTIC ICHTHYOSAURS FROM
ANATOMICAL CHARACTERISTICS Snout slender, teeth THE EARLY TO MIDDLE TRIASSIC OF NORTH
fairly numerous, small. Foreflipper large, much larger than AMERICA AND EUROPE
aft, distal elements in both poorly ossified.
AGE Middle Triassic; late Olenekian. ANATOMICAL CHARACTERISTICS Heads solidly
DISTRIBUTION AND FORMATIONS Eastern China; constructed, snouts long, temporal region greatly
upper Nanlinghu. expanded. Trunks elongated and very shallow. Tall
HABITAT Continental shallows. vertebrae at modest downward flexure of aft tail
NOTES May include C. chaoxianensis and C. supported a modest upper tail fin. Distal flipper elements
brevifemoralis. Shared its habitat with Majiashanosaurus, poorly ossified. Hydrodynamically well-streamlined,
Sclerocormus. transitional anguilliform-carangiform swimmers.
HABITAT Coastal to deep ocean.
ICHTHYOSAURS HABITS Swimming performance good to very good.
Pursuit fishers and hunters of medium-sized and especially
SMALL TO GIGANTIC EOICHTHYOSAURS FROM big game.
THE EARLY TRIASSIC TO THE END OF THE NOTES First large, big-game-hunting marine reptiles.
MESOZOIC, GLOBAL Placement of many genera in family questionable.
Absence from at least some other seas may reflect lack of
sufficient sampling.
Quasianosteosaurus vikinghoegdai
5 m (16 ft) TL, 500 kg (1,000 lb)
FOSSIL REMAINS Partial skull.
information.
AGE Early Triassic; late Olenekian.
DISTRIBUTION AND FORMATIONS Svalbard; upper
Cymbospondylus (Ichthyosaurs)
Sticky Keep.
ANATOMICAL CHARACTERISTICS Pelvis not attached to NOTES Earliest known large ichthyosaur and sea reptile.
vertebral column, upper pelvic element a splint. Flippers
better developed, with shortened lower arm and increased Unnamed genus and species
distal complexes. 9.6 m (31 ft) TL, 2.8 tonnes
HABITAT Coastal nearshore to deep oceans. FOSSIL REMAINS Nearly complete skull and skeleton.
HABITS Swimming performance good to extremely high. ANATOMICAL CHARACTERISTICS Head medium sized,
Ambush and/or pursuit fishers and hunters of small to moderately broad, snout very long, robust, teeth
big game, some crushers. numerous, fairly small.
NOTES Includes the largest and the fastest sea reptiles. AGE Middle Triassic; late Anisian.
Unnamed genus
and species
131

Unnamed genus
and species
DISTRIBUTION AND FORMATIONS Nevada; upper NOTES Placement in same genus as above unnamed
Favret. genus and species incorrect because of very different
HABITAT Continental shelf. shoulder girdles, may not be member of same family.
HABITS Swimming performance good.
NOTES Has long been Cymbospondylus petrinus, but Unnamed genus buchseri
original specimens too fragmentary to base a genus and 8 m (26 ft) TL, 1.5 tonnes
species on. C. nichollsi may be young of this or other species FOSSIL REMAINS Majority of skull and partial skeleton.
from the same level of the Favret Formation. A partial ANATOMICAL CHARACTERISTICS Foreflipper fairly
skull may be an individual 14 m (46 ft) long. Shared its large.
habitat with Augustasaurus, Phalarodon, Thalattoarchon, AGE Middle Triassic; latest Anisian and/or earliest
unnamed genus duelferi, unnamed genus youngorum. Ladinian.
DISTRIBUTION AND FORMATIONS Switzerland; Besano.
Unnamed genus youngorum HABITAT Island archipelago shallows.
14 m (42 ft) TL, 9 tonnes NOTES Original placement in Cymbospondylus incorrect,
FOSSIL REMAINS Skull and minority of skeleton. nearly identical shoulder girdles indiacate is close relative
ANATOMICAL CHARACTERISTICS Head moderately of above unnamend genus and species. Shared its habitat
broad, snout long, robust, teeth numerous, fairly small. with Askeptosaurus, Helveticosaurus, Paraplacodus,
AGE Middle Triassic; late Anisian. Serpianosaurus, Wimanius, Besanosaurus, Mixosaurus
DISTRIBUTION AND FORMATIONS Nevada; upper cornalianus, M.? kuhnschnyderi.
Favret.
HABITAT Continental shelf. Thalattoarchon saurophagis
NOTES Placement in same genus as prior unnamed taxon 8.6 m (28 ft) TL, 2 tonnes
incorrect because of very different shoulder girdles, may FOSSIL REMAINS Partial skull and skeleton.
not be member of same family. Earliest known gigantic ANATOMICAL CHARACTERISTICS Head robustly
sea reptile, but claim of mass of 45 tonnes is greatly constructed, teeth large.
exaggerated. AGE Middle Triassic; middle Anisian.
DISTRIBUTION AND FORMATIONS Nevada; upper
Favret.
HABITAT Continental shelf.
unnamed genus NOTES Shared its habitat withAugustasaurus, unnamed
youngorum genus and species, unnamed genus duelferi, unnamed
genus youngorum, Phalarodon. First known archpredatory
marine reptile.
Unnamed genus duelferi Phantomosaurus neubigi

5 m (17 ft) TL, 400 kg (900 lb) 6 m (20 ft) TL, 700 kg (1,500 lb)
FOSSIL REMAINS Skull and minority of skeleton. FOSSIL REMAINS Partial skull and skeleton, badly
ANATOMICAL CHARACTERISTICS Head moderately damaged.
broad, snout long, robust, teeth fairly large. ANATOMICAL CHARACTERISTICS Insufficient
AGE Middle Triassic; late Anisian. information.
DISTRIBUTION AND FORMATIONS Nevada; upper AGE Middle Triassic; late Anisian.
Favret. DISTRIBUTION AND FORMATIONS Southern Germany;
HABITAT Continental shelf. upper Muschelkalk.
132

HABITAT Island archipelago shallows. Contectopalatus atavus

NOTES Shared its habitat with Placodus, Nothosaurus 5 m (16 ft) TL, 400 kg (900 lb)
mirablis, N. giganteus, Pistosaurus. FOSSIL REMAINS Partial skull.
ANATOMICAL CHARACTERISTICS Head possibly very
MIXOSAURIDS narrow, tall subtriangular midline crest atop temporal
region, teeth medium sized.
SMALL TO MEDIUM-SIZED ICHTHYOSAURS AGE Middle Triassic; middle Anisian.
FROM THE MIDDLE TO LATE TRIASSIC DISTRIBUTION AND FORMATIONS Southern Germany;
OF THE NORTHERN HEMISPHERE lower Muschelkalk.
ANATOMICAL CHARACTERISTICS Heads large, snouts NOTES Shared its habitat with Cyamodus tarnowitzensis,
long and slender. Tall vertebrae at modest downward Tholodus.
flexure of aft tail supported a modest upper tail fin. Distal
flipper elements ossified, foreflippers much larger than aft. Barracudasauroides panxianensis
Dorsal fin may be present. Transitional anguilliform- 1.23 m (4 ft) TL, 8 kg (18 lb)
carangiform swimmers. FOSSIL REMAINS Three nearly complete or partial skulls
HABITAT Coastal shallows. and skeletons.
HABITS Swimming performance good. Ambush and ANATOMICAL CHARACTERISTICS Head robust, orbits
pursuit fishers of small game, some crushers. very large, low midline crest atop temporal region, teeth
NOTES A poorly preserved dorsal fin may indicate moderate in number and robust.
presence on at least some basal ichthyosaurs. Absence AGE Middle Triassic; middle Anisian.
from Southern Hemisphere may reflect lack of sufficient DISTRIBUTION AND FORMATIONS Southern China;
sampling. upper Guanling.
HABITS Hunted larger game.
Barracudasauroides
panxianensis
133

Barracudasauroides
panxianensis
134

Mixosaurus cornalianus
NOTES Shared its habitat with Largocephalosaurus, DISTRIBUTION AND FORMATIONS Switzerland-Italy
Sinosaurosphargis, Panzhousaurus, Wumengosaurus, border; Besano.
Nothosaurus yangjuanensis, Xinminosaurus. HABITAT Island archipelago shallows.
HABITS Also crushed small mollusks.
Mixosaurus cornalianus
1.85 m (6 ft) TL, 24 kg (50 lb) Phalarodon fraasi
FOSSIL REMAINS Numerous skulls and skeletons, 1.25 m (4 ft) TL, 7 kg (15 lb)
including possible fetuses, soft tissues. FOSSIL REMAINS A few skulls and skeletons, complete
ANATOMICAL CHARACTERISTICS Head not robustly to partial.
constructed, snout quite long and slender, no crest atop ANATOMICAL CHARACTERISTICS Snout fairly long
temporal region, teeth numerous and small. and shallow, orbits very large, tips of jaws toothless,
AGE Middle Triassic; latest Anisian and earliest Ladinian. front teeth small robust spikes, aft teeth somewhat
DISTRIBUTION AND FORMATIONS Switzerland-Italy larger blunt knobs.
border; Besano. AGE Middle Triassic; middle Anisian.
HABITAT Island archipelago shallows. DISTRIBUTION AND FORMATIONS Nevada; upper
NOTES The classic early small ichthyosaur. Possible fetuses Favret, lower Prida.
may indicate live birth. Dorsal fin may be present on a HABITAT Continental shallows.
specimen. Shared its habitat with Askeptosaurus, HABITS Also crushed mollusks.
Helveticosaurus, Paraplacodus, Serpianosaurus, Wimanius, NOTES Probably includes P. callawayi. Is not certain
Besanosaurus, unnamed genus buchseri, M.? kuhnschnyderi. whether this species had a midline aft head crest. Shared
its habitat with Augustasaurus, unnamed genus and
Mixosaurus (or Sangiorgiosaurus) species, unnamed genus duelferi, unnamed genus
kuhnschnyderi youngorum, Thalattoarchon.
1 m (3.3 ft) TL, 4 kg (9 lb)
FOSSIL REMAINS Skull and skeletons, severely flattened
top to bottom.
ANATOMICAL CHARACTERISTICS A few aft teeth are Phalarodon fraasi
stout knobs.
AGE Middle Triassic; latest Anisian and earliest Ladinian.
135

Phalarodon? nordenskioeldii HABITS Swimming performance good. Ambush and

1.5 m (5 ft) TL, 10 kg (20 lb) pursuit fishers of small game.
FOSSIL REMAINS A few partial skulls and skeletons. NOTES Absence from at least some other seas may reflect
ANATOMICAL CHARACTERISTICS Snout fairly long and lack of sufficient sampling.
shallow, orbits very large; large, long, subtriangular
midline crest atop temporal region, tips of jaws toothless, Qianichthyosaurus xingyiensis
front teeth small robust spikes, aft teeth markedly larger 1.3 m (4 ft) TL, 10 kg (20 lb)
blunt knobs. FOSSIL REMAINS Two skulls and skeletons.
AGE Late Triassic; early Carnian. ANATOMICAL CHARACTERISTICS Standard for group.
DISTRIBUTION AND FORMATIONS Svalbard; AGE Middle Triassic; late Ladinian.
Tschermakfjellet. DISTRIBUTION AND FORMATIONS Southeastern China;
HABITAT Open continental shelf, polar. lower Falang.
HABITS Specialized for crushing mollusks. HABITAT Continental nearshore.
NOTES Whether this species belongs in the much earlier NOTES Shared its habitat with Keichousaurus, Nothosaurus
and distinctive Phalarodon is problematic. youngi, Lariosaurus, Qianxisaurus, Wangosaurus. May be
the direct ancestor of Qianichthyosaurus zhoui.
Phalarodon? Qianichthyosaurus
nordenskioeldii xingyiensis
Wimanius odontopalatus Qianichthyosaurus zhoui

1 m (3.3 ft) TL, 4 kg (9 lb) 2 m (6.5 ft) TL, 35 kg (75 lb)
FOSSIL REMAINS Majority of skull. FOSSIL REMAINS Numerous skulls and skeletons.
ANATOMICAL CHARACTERISTICS Snout slender, orbits ANATOMICAL CHARACTERISTICS Standard for group.
very large, aft skull deep, teeth numerous and rather AGE Late Triassic; Carnian.
small. DISTRIBUTION AND FORMATIONS Southeastern China;
AGE Middle Triassic; latest Anisian and earliest Ladinian. upper Falang.
DISTRIBUTION AND FORMATIONS Switzerland-Italy HABITAT Continental nearshore. Shared its habitat with
border; Besano. Anshunsaurus huangguoshuensis, Miodentosaurus,
HABITAT Island archipelago shallows. Yunguisaurus, Guanlingsaurus, Guizhouichthyosaurus,
NOTES May be in its own family. Shared its habitat with unnamed genus orientalis.
Askeptosaurus, Helveticosaurus, Paraplacodus,
Serpianosaurus, Besanosaurus, unnamed genus buchseri, Toretocnemus californicus
Mixosaurus cornalianus, M.? kuhnschnyderi. 2 m (6.5 ft) TL, 35 kg (75 lb)
FOSSIL REMAINS Skeleton lacking skull.
TORETOCNEMIDS ANATOMICAL CHARACTERISTICS Insufficient
information.
SMALL TO LARGE ICHTHYOSAURS FROM AGE Late Triassic; Carnian.
THE MIDDLE TO LATE TRIASSIC OF DISTRIBUTION AND FORMATIONS Northern California;
NORTH AMERICA AND ASIA Hosselkus Limestone.
ANATOMICAL CHARACTERISTICS Heads large, snouts NOTES May include T. zitelli.
long and slender, orbits very large. Aft head deep, teeth
numerous and small. Tall vertebrae at modest downward Besanosaurus leptorhynchus
flexure of aft tail supported a modest upper tail fin. Distal 6 m (20 ft) TL, 1 tonne
flipper elements ossified, flippers fairly large, fore not FOSSIL REMAINS A few specimens from nearly complete
much larger than aft. Transitional anguilliform- to partial skull and skeleton, other skeleton severely
carangiform swimmers. disrupted, and partial probable fetus.
HABITAT Coastal shallows. ANATOMICAL CHARACTERISTICS As for group.
136

Qianichthyosaurus zhoui
AGE Middle Triassic; latest Anisian and earliest Ladinian. NOTES Usually considerd shastasaurids, but differences in
DISTRIBUTION AND FORMATIONS Northern Italy; shoulder girdle indicate otherwise. Length of snout and
Besano. presence/absence of teeth not known. Estimates that
HABITAT Continental coastal. fragmentary remains indicate that some shastasaurs
NOTES Includes Mikadocephalus. Shared its habitat with reached 26 m (85 ft) and the mass of blue whales are
Askeptosaurus, Helveticosaurus, Paraplacodus, highly exaggerated. Absence from at least some other
Serpianosaurus, Wimanius, unnamed genus buchseri, seas may reflect lack of sufficient sampling.
Mixosaurus cornalianus, M.? kuhnschnyderi.
Unnamed genus sikanniensis
16.6–17.9 m (54–59 ft) TL, 16 tonnes
Besanosaurus ANATOMICAL CHARACTERISTICS Head very large.
leptorhynchus AGE Late Triassic; middle Norian.
upper Pardonet.
MEGAMARINASAURIDS NOTES Belongs to neither much earlier and significantly
different genus Shastasaurus, nor earlier and very different
GIGANTIC ICHTHYOSAURS FROM THE Shonisaurus, both of which this species has been placed
LATE TRIASSIC OF NORTH AMERICA in. Shared its habitat with Macgowania, Hudsonelpidia.
The longest and most massive known sea reptile—or
ANATOMICAL CHARACTERISTICS Trunks shallow, nondinosaur “reptile”—but greatly exaggerated claims
downward flexure of aft tail modest. Distal flipper of 21 m (70 ft) and over 80 tonnes stem from major
elements probably unossified. Transitional anguilliform- misreadings of specimen measurements; uncertain total
carangiform swimmers. length depends in part on unknown length of snout.
137

GUANLINGASAURIDS HABITAT Continental nearshore.

NOTES Shared its habitat with Anshunsaurus
LARGE ICHTHYOSAURS FROM THE huangguoshuensis, Miodentosaurus, Yunguisaurus,
LATE TRIASSIC OF NORTH AMERICA Qianichthyosaurus zhoui, Guizhouichthyosaurus, unnamed
genus orientalis.
ANATOMICAL CHARACTERISTICS Snouts short and
triangular in top view, aft head broad, toothless. Trunks SHONISAURIDS
elongated and very shallow, downward flexure of aft tail
slight. Distal flipper elements poorly ossified. Anguilliform GIGANTIC ICHTHYOSAURS FROM THE
swimmers. LATE TRIASSIC OF NORTH AMERICA
HABITS May have been suction feeders. Swimming
performance good. ANATOMICAL CHARACTERISTICS Heads long, shallow,
NOTES Usually considerd shastasaurids, but differences in not lightly built, snouts very long, orbits modest sized,
aft skull and shoulder girdle indicate otherwise. Feeding lower jaw a little deeper than skull, teeth few, limited to
mode disputed. Absence from at least some other seas front of jaws, small. Trunk fairly deep, downward flexure
may reflect lack of sufficient sampling. of aft tail modest.
HABITS Fishers of medium-sized game. Swimming
Guanlingsaurus liangae performance moderate.
8.3 (27 ft) TL, 2.5 tonnes NOTES Not shastasaurids, as has been proposed.
FOSSIL REMAINS A few skulls and skeletons complete to
nearly so, adult and juvenile, skeletons severely crushed Shonisaurus popularis
top to bottom. 13.7 m (45 ft) TL, 16 tonnes
ANATOMICAL CHARACTERISTICS Trunk very shallow. FOSSIL REMAINS Over three dozen partial specimens.
Foreflippers long, hind flippers medium sized. ANATOMICAL CHARACTERISTICS Flippers fairly long,
AGE Late Triassic; Carnian. fore a little larger than aft, tail not large.
DISTRIBUTION AND FORMATIONS Southeastern China; AGE Late Triassic; latest Carnian, possibly earliest Norian.
upper Falang. DISTRIBUTION AND FORMATIONS Nevada; middle
Luning.
NOTES The classic gigantic early ichthyosaur. Past
restorations have tended to exaggerate the depth of the
body, in part because the dorsal column was posed too
Guanlingsaurus straight while the ribs were oriented too vertically. May
liangae compete with longer but more slenderly built unnamed
genus sikanniensis as most massive known pelagic reptile,
but markedly higher size estimates for known specimens
are unlikely.
Shonisaurus popularis
138

SHASTASAURIDS NONPARVIPELVIAN
ICHTHYOSAUR MISCELLANEA
LARGE TO GIGANTIC ICHTHYOSAURS FROM
THE LATE TRIASSIC OF NORTH AMERICA
AND ASIA Californosaurus perrini
3 m (10 ft) TL, 100 kg (200 lb)
ANATOMICAL CHARACTERISTICS Snouts slender and FOSSIL REMAINS Majority of skeleton lacking skull.
pointed. Trunks elongated and shallow, tail fairly long, aft ANATOMICAL CHARACTERISTICS Insufficient
portion flexed more downward than usual in basal information.
ichthyosaurs. Distal flipper elements ossified. Anguilliform AGE Late Triassic; middle Carnian.
swimmers. DISTRIBUTION AND FORMATIONS Northern California;
HABITAT Coastal, possibly deeper ocean. lower Hosselkus Limestone.
HABITS Swimming performance good. HABITAT Continental coastal.
NOTES Often considered to contain short-snouted, HABITS Swimming performance very good or excellent.
toothless guanlingsaurs, but differences in aft skull and Fisher of small to medium-sized game.
shoulder girdle indicate otherwise. Absence from at least
some other seas may reflect lack of sufficient sampling. PARVIPELVIANS
Shastasaurus pacificus SMALL TO GIGANTIC ICHTHYOSAURS FROM
7 m (23 ft) TL, 1.5 tonnes THE LATE TRIASSIC TO THE END OF THE
FOSSIL REMAINS Partial skull and minority of skeleton. MESOZOIC, GLOBAL
information. ANATOMICAL CHARACTERISTICS Fairly uniform. Heads
AGE Late Triassic; late Carnian. narrow. Trunks compact, gastralia do not continue aft to
DISTRIBUTION AND FORMATIONS Northern California; pelvis. Slender vertebrae of aft tail sharply flexed
middle Hosselkus Limestone. downward, supported large, soft upper fin that was dorsal
HABITAT Continental coastal. half of half-moon-shaped tail fluke. Shoulder and pelvic
NOTES Shared its habitat with Nectosaurus, girdles reduced. Lower flipper elements ossified and
Thalattosaurus, Toretocnemus? reduced to blocks that formed part of a distal pavement
flipper complex. Prominent dorsal fin present. Highly
Guizhouichthyosaurus tangae hydrodynamically streamlined, carangiform to thunniform
10 m (33 ft) TL, 4 tonnes swimmers.
FOSSIL REMAINS A few complete and partial skulls, HABITAT Coastal nearshore to deep oceans.
and skeletons severely flattened top to bottom. HABITS Swimming performance very good to extremely
ANATOMICAL CHARACTERISTICS Head elongated, high. Pursuit fishers and hunters of small to big game.
snout very long and slender, robust, medium-sized teeth NOTES The fastest fish- and cetacean-like sea reptile.
numerous and line most of jaws. Very high number of
vertebrae, trunk very shallow. Flippers elongated, narrow. MACGOWANIIDS
AGE Late Triassic; Carnian.
DISTRIBUTION AND FORMATIONS Southeastern China; MEDIUM-SIZED PARVIPELVIANS FROM
upper Falang. THE LATE TRIASSIC OF NORTH AMERICA
NOTES Placement in shastasaurids not certain. ANATOMICAL CHARACTERISTICS Snout rather short,
Shared its habitat with Anshunsaurus huangguoshuensis, fairly sharp tipped, teeth numerous, medium sized.
Miodentosaurus, Yunguisaurus, Qianichthyosaurus zhoui, Carangiform swimmers.
Guanlingsaurus, unnamed genus orientalis. HABITAT Coastal.
HABITS Swimming performance very good. Fishers of
small to medium-sized game.
Guizhouichthyosaurus lack of sufficient sampling.
tangae
139

Macgowania janiceps Hudsonelpidia brevirostris

3.5 m (11 ft) TL, 125 kg (275 lb) Adult size uncertain
FOSSIL REMAINS Skull and partial skeleton, badly FOSSIL REMAINS Skull and partial skeleton, badly
damaged. damaged, probably juvenile.
ANATOMICAL CHARACTERISTICS As for group. ANATOMICAL CHARACTERISTICS Standard for group.
AGE Late Triassic; middle Norian. AGE Late Triassic; middle Norian.
DISTRIBUTION AND FORMATIONS British Columbia; DISTRIBUTION AND FORMATIONS British Columbia;
upper Pardonet. upper Pardonet.
HABITAT Continental coastal. HABITAT Continental coastal.
NOTES Shared its habitat with unnamed genus NOTES Shared its habitat with unnamed genus
sikanniensis, Hudsonelpidia. sikanniensis, Macgowania.
HUDSONELPIDIDS HAUFFIOPTERYGIANS
SMALL PARVIPELVIANS FROM THE MEDIUM-SIZED PARVIPELVIANS FROM
LATE TRIASSIC OF NORTH AMERICA THE EARLY JURASSIC OF EUROPE
ANATOMICAL CHARACTERISTICS Snout rather short, ANATOMICAL CHARACTERISTICS Heads large, orbits
fairly sharp tipped. Flippers appear rather small. extremely large, teeth numerous and small. Transitional
Carangiform swimmers. carangiform-thunniform swimmers.
HABITAT Coastal. NOTES Absence from at least some other seas may reflect
HABITS Swimming performance very good. Fishers of lack of sufficient sampling.
small to medium-sized game. HABITAT Coastal.
NOTES Absence from at least some other seas may reflect HABITS Swimming performance very good. Fishers of
lack of sufficient sampling. small to medium-sized game.
Hauffiopteryx typicus
140

NOTES Absence from at least some other seas may reflect TEMNODONTOSAURIDS
lack of sufficient sampling.
MEDIUM-SIZED TO GIGANTIC PARVIPELVIANS
Hauffiopteryx typicus FROM THE EARLY JURASSIC OF EUROPE
2.9 m (9.5 ft) TL, 110 kg (250 lb)
FOSSIL REMAINS A number of complete skulls and ANATOMICAL CHARACTERISTICS Heads large,
skeletons, adult and juvenile. massively constructed, snouts very long, robust, orbits
ANATOMICAL CHARACTERISTICS Flippers medium very large, temporal region greatly expanded, teeth
sized, fore much larger than aft. numerous. Trunks long. Tails fairly long. Flippers medium
AGE Early Jurassic; late Pliensbachian to early Toarcian. sized, fore considerably larger than aft, pavement elements
DISTRIBUTION AND FORMATIONS Southern Germany, often poorly ossified. Carangiform swimmers.
Switzerland, southern England; Posidonienschiefer, HABITAT Island archipelago shallows.
unnamed. HABITS Swimming performance very good. Pursuit fishers
NOTES H. altera probably a juvenile of this species. and hunters of medium-sized and especially big game.
Shared its habitat with Meyerasaurus, Hauffiosaurus, NOTES Had the largest eyes among known sea reptiles
Seeleyosaurus, Hydrorion, unnamed genus trigonodon, and possibly all animals. Absence from at least some other
Suevoleviathan, Eurhinosaurus, Stenopterygius, seas probably reflects lack of sufficient sampling.
Mystriosaurus, Platysuchus, Macrospondylus, Hauffiopteryx
altera. Temnodontosaurus platyodon
8.5 m (28 ft) TL, 2.6 tonnes
FOSSIL REMAINS Skulls and skeletons, complete to
partial, some juvenile.
ANATOMICAL CHARACTERISTICS Head very large, very
massive including snout, teeth medium sized. Trunk fairly
Temnodontosaurus platyodon robust. Forefin somewhat larger than aft.
141

Temnodontosaurus platyodon
AGE Early Jurassic; late Hettangian and early Sinemurian. ANATOMICAL CHARACTERISTICS Head not massive,
DISTRIBUTION AND FORMATIONS Southern England; snout rather slender, teeth small. Trunk shallower.
upper Blue Lias. Forefin somewhat larger than aft.
HABITAT Island archipelago shallows. AGE Early Jurassic; early Toarcian.
NOTES T. risor may be the juvenile of this species. Shared DISTRIBUTION AND FORMATIONS Southern Germany,
its habitat with Excalibosaurus, Plesiosaurus, Eretmosaurus, northern, eastern France, central England?;
Leptonectes tenuirostris, Ichthyosaurus communis, unnamed Posidonienschiefer, unnamed, Whitby Mudstone?
genus eurycephalus. NOTES Placement in much earlier Temnodontosaurus is
problematic, may be more than one species. Eye nearly
Unnamed genus? trigonodon 0.3 m (1 ft) across. Newly found complete English
11.2 m (37 ft) TL, 4.5 tonnes specimen 10 m (32.8 ft) long assigned to this species is
FOSSIL REMAINS Skulls and skeletons, complete to not the largest. Shared its habitat with Meyerasaurus,
partial. Hauffiosaurus, Seeleyosaurus, Hydrorion, Hauffiopteryx,
Unnamed genus?
trigonodon
142

Unnamed genus?
trigonodon
Suevoleviathan, Eurhinosaurus, Stenopterygius, Suevoleviathan integer

Mystriosaurus, Platysuchus, Macrospondylus. 3.9 m (13 ft) TL, 220 kg (480 lb)
Unnamed genus? crassimanus ANATOMICAL CHARACTERISTICS Head massive,
9 m (30 ft) TL, 3 tonnes teeth medium sized. Foreflipper large.
FOSSIL REMAINS Majority of skull and skeleton. AGE Early Jurassic; early Toarcian.
ANATOMICAL CHARACTERISTICS Head very large. DISTRIBUTION AND FORMATIONS Southern Germany;
Trunk rather slender. Flippers rather small, fore much Posidonienschiefer.
larger than aft. NOTES S. disinteger is probably a growth stage of this
AGE Early Jurassic; middle Toarcian. species. Shared its habitat with Meyerasaurus, Hauffiosaurus,
DISTRIBUTION AND FORMATIONS Northeastern Seeleyosaurus, Hydrorion, Hauffiopteryx, unnamed genus
England; middle Whitby Mudstone. trigonodon, Eurhinosaurus, Stenopterygius, Mystriosaurus,
HABITAT Island archipelago shallows. Platysuchus, Macrospondylus.
NOTES Placement in much earlier Temnodontosaurus is
problematic, may not be same genus as trigonodon.
143

Suevoleviathan integer
LEPTONECTIDS DISTRIBUTION AND FORMATIONS Southern England;

upper Charmouth Mudstone, upper Belemnite Marls.
MEDIUM-SIZED TO GIGANTIC PARVIPELVIANS HABITAT Island archipelago shallows.
FROM THE EARLY JURASSIC OF EUROPE NOTES L. moorei may be juvenile of this species. Shared
its habitat with Ichthyosaurus anningae. May be the direct
ANATOMICAL CHARACTERISTICS Heads very long ancestor of L. tenuirostris.
because snout is very elongated into a slender bill, orbits
large, head not deep aft, teeth numerous and small. Leptonectes tenuirostris
Trunks fairly compact but not deep. Foreflippers markedly Adult size uncertain
larger than aft. Carangiform swimmers. FOSSIL REMAINS Several skulls and skeletons, complete
HABITAT Coastal to deep oceans. and partial.
HABITS Swimming performance high to very high. ANATOMICAL CHARACTERISTICS Head large, upper
Fishers of medium-sized game. and lower bills same length, teeth highly numerous and
NOTES Convergent with billfish, classic high-speed small. Tail rather short, lunate fluke very large. Foreflipper
carangiform swimmers. Absence from at least some other very large.
seas probably reflects lack of sufficient sampling. AGE Early Jurassic; early Sinemurian.
DISTRIBUTION AND FORMATIONS Southern England,
Leptonectes? solei Belgium; upper Blue Lias.
7 m (23 ft) TL, 1.1 tonnes HABITAT Island archipelago shallows.
FOSSIL REMAINS Majority of two skulls and skeletons, HABITS Highly maneuverable.
severely flattened top to bottom. NOTES Whether adults reached 5 m (16 ft) is not certain.
ANATOMICAL CHARACTERISTICS Teeth highly Shared its habitat with Excalibosaurus, Plesiosaurus,
numerous and small. Eretmosaurus, Temnodontosaurus, Ichthyosaurus
AGE Early Jurassic; early Pliensbachian. communis, unnamed genus eurycephalus.
144

Leptonectes tenuirostris
Leptonectes unnamed species Unnamed genus nuertingensis

Adult size uncertain 10 m (33 ft) TL, 3 tonnes
FOSSIL REMAINS Skull, probably juvenile. FOSSIL REMAINS Nearly complete skull and skeleton,
ANATOMICAL CHARACTERISTICS Insufficient severely flattened top to bottom, partial juvenile
information. remains.
AGE Early Jurassic; late Pliensbachian. ANATOMICAL CHARACTERISTICS Upper jaw slightly
DISTRIBUTION AND FORMATIONS Switzerland; unnamed. longer than lower.
HABITAT Island archipelago shallows. AGE Early Jurassic; middle Toarcian.
NOTES Placement in much earlier L. tenuirostris highly DISTRIBUTION AND FORMATIONS Eastern France;
problematic. unnamed.
Wahlisaurus massare NOTES Slender snout indicates does not belong to
Adult size uncertain Temnodontosaurus, azerguensis may be the juvenile of
FOSSIL REMAINS Skull and partial skeleton, possibly nuertingensis.
juvenile.
ANATOMICAL CHARACTERISTICS Insufficient Excalibosaurus costini
information. 6 m (20 ft) TL, 550 kg (1,200 lb)
AGE Early Jurassic; early Hettangian. FOSSIL REMAINS Two skulls and skeletons.
DISTRIBUTION AND FORMATIONS Central England; ANATOMICAL CHARACTERISTICS Upper bill elongated
middle Scunthorpe Mudstone. into a slender sword much longer than lower.
HABITAT Island archipelago shallows. AGE Early Jurassic; early Sinemurian.
145

Excalibosaurus costini
DISTRIBUTION AND FORMATIONS Southern England; AGE Early Jurassic; early Toarcian.
upper Blue Lias. DISTRIBUTION AND FORMATIONS Northeastern
HABITAT Island archipelago shallows. England, southern Germany, southeastern France,
NOTES Shared its habitat with Plesiosaurus, Eretmosaurus, Switzerland; lower Whitby Mudstone, Posidonienschiefer.
Leptonectes tenuirostris, Temnodontosaurus, Ichthyosaurus HABITAT Island archipelago shallows.
communis, unnamed genus eurycephalus. HABITS Oversized flippers indicate high maneuverability
combined with speed driven by oversized tail.
Eurhinosaurus longirostris NOTES Shared its habitat with Meyerasaurus, Hauffiosaurus,
3.4 m (11 ft) TL, 121 kg (260 lb) Rhomaleosaurus, Seeleyosaurus, Hydrorion, Hauffiopteryx,
FOSSIL REMAINS A number of skulls and skeletons of unnamed genus trigonodon, Suevoleviathan, Stenopterygius,
varying completeness. Plagiophthalmosuchus, Mystriosaurus, Platysuchus,
ANATOMICAL CHARACTERISTICS Upper bill elongated Macrospondylus. The sea reptile closest to modern
into a supersword many times longer than reduced lower swordfish in form and swimming performance.
bill, teeth very numerous and very small. Lunate fluke
very large. Foreflipper very long.
Eurhinosaurus longirostris
146

147

THUNNOSAURS ICHTHYOSAURIDS
SMALL TO LARGE PARVIPELVIANS FROM SMALL TO MEDIUM-SIZED THUNNOSAURS
THE EARLY JURASSIC TO THE END OF FROM THE EARLY JURASSIC OF EUROPE
ANATOMICAL CHARACTERISTICS Heads large, snouts
ANATOMICAL CHARACTERISTICS Uniform. Trunks fairly long, narrow, orbits large, teeth fairly large. Flippers
to very compact. Foreflippers with at least six long lines of broad, foreflipper medium sized and much larger than
pavement elements. Carangiform to thunniform swimmers. hind. High-speed thunniform swimmers.
HABITAT Coastal shallows to deep oceans. HABITS Swimming performance high to extremely high.
HABITS Swimming performance good to extremely high. Fishers of small game.
Fishers of small to medium-sized game. NOTES Absence from at least some other seas probably
NOTES The only marine reptiles to evolve thunniform reflects lack of sufficient sampling.
bodies.
Protoichthyosaurus prostaxalis
2.5 m (8 ft) TL, 100 kg (220 lb)
Stenopterygius
(Thunnosaurs) FOSSIL REMAINS Partial skulls and skeletons.
ANATOMICAL CHARACTERISTICS Snout very long,
fairly robust, orbit medium sized, lower jaw fairly robust,
upcurved, teeth numerous, stout, vertical.
AGE Early Jurassic; early Hettangian.
148

Stenopterygius
muscle study
Protoichthyosaurus?
applebyi
Protoichthyosaurus
prostaxalis
AGE Early Jurassic.

unknown.
DISTRIBUTION AND FORMATIONS Central England; HABITAT Island archipelago shallows.
lower Blue Lias, Hydraulic Limestone. HABITS Probably favored somewhat large prey.
HABITAT Island archipelago shallows. NOTES Whether this is in genus Protoichthyosaurus is
NOTES Shared its habitat with Eoplesiosaurus, problematic.
Stratesaurus, Avalonnectes, Eurycleidus, Atychodracon,
Thalassiodracon. Ichthyosaurus somersetensis
3.2 m (10 ft) TL, 220 kg (480 lb)
Protoichthyosaurus? applebyi FOSSIL REMAINS A few complete and partial skulls and
2 m (6.5 ft) TL, 50 kg (110 lb) skeletons.
FOSSIL REMAINS Two skulls and majority of a skeleton. ANATOMICAL CHARACTERISTICS Head fairly robust,
ANATOMICAL CHARACTERISTICS Snout moderately teeth moderate in number, medium sized, not too stout,
long, fairly shallow, orbits large, lower jaw shallow, somewhat procumbent.
straight, teeth numerous, stout, vertical. AGE Early Jurassic; Hettangian.
Ichthyosaurus somersetensis
149

Ichthyosaurus somersetensis
DISTRIBUTION AND FORMATIONS Southern England; Ichthyosaurus communis

lower and/or middle Blue Lias. 2 m (6.5 ft) TL, 50 kg (110 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS A number of complete and partial
skulls and skeletons, including probable fetuses.
Ichthyosaurus larkini ANATOMICAL CHARACTERISTICS Teeth numerous,
2 m (6.5 ft) TL, 50 kg (110 lb) rather small, vertical. Shoulder vertebrae spines tall.
FOSSIL REMAINS Three nearly complete skulls and AGE Early Jurassic; early Hettangian to early Sinemurian.
skeletons. DISTRIBUTION AND FORMATIONS Southern England;
ANATOMICAL CHARACTERISTICS Snout downcurved, middle to upper Blue Lias.
teeth moderate in number, fairly large, stout, vertical. HABITAT Island archipelago shallows.
AGE Early Jurassic; Hettangian. NOTES Assignment of some other species to this genus
DISTRIBUTION AND FORMATIONS Southern England; questionable. Presence of small fetus exiting skeleton of
lower and/or middle Blue Lias. an adult indicates live birth tailfirst. Shared its habitat with
HABITAT Island archipelago shallows. Excalibosaurus, Plesiosaurus, Eretmosaurus, Leptonectes
tenuirostris, unnamed genus eurycephalus.
Ichthyosaurus conybeari
Ichthyosaurus larkini 1.6 m (5.3 ft) TL, 30 kg (65 lb)
FOSSIL REMAINS Skulls and skeletons.
ANATOMICAL CHARACTERISTICS Teeth numerous,
rather small, vertical.
150

AGE Early Jurassic; late Hettangian to early Pliensbachian. DISTRIBUTION AND FORMATIONS Southern England;
DISTRIBUTION AND FORMATIONS Southern England; Blue Lias, level unknown.
lower to uppermost Blue Lias. HABITAT Island archipelago shallows.
HABITAT Island archipelago shallows. NOTES Placement in Ichthyosaurus dubious.
Unnamed genus eurycephalus

Ichthyosaurus 5 m (16 ft) TL, 1 tonne
conybeari
FOSSIL REMAINS Skull.
ANATOMICAL CHARACTERISTICS Head quite deep
along entire length, lower jaw fairly stout, teeth limited to
Ichthyosaurus anningae front halves of jaws, quite large and robust.
Adult size uncertain AGE Early Jurassic; early Sinemurian.
FOSSIL REMAINS Nearly complete skull and skeleton, DISTRIBUTION AND FORMATIONS Southern England;
two skulls and partial skeletons, all immature. upper Blue Lias.
ANATOMICAL CHARACTERISTICS Teeth moderate in NOTES Deep snout, large teeth, etc., indicate does not
number, fairly large, stout, vertical. belong in Temnodontosaurus, where is usually placed, may
AGE Early Jurassic; early Pliensbachian. be closer to ichthyosaurids. Shared its habitat with that,
DISTRIBUTION AND FORMATIONS Southern England; Excalibosaurus, Plesiosaurus, Eretmosaurus, Leptonectes
upper Charmouth Mudstone. tenuirostris, Ichthyosaurus communis.
HABITS Probably favored somewhat large prey.
NOTES Shared its habitat with Leptonectes solei.
Unnamed genus
eurycephalus
Ichthyosaurus
anningae
MALAWANIANS
Ichthyosaurus? breviceps SMALL THUNNOSAURS FROM THE EARLY
1.6 m (5.3 ft) TL, 30 kg (65 lb) CRETACEOUS OF THE MIDDLE EAST
FOSSIL REMAINS Some complete skulls and skeletons.
ANATOMICAL CHARACTERISTICS Head short, fairly Malawania anachronus
deep, snout not shallow, eyes large, teeth not numerous 2 m (6.5 ft) TL, 50 kg (110 lb)
and stout. Trunk vertebrae spines tall, body compact and FOSSIL REMAINS Partial skull and skeleton.
highly thunniform. Foreflipper large. ANATOMICAL CHARACTERISTICS Insufficient information.
AGE Early Jurassic. AGE Early Cretaceous; late Hauterivian or Barremian.
Ichthyosaurus? breviceps
151

Stenopterygius quadriscissus
DISTRIBUTION AND FORMATIONS Iraq; unnamed. Stenopterygius quadriscissus

HABITAT Continental shallows. 3.5 m (11.5 ft) TL, 270 kg (600 lb)
NOTES Placement in the much earlier ichthyosaurids is FOSSIL REMAINS A number of complete skulls and
problematic. skeletons of various ages, some with preserved soft tissues.
ANATOMICAL CHARACTERISTICS Snout not very
BARCROMIANS elongated, teeth functionally absent in adults. Body
plump and highly thunniform, lower tail lobe not strongly
SMALL TO LARGE THUNNOSAURS FROM elongated. Fins rather short.
THE EARLY JURASSIC TO THE END OF AGE Early Jurassic; early Toarcian.
THE MESOZOIC, GLOBAL DISTRIBUTION AND FORMATIONS Southern Germany;
Posidonienschiefer.
ANATOMICAL CHARACTERISTICS Uniform. Ventral HABITAT Island archipelago shallows.
pelvic elements fused to one another. NOTES Internally positioned fetuses in this genus indicate
HABITAT Coastal shallows to deep oceans. live birth. The profile-skeletal is based on a specimen with
HABITS Swimming performance good to extremely high. a well-preserved body outline; preserved pigmentation
Fishers of small to medium-sized game. patterns indicate countershading in at least some species
of genus. Later examples shared their habitat with less
STENOPTERYGIDS numerous S. triscissus and S. uniter, and Meyerasaurus,
Hauffiosaurus, Seeleyosaurus, Hydrorion, Hauffiopteryx,
MEDIUM-SIZED BARCROMIANS FROM THE unnamed genus trigonodon, Suevoleviathan,
EARLY TO MIDDLE JURASSIC OF EUROPE Eurhinosaurus, Mystriosaurus, Platysuchus,
AND SOUTH AMERICA Macrospondylus.
ANATOMICAL CHARACTERISTICS Very uniform. Heads Stenopterygius triscissus

large, snouts long, narrow, orbits large, teeth very reduced 3.3 m (11 ft) TL, 210 kg (450 lb)
to absent, especially in adults. Trunk width less than two- FOSSIL REMAINS A number of complete skulls and
thirds its own depth. Flippers broad, foreflippers medium skeletons of various ages.
sized and much larger than aft. High-speed thunniform ANATOMICAL CHARACTERISTICS Snout elongated,
swimmers. adult tooth reduction highly variable. Body not extremely
HABITS Swimming performance very to extremely high. plump, lower tail lobe not strongly elongated. Fins rather
Fishers of small game. short.
NOTES Highly convergent with lamnid sharks, tuna, and AGE Early Jurassic; early Toarcian.
dolphins, perhaps the acme of high-speed marine reptiles. DISTRIBUTION AND FORMATIONS Southern Germany,
Absence from at least some other seas probably reflects southern England; Posidonienschiefer, unnamed.
lack of sufficient sampling. HABITAT Island archipelago shallows.
152

Stenopterygius quadriscissus
153

Stenopterygius triscissus
NOTES Apparently did not appear as early as S. quadriscissus NOTES Apparently did not appear as early as S. triscissus
and may have been direct descendant of early examples of and may have been direct descendant of early examples of
that. that or of S. quadriscissus.
Stenopterygius uniter Stenopterygius aaleniensis

3.8 m (12.5 ft) TL, 310 kg (680 lb) 3 m (10 ft) TL, 150 kg (330 lb)
FOSSIL REMAINS A number of complete skulls and FOSSIL REMAINS Majority of skull and skeleton.
skeletons of various ages. ANATOMICAL CHARACTERISTICS Insufficient
ANATOMICAL CHARACTERISTICS Snout not very information.
elongated, adult tooth reduction variable. Body not AGE Middle Jurassic; early Aalenian.
extremely plump, lower tail lobe long. Fins fairly long, DISTRIBUTION AND FORMATIONS Southern Germany;
slender, and sharp tipped. lowest Opalinuston.
AGE Early Jurassic; early Toarcian. HABITAT Island archipelago shallows.
DISTRIBUTION AND FORMATIONS Southern Germany;
Posidonienschiefer. Chacaicosaurus cayi
HABITAT Island archipelago shallows. 3.5 m (11.5 ft) TL, 270 kg (600 lb)
HABITS Large fins indicate most maneuverable swimmer FOSSIL REMAINS Skull and minority of skeleton.
of this genus. ANATOMICAL CHARACTERISTICS Snout very long,
moderately robust, tapering.
Stenopterygius uniter
154


Chacaicosaurus
NOTES Shared its habitat with Pliosaurus? funkei,
cayi
Colymbosaurus? svalbardensis, Spitrasaurus, Palvennia,
Janusaurus, Cryopterygius.
AGE Middle Jurassic; early Bajocian. Palvennia hoybergeti

DISTRIBUTION AND FORMATIONS Southern Argentina; 4 m (13 ft) TL, 300 kg (650 lb)
upper Los Molles. FOSSIL REMAINS Complete skull, partial skull and
HABITAT Continental shelf. skeleton.
NOTES Shared its habitat with Maresaurus and ANATOMICAL CHARACTERISTICS Snout not elongated,
Mollesaurus. rather sharp tipped, teeth numerous and rather small.
AGE Late Jurassic; late Tithonian.
OPHTHALMOSAURIDS DISTRIBUTION AND FORMATIONS Svalbard; upper
Agardhfjellet.
SMALL TO LARGE BARCROMIANS FROM THE HABITAT Open continental shelf, polar.
MIDDLE JURASSIC TO LATE CRETACEOUS,
GLOBAL Janusaurus lundi
3 m (10 ft) TL, 135 kg (300 lb)
ANATOMICAL CHARACTERISTICS Foreflipper pavements FOSSIL REMAINS Partial skull and skeleton.
very highly developed. High-performance carangiform ANATOMICAL CHARACTERISTICS Teeth very small.
and thunniform swimmers. AGE Late Jurassic; late Tithonian.
HABITAT Coastal shallows to deep oceans. DISTRIBUTION AND FORMATIONS Svalbard; upper
HABITS Swimming performance good to extremely high. Agardhfjellet.
Fishers of small to medium-sized game. HABITAT Open continental shelf, polar.
OPHTHALMOSAURID Cryopterygius kristiansenae

5.3 m (17 ft) TL, 400 kg (880 lb)
MISCELLANEA
FOSSIL REMAINS Nearly complete skull and skeleton.
ANATOMICAL CHARACTERISTICS Head large, snout
Keilhauia nui quite long, teeth fairly numerous, rather small. Trunk
4 m (13 ft) TL, 300 kg (650 lb) fairly long. Flippers rather small. Transitional carangiform-
FOSSIL REMAINS Partial skull and majority of skeleton. thunniform swimmer.
ANATOMICAL CHARACTERISTICS Insufficient AGE Late Jurassic; late Tithonian.
information. DISTRIBUTION AND FORMATIONS Svalbard; upper
AGE Late Jurassic; late Tithonian. Agardhfjellet.
DISTRIBUTION AND FORMATIONS Svalbard; upper HABITAT Open continental shelf, polar.
Agardhfjellet. HABITS Swimming performance modest.
Cryopterygius kristiansenae
155

Cryopterygius kristiansenae
Arthropterygius chrisorum
2 m (6.5 ft) TL, 40 kg (90 lb)
NANNOPTERYGINES
FOSSIL REMAINS Skull and partial skeletons, adult to MEDIUM-SIZED OPHTHALMOSAURIDS FROM
juvenile. THE LATE JURASSIC AND POSSIBLY EARLY
ANATOMICAL CHARACTERISTICS Snout rather short, CRETACEOUS OF EUROPE
orbit large, teeth numerous.
AGE Late Jurassic; Oxfordian or Kimmeridgian. ANATOMICAL CHARACTERISTICS Heads fairly small,
DISTRIBUTION AND FORMATIONS Northern Northwest snouts rather short, narrow, orbits large, teeth small and
Territories; Ringnes. numerous. Trunk fairly long. Flippers quite small.
HABITAT Continental shelf. HABITS Not highly maneuverable. Fishers of small game.
NOTES Had the smallest flippers of any known ichthyosaur.
Unnamed genus or The purpose of such small steering and stabilizing surfaces
Arthropterygius thalassonotus is not clear. Fragmentary remains may indicate presence in
3 m (10 ft) TL, 140 kg (300 lb) Early Cretaceous. Absence from at least some other seas
FOSSIL REMAINS Partial skull and minority of skeleton. probably reflects lack of sufficient sampling.
information. Nannopterygius enthekiodon
AGE Late Jurassic; late Tithonian. 4.2 m (13.8 ft) TL, 375 (800 lb)
DISTRIBUTION AND FORMATIONS Western Argentina; FOSSIL REMAINS Majority of two skulls and skeleton,
middle Vaca Muerta. other skeletal remains.
HABITAT Continental shelf. ANATOMICAL CHARACTERISTICS Transitional
NOTES This species belonging to earlier and also poorly carangiform-thunniform.
preserved and remote Arthropterygius is problematic. AGE Late Jurassic; late Kimmeridgian or early Tithonian.
Shared its habitat with Dakosaurus andiniensis, unnamed DISTRIBUTION AND FORMATIONS Southern England;
genus and species, Sumpalla? middle Kimmeridge Clay.
Gengasaurus nicosiai HABITS Swimming performance modest.
3 m (10 ft) TL, 135 kg (300 lb) NOTES Is possible that some specimens do not belong to
FOSSIL REMAINS Skull disarticulated and majority of N. enthekiodon. Shared its habitat with Pliosaurus
skeleton. westburyensis, Grendelius mordax.
information. Thalassodraco etchesi
AGE Late Jurassic; late Kimmeridgian and/or earliest 3.5 m (10 ft) TL, 250 kg (500 lb)
Tithonian. FOSSIL REMAINS Majority of skull and partial skeleton.
DISTRIBUTION AND FORMATIONS Central Italy; ANATOMICAL CHARACTERISTICS May have been more
unnamed. thunniform.
HABITAT Island archipelago shallows. AGE Late Jurassic; early Tithonian.
NOTES Majority of skull is not visible.
156

Nannopterygius enthekiodon
DISTRIBUTION AND FORMATIONS Southern England; OPHTHALMOSAURINES

middle Kimmeridge Clay.
HABITAT Island archipelago shallows. MEDIUM-SIZED TO LARGE OPHTHALMOSAURIDS
FROM THE MIDDLE JURASSIC TO EARLY
Unnamed genus saveljeviensis CRETACEOUS OF EUROPE AND THE AMERICAS
5 m (16 ft) TL, 800 kg (1,800 lb)
FOSSIL REMAINS Majority of skull and partial skeleton, ANATOMICAL CHARACTERISTICS Heads large, snouts
partial skull and skeletons, other remains, some juvenile. long, narrow, orbits large, teeth reduced to absent,
ANATOMICAL CHARACTERISTICS May have been more especially in adults. Body cross section nearly circular.
thunniform. Flippers broad, foreflippers medium sized and much
AGE Late Jurassic; late middle to early late Tithonian. larger than aft. High-speed thunniform swimmers.
DISTRIBUTION AND FORMATIONS Southwestern HABITS Swimming performance very to extremely high.
Russia; unnamed. Fishers of small game.
HABITAT Continental shelf. NOTES Highly convergent with lamnid sharks, tuna, and
NOTES Nannopterygius yasykovi may be a juvenile of this dolphins. Absence from at least some other seas probably
species. Placement in earlier Nannopterygius highly reflects lack of sufficient sampling.
problematic.
Ophthalmosaurus icenicus
FOSSIL REMAINS Numerous incomplete specimens.
ANATOMICAL CHARACTERISTICS Teeth limited to front
Unnamed genus of jaws.
saveljeviensis
Ophthalmosaurus icenicus
157

AGE Middle Jurassic; middle Callovian. DISTRIBUTION AND FORMATIONS Eastern England;
DISTRIBUTION AND FORMATIONS Eastern England; Speeton Clay.
lower Oxford Clay. HABITAT Island archipelago shallows.
HABITAT Island archipelago shallows. Muiscasaurus catheti
NOTES Assignment of a number of specimens to this Adult size uncertain
species from other locations and times is highly FOSSIL REMAINS Skull and partial skeleton, partial skull
problematic. Whether adults reached 6 m (20 ft) is not and skeleton, both juvenile.
certain. Had the largest eyes, 0.25 m (0.6 ft) across, ANATOMICAL CHARACTERISTICS Teeth along entire
relative to its size of known sea reptiles. Shared its habitat length of jaws, rather small and numerous.
with Peloneustes, Pachycostasaurus, Simolestes, AGE Early Cretaceous; Barremian or Aptian.
Liopleurodon, Cryptoclidus, Muraenosaurus, Tricleidus, DISTRIBUTION AND FORMATIONS Colombia; Paja.
Tyrannoneustes, Suchodus, Gracilineustes. HABITAT Continental shelf.
NOTES May be a platypterygiine.
Mollesaurus periallus
5 m (16 ft) TL, 800 kg (1,800 lb)
PLATYPTERYGIINES
ANATOMICAL CHARACTERISTICS Insufficient SMALL TO LARGE OPHTHALMOSAURIDS
information. FROM THE LATE JURASSIC TO EARLY LATE
AGE Middle Jurassic; early Bajocian. CRETACEOUS, GLOBAL
DISTRIBUTION AND FORMATIONS Central Argentina;
upper Los Molles. ANATOMICAL CHARACTERISTICS Trunks not highly
HABITAT Continental shelf. compact. Foreflipper pavements uniquely developed.
NOTES Shared its habitat with Chacaicosaurus and High-performance carangiform swimmers.
Maresaurus. HABITAT Coastal shallows to deep oceans.
HABITS Swimming performance high to very high.
Baptanodon natans Fishers of small to medium-sized game.
6 m (20 ft) TL, 1 tonne NOTES Thunniforms not yet identified among these often
FOSSIL REMAINS Skull and partial skeletal remains. Cretaceous ichthyosaurs. Possess the most sophisticated
ANATOMICAL CHARACTERISTICS Teeth functionally forefins among marine tetrapods, if not vertebrates.
absent in adults. Includes the last ichthyosaurs.
AGE Late Jurassic; early Oxfordian.
DISTRIBUTION AND FORMATIONS Wyoming; upper Sumpalla argentina
Sundance. 2.1 m (7 ft) TL, 30 kg (60 lb)
HABITAT Interior seaway. FOSSIL REMAINS Skull and skeleton severely flattened
NOTES Shared its habitat with Pantosaurus, Tatenectes. top to bottom.
ANATOMICAL CHARACTERISTICS Head long, snout
fairly slender.
AGE Late Jurassic; middle or late Tithonian.
DISTRIBUTION AND FORMATIONS Western Argentina;
Baptanodon natans middle Vaca Muerta.
NOTES Shared its habitat with Arthropterygius?
thalassonotus, Dakosaurus andiniensis?, unnamed genus
and species?
Grendelius mordax
Acamptonectes densus 5.5 m (18 ft) TL, 800 kg (1,700 lb)
4.5 m (15 ft) TL, 400 kg (900 lb) FOSSIL REMAINS Skull, skeletal remains.
FOSSIL REMAINS Skull and partial skeleton, severely ANATOMICAL CHARACTERISTICS Snout very long,
flattened top to bottom and otherwise disrupted. robust, teeth fairly large, stout.
ANATOMICAL CHARACTERISTICS Insufficient AGE Late Jurassic; late Kimmeridgian and/or early
information. Tithonian.
AGE Early Cretaceous; Hauterivian.
158

Maiaspondylus lindoei
Grendelius mordax
FOSSIL REMAINS Partial skull and skeleton, other
remains including juvenile.
DISTRIBUTION AND FORMATIONS Southern England; information.
middle Kimmeridge Clay. AGE Early Cretaceous; early Albian.
HABITAT Island archipelago shallows. DISTRIBUTION AND FORMATIONS Southern Northwest
NOTES Some researchers consider this to be same as poorly Territories; upper Loon River.
preserved Brachypterygius extremus. Shared its habitat with HABITAT Continental shelf.
Pliosaurus westburyensis, Nannopterygius enthekiodon.
Unnamed genus cantabrigiensis
Grendelius? alekseevi Adult size uncertain
4 m (13 ft) TL, 300 kg (650 lb) FOSSIL REMAINS Minority of skull and skeletons,
FOSSIL REMAINS Partial skull and skeleton. possibly immature.
ANATOMICAL CHARACTERISTICS Insufficient ANATOMICAL CHARACTERISTICS Insufficient
information. information.
AGE Late Jurassic; early Tithonian. AGE Early Cretaceous; Cenomanian.
DISTRIBUTION AND FORMATIONS Western Russia; DISTRIBUTION AND FORMATIONS Eastern England;
unnamed. Greensand.
NOTES Placement in Grendelius not certain. HABITAT Island archipelago shallows.
NOTES Assignment of these poorly preserved remains to
Grendelius? zhuravlevi much earlier and remote Maiaspondylus, and of poorly
3.5 m (11 ft) TL, 200 kg (450 lb) preserved Russian remains to this species, is at best
FOSSIL REMAINS Partial skull and skeletons. extremely problematic.
information. Aegirosaurus leptospondylus
AGE Late Jurassic; early Tithonian. 1.6 m (5.2 ft) TL, 15 kg (30 lb)
DISTRIBUTION AND FORMATIONS Southwestern FOSSIL REMAINS A few skulls and skeletons, some with
Russia; unnamed. soft tissues.
NOTES Placement in Grendelius not certain. ANATOMICAL CHARACTERISTICS Head extremely long,
snout very long, fairly robust, teeth moderate in number,
Undorosaurus gorodischensis medium sized. Trunk fairly compact. Lunate fluke large.
4 m (13 ft) TL, 300 kg (650 lb) Foreflipper medium sized.
FOSSIL REMAINS Partial skull and skeleton. AGE Late Jurassic; early Tithonian.
ANATOMICAL CHARACTERISTICS Insufficient DISTRIBUTION AND FORMATIONS Southern Germany;
information. Altmühltal (Solnhofen).
AGE Late Jurassic; middle Tithonian. HABITAT Island archipelago shallows.
DISTRIBUTION AND FORMATIONS Southern Russia;
unnamed.
Aegirosaurus leptospondylus
159

Sveltonectes insolitus Unnamed genus hercynicus

3 m (10 ft) TL, 125 kg (275 lb) 7 m (23 ft) TL, 1.5 tonnes
FOSSIL REMAINS Skull and majority of skeleton. FOSSIL REMAINS Two partial skulls and partial skeleton.
ANATOMICAL CHARACTERISTICS Snout fairly robust, ANATOMICAL CHARACTERISTICS Insufficient
teeth fairly numerous, rather small, stout. information.
AGE Early Cretaceous; late Barremian. AGE Early Cretaceous; Aptian.
DISTRIBUTION AND FORMATIONS Western Russia; DISTRIBUTION AND FORMATIONS Northern Germany,
unnamed. northwestern France; unnamed.
HABITAT Continental shelf. HABITAT Island archipelago shallows.
NOTES Usually placed, as many mid-Cretaceous
ichthyosaurs have been, in Platypterygius, even though
the original specimens of P. platydactylus were
Sveltonectes insolitus
fragmentary and destroyed by Allied bombardment in
World War II.
Longirostra australis or longmani

Kyhytysuka sachicarum 6 m (20 ft) TL, 1 tonne
5.5 m (18 ft) TL, 800 kg (1,700 lb) FOSSIL REMAINS Skull and majority of skeleton, other
FOSSIL REMAINS Two skulls and partial of skeleton. remains.
ANATOMICAL CHARACTERISTICS Head large, snout ANATOMICAL CHARACTERISTICS Head large, snout
very long, fairly robust, teeth numerous, medium sized, very long, fairly robust, teeth numerous, medium sized,
range progressing aft from slender piercing to cutting to stout. Foreflipper very large, consists of a pavement of
stouter crushing. nine rows of elements in which radius and ulna are barely
AGE Early Cretaceous; latest Barremian and/or early larger than rest.
Aptian. AGE Early Cretaceous; middle to late Albian.
DISTRIBUTION AND FORMATIONS Colombia; upper DISTRIBUTION AND FORMATIONS Northeastern
Paja. Australia; Toolebuc, Allaru Mudstone.
HABITAT Continental shelf. HABITAT Shallow seaway, temperate.
NOTES Has incorrectly been placed in Platypterygius. HABITS Combined high speed and maneuverability.
Shared its habitat with Monquirasaurus, Callawayasaurus. NOTES Whether all specimens are one species is not
certain, correct species designation also uncertain. Usual
placement in Platypterygius not correct, so subgeneric
name applies. Remains of small enantiornithine coastal
Kyhytysuka sachicarum
bird have been found in gut region of one specimen.
Shared its habitat with ?Kronosaurus, Eromangasaurus,
Cratochelone, Notochelone, Bouliachelys.
Longirostra australis or longmani
160

Longirostra australis
or longmani
Caypullisaurus bonapartei Caypullisaurus

0.9 m (3 ft) TL, 4 kg (8 lb) bonapartei
FOSSIL REMAINS Two skulls and partial skeleton.
ANATOMICAL CHARACTERISTICS Snout not particularly
elongated and sharp tipped, toothless. Flippers not large. Unnamed genus and
AGE Early Cretaceous; early Tithonian.
species (Tithonian age)
lower Vaca Muerta.
NOTES Shared its habitat with Pliosaurus? patagonicus, Unnamed genus and
Cricosaurus araucanensis. species (Berriasian age)
Unnamed genus and species

1.2 m (4 ft) TL, 8 kg (18 lb)
FOSSIL REMAINS Two skulls and partial skeleton. Unnamed genus and species
ANATOMICAL CHARACTERISTICS Snout moderately 1.5 m (5 ft) TL, 15 kg (30 lb)
elongated and fairly robust with blunt tip, toothless. FOSSIL REMAINS Two skulls and partial skeleton.
AGE Late Jurassic; late Tithonian. ANATOMICAL CHARACTERISTICS Snout moderately
DISTRIBUTION AND FORMATIONS Western Argentina; elongated and more robust, with blunt tip, toothless.
middle Vaca Muerta. AGE Early Cretaceous; lower Berriasian.
HABITAT Continental shelf. DISTRIBUTION AND FORMATIONS Western Argentina;
NOTES Placement in Caypullisaurus is problematic, may upper Vaca Muerta.
be the same genus as the next entry. Shared its habitat HABITAT Continental shelf.
with Arthropterygius? thalassonotus, Dakosaurus NOTES As per previous entry. Shared its habitat with
andiniensis, Sumpalla? Cricosaurus puelchorum, Purranisaurus potens.
161

Simbirskiasaurus birjukovi
3.5 m (12 ft) TL, 200 kg (450 lb) S Q U A M AT E S
FOSSIL REMAINS Partial skull. (LIZARDS AND
ANATOMICAL CHARACTERISTICS
front and rear opening.
Nostrils divided into
SNAKES)
AGE Early Cretaceous; lower Barremian. SMALL TO GIGANTIC NEODIAPSIDS FROM
DISTRIBUTION AND FORMATIONS Western Russia; THE EARLY TRIASSIC TO THE MODERN
unnamed. ERA, GLOBAL
ANATOMICAL CHARACTERISTICS Diverse. Fully
Athabascasaurus bitumineus terrestrial to fully marine. Usually a flexible transverse
3.5 m (12 ft) TL, 200 kg (450 lb) joint at top of flat-roofed skull between snout and
FOSSIL REMAINS Partial skull and skeleton. temporal region, lower temporal bar absent, so skulls
ANATOMICAL CHARACTERISTICS Teeth moderate in usually strongly kinetic, coronoid process prominent.
number and rather small. Gastralia absent. Some limbless. Extremely variable. Two
AGE Early Cretaceous; earliest Albian. temporal openings in the skull, often lost by partial
DISTRIBUTION AND FORMATIONS Alberta; lower opening or by closure. Ribs usually swept backward.
Clearwater. HABITAT AND HABITS Highly variable, fully aquatic to
HABITAT Interior seaway, polar. fully terrestrial to aerial, herbivorous to archpredatory.
NOTES Shared its habitat with Nichollssaura, Lay soft-shelled eggs and give live birth, no parental care.
Wapuskanectes.
Pervushovisaurus bannovkensis LACERTILIANS

4 m (13 ft) TL, 300 kg (650 lb) (LIZARDS)
ANATOMICAL CHARACTERISTICS Snout very long, SMALL TO GIGANTIC SQUAMATES FROM THE
fairly robust, teeth numerous, rather small, robust, LATE TRIASSIC TO THE MODERN ERA, GLOBAL
intermeshing.
AGE Late Cretaceous; middle Cenomanian. ANATOMICAL CHARACTERISTICS Trunks not
DISTRIBUTION AND FORMATIONS Southwestern hyperelongated. All have limbs.
Russia; unnamed. HABITAT AND HABITS Very variable, fully aquatic to
HABITAT Continental shelf. fully terrestrial to aerial, herbivorous to archpredatory.
Tenuirostria americanus
3 m (10 ft) TL, 125 kg (275 lb)
TOXICOFERANS
FOSSIL REMAINS Skull, partial skeletal remains. SMALL TO GIGANTIC LACERTILIANS FROM
ANATOMICAL CHARACTERISTICS Head large, snout THE EARLY CRETACEOUS TO THE MODERN
very long, fairly robust, teeth numerous, medium sized, ERA, GLOBAL
stout.
AGE Late Cretaceous; middle Cenomanian. HABITAT Terrestrial to marine.
DISTRIBUTION AND FORMATIONS Wyoming; upper HABITS Small to big game hunters.
Mowry Shale. NOTES The existence of this very diverse group is not
HABITAT Interior seaway. certain.
NOTES Usual placement in much earlier Platypterygius
not correct, so subgeneric name applies. Last known MOSASAUROIDS (SEA LIZARDS)
ichthyosaur.
SMALL TO GIGANTIC TOXICOFERANS
FROM THE LATE CRETACEOUS, GLOBAL
Tenuirostria
americanus ANATOMICAL CHARACTERISTICS Fairly uniform.
Highly aquatic. Heads fairly to quite large, generally
subtriangular, neither very narrow nor very broad, snout
at least nearly half head length, quadrate modified into
162

S Q U A M A T E S ( L I Z A R D S A N D S N A K E S )
Tylosaurus muscle study
subcircular auditory depression, teeth modest in number, HABITAT Coastal and brackish shorelines, lagoons,
never very small, with bulbous roots, some on mouth reefs, estuaries.
roof. Aft tail flattened. Small bony scapula blade greatly HABITS Swimming performance good. Shallow-water
expanded by large cartilage fan, clavicles reduced or ambush and pursuit fishers of small to medium-sized
absent. Limbs modified at least into stiff-jointed partial game. Possibly bred and nested on beaches.
hydrofoils. Hydrodynamically streamlined, primarily axial NOTES Absence from at least some other seas may
undulators of anguilliform to transitional anguilliform- reflect lack of sufficient sampling.
carangiform grade, flippers primarily for stability and
maneuvering. Aigialosaurus dalmaticus
HABITAT Freshwaters to oceans, although dependence 1.15 m (3.8 ft) TL, 3 kg (6.5 lb)
on periodic access to freshwater may preclude dwelling in FOSSIL REMAINS Majority of two skulls and skeletons.
deep oceans in most or all cases. ANATOMICAL CHARACTERISTICS Teeth rather small
HABITS Swimming performance good to high. Ambush and widely spaced. Arm and leg similar in size.
and/or pursuit fishers and hunters of small to large game. AGE Late Cretaceous; late Cenomanian.
NOTES Relationships to other lizards and snakes uncertain. DISTRIBUTION AND FORMATIONS Croatia; unnamed.
AIGIALOSAURIDS NOTES Opetiosaurus bucchichi may be the same species.
May have shared its habitat with Pontosaurus lesinensis.
SMALL MOSASAUROIDS FROM THE LATE
CRETACEOUS OF NORTH AMERICA AND Komensaurus carrolli
EUROPE 1 m (3.3 ft) TL, 2 kg (4.5 lb)
FOSSIL REMAINS Minority of skull and majority of
ANATOMICAL CHARACTERISTICS Aft tail straight. skeleton.
Limbs short, modified into stiffened paddles, fingers and ANATOMICAL CHARACTERISTICS Insufficient
toes present but not highly elongated, presumably information.
webbed. Primarily axial undulators of anguilliform grade, AGE Late Cretaceous; late Cenomanian.
limbs primarily for stability and maneuvering, pressed DISTRIBUTION AND FORMATIONS Slovenia; upper Povir.
tight to body when hydrocruising. HABITAT Island archipelago shallows.
Aigialosaurus dalmaticus
163

Aigialosaurus dalmaticus
NOTES Shared its habitat with Adriosaurus suessi, A.? ANATOMICAL CHARACTERISTICS Fingers elongated
skrbinensis, A.? microbrachis, Acteosaurus, Eidolosaurus, into protoflipper.
Mesoleptos, Carsosaurus. AGE Late Cretaceous; Cenomanian or Turonian.
DISTRIBUTION AND FORMATIONS Croatia; unnamed.
Carsosaurus marchesetti HABITAT Island archipelago shallows.
1.5 m (5 ft) TL, 6 kg (13 lb) NOTES Phylogenetic position not certain.
FOSSIL REMAINS Majority of skeleton.
ANATOMICAL CHARACTERISTICS Insufficient MOSASAURIDS
information.
AGE Late Cretaceous; late Cenomanian. MEDIUM-SIZED TO GIGANTIC MOSASAUROIDS
DISTRIBUTION AND FORMATIONS Slovenia; upper Povir. FROM THE LATE CRETACEOUS, GLOBAL
ANATOMICAL CHARACTERISTICS Tongues probably
Vallecillosaurus donrobertoi stout. Ribcage deep only at chest, short-ribbed lumbar
1.3 m (4.3 ft) TL, 4.5 kg (10 lb)
FOSSIL REMAINS Partial skeleton.
information.
AGE Late Cretaceous; early Turonian.
DISTRIBUTION AND FORMATIONS Northern Mexico;
Agua Nueva.
NOTES Phylogenetic position not certain.
Portunatasaurus krambergeri
1 m (3.3 ft) TL, 2 kg (4.5 lb)
FOSSIL REMAINS Skull and majority of skeleton, severely Prognathodon
flattened top to bottom. (Mosasaurids)
164

region aft. Vertebrae at downward flexure of aft tail

supported an upper tail fin. Limbs clawless full flippers
with increased number of distal elements, outer finger and
toe divergent from others. Hydrodynamically well- Tethysaurus nopscai
streamlined, anguilliform to transitional anguilliform-
carangiform swimmers. Skin scales small, well ordered,
diamond shaped, subtly keeled fore and aft on top of
body, smooth on underside.
HABITAT Freshwaters to deep oceans.
HABITS Swimming performance very good to high.
Fishers and hunters of small to big game, including large Pannoniasaurus inexceptatus
shelled ammonoids. Probably gave live birth. 6 m (20 ft) TL, 550 kg (1,200 kg)
FOSSIL REMAINS Partial specimens.
Goronyosaurus nigeriensis ANATOMICAL CHARACTERISTICS Head somewhat
5 m (16 ft) TL, 300 kg (650 lb) shallow.
FOSSIL REMAINS Majority of skull and minority of AGE Late Cretaceous; Santonian.
skeleton. DISTRIBUTION AND FORMATIONS Hungary; upper
ANATOMICAL CHARACTERISTICS Snout long, robust, Csehbanya.
subrectangular, lower jaw same, teeth large. HABITAT Rivers and lakes.
AGE Late Cretaceous; middle Maastrichtian. HABITS Flattened head and river habitat suggest
DISTRIBUTION AND FORMATIONS Nigeria; Dukamaje. crocodile-like aquatic lifestyle, may have snatched land
HABITAT Continental coastal. animals when available.
NOTES Indicates that some mosasaurs inhabited
RUSSELLOSAURANS freshwaters.
MEDIUM-SIZED TO GIGANTIC MOSASAURIDS YAGUARASAURINES

FROM THE LATE CRETACEOUS, GLOBAL
MEDIUM-SIZED RUSSELLOSAURANS FROM
ANATOMICAL CHARACTERISTICS Anguilliform to THE LATE CRETACEOUS OF THE AMERICAS
transitional anguilliform-carangiform swimmers.
ANATOMICAL CHARACTERISTICS Tail vertebrae at
TETHYSAURINES modest downward flexure supported a modest upper tail
fin. Anguilliform swimmers.
MEDIUM-SIZED TO LARGE RUSSELLOSAURANS HABITAT Coastal.
FROM THE LATE CRETACEOUS OF EUROPE HABITS Swimming performance very good. Ambush and
AND AFRICA pursuit fishers and hunters of small to big game.
ANATOMICAL CHARACTERISTICS Tail vertebrae at Yaguarasaurus columbianus

modest downward flexure supported a modest upper tail 5 m (16.5 ft) TL, 300 kg (650 lb)
fin. Anguilliform swimmers. FOSSIL REMAINS Skull and minority of skeleton, severely
HABITAT Freshwaters to continental coastal. flattened top to bottom.
HABITS Swimming performance very good. Ambush and ANATOMICAL CHARACTERISTICS Standard for group.
pursuit fishers and hunters of small to big game. AGE Late Cretaceous; late Turonian.
DISTRIBUTION AND FORMATIONS Colombia; Hondita
Tethysaurus nopscai or La Frontera.
3 m (10 ft) TL, 70 kg (150 lb) HABITAT Continental coastal.
FOSSIL REMAINS Skull and partial skeleton.
ANATOMICAL CHARACTERISTICS Head fairly narrow, Russellosaurus coheni
snout somewhat delicate, teeth medium sized. 3 m (10 ft) TL, 70 kg (150 lb)
AGE Late Cretaceous; early Turonian. FOSSIL REMAINS Majority of skull.
DISTRIBUTION AND FORMATIONS Morocco; unnamed. ANATOMICAL CHARACTERISTICS Head somewhat
HABITAT Continental shallows. shallow, teeth large.
AGE Late Cretaceous; middle Turonian.
165

Tylosaurus nepaeolicus
8.5 m (28 ft) TL, 1.8 tonnes
Russellosaurus coheni FOSSIL REMAINS Two skulls, adult partial, minority of
skeleton, some soft tissues.
ANATOMICAL CHARACTERISTICS Snout shallow, teeth
medium sized, intermeshing. Coloration appears to have
DISTRIBUTION AND FORMATIONS Northeastern Texas; been dark overall.
lower Arcadia Park. AGE Late Cretaceous; late Coniacian to late Santonian?
HABITAT Interior seaway, coastal. DISTRIBUTION AND FORMATIONS Kansas; lower to
NOTES Shared its habitat with Dallasaurus. upper Niobrara?
HABITAT Interior seaway was shifting from maximum to
TYLOSAURINES less broad and deep.
NOTES T. kansasensis is probably a juvenile of this species.
LARGE TO GIGANTIC RUSSELLOSAURANS Some of the later specimens assigned to this species are
FROM THE LATE CRETACEOUS, GLOBAL probably unnamed species. May be the direct ancestor of
T. proriger.
ANATOMICAL CHARACTERISTICS Heads large,
somewhat shallow, snouts long, jaw tips somewhat squared Tylosaurus proriger
off and without front teeth. Fore and aft flippers similar in 13 m (43 ft) TL, 6.7 tonnes
size. Tail vertebrae at modest downward flexure supported FOSSIL REMAINS Several skulls and skeletons of varying
a modest upper tail fin. Anguilliform swimmers. completeness.
HABITAT Coastal. ANATOMICAL CHARACTERISTICS Snout robust, teeth
HABITS Swimming performance very good. Ambush and large.
pursuit fishers and hunters of small to big game. AGE Late Cretaceous; middle Santonian to early
Campanian.
DISTRIBUTION AND FORMATIONS Kansas, Manitoba;
Tylosaurus nepaeolicus upper Niobrara, lower Pierre Shale.
Tylosaurus proriger
166

Tylosaurus proriger and Dolichorhynchops bonneri
167

HABITAT Interior seaway was continuing to become less Tylosaurus or Hainosaurus bernardi
broad and deep. 12 m (40 ft) TL, 5 tonnes
NOTES Shared its habitat with Styxosaurus browni, FOSSIL REMAINS Majority of skull and minority of
Elasmosaurus, Dolichorhynchops bonneri, Latoplatecarpus, skeleton.
Globidens? dakotensis, Toxochelys latiremus. May be the ANATOMICAL CHARACTERISTICS Snout deep, teeth
direct ancestor of Tylosaurus pembinensis. rather small.
AGE Late Cretaceous; early Maastrichtian.
Tylosaurus pembinensis DISTRIBUTION AND FORMATIONS Belgium; Ciply
12 m (40 ft) TL, 5 tonnes Phosphatic Chalk.
FOSSIL REMAINS Majority of skull and partial skeleton. HABITAT Continental shelf.
ANATOMICAL CHARACTERISTICS Teeth medium sized, NOTES Shared its habitat with Prognathodon solvayi,
tightly intermeshing. Phosphorosaurus, Mosasaurus lemonnieri.
AGE Late Cretaceous; middle Campanian.
DISTRIBUTION AND FORMATIONS Manitoba; middle
Pierre Shale.
HABITAT Interior seaway was becoming much narrower. Tylosaurus or
NOTES Shared its habitat with Plioplatecarpus? primaevus. Hainosaurus bernardi
May be the direct ancestor of T. saskatchewanensis.
Taniwhasaurus oweni
7 m (23 ft) TL, 3 tonnes
Tylosaurus FOSSIL REMAINS Partial skull and skeleton.
pembinensis ANATOMICAL CHARACTERISTICS Insufficient
information.
AGE Late Cretaceous; earliest Maastrichtian.
DISTRIBUTION AND FORMATIONS Southern New
Zealand; lower Conway.
HABITAT Island coastal.
Taniwhasaurus (or Lakumasaurus)

antarcticus
Tylosaurus Adult size uncertain
saskatchewanensis
FOSSIL REMAINS Majority of skull and minority of
skeleton, possibly juvenile.
ANATOMICAL CHARACTERISTICS Teeth large.
AGE Late Cretaceous; late Campanian.
Tylosaurus saskatchewanensis DISTRIBUTION AND FORMATIONS Antarctic Peninsula;
9 m (30 ft) TL, 2 tonnes Santa Marta.
FOSSIL REMAINS Skull and partial skeleton. HABITAT Continental coastal, polar.
ANATOMICAL CHARACTERISTICS Teeth quite large.
AGE Late Cretaceous; late Campanian. Kaikaifilu hervei
DISTRIBUTION AND FORMATIONS Saskatchewan; upper 10 m (33 ft) TL, 3 tonnes
Bearpaw Shale. FOSSIL REMAINS Partial skull and minority of skeleton.
HABITAT Interior seaway was becoming very narrow. ANATOMICAL CHARACTERISTICS Insufficient
NOTES Shared its habitat with Nakonanectes, information.
Terminonatator, Plioplatecarpus? primaevus. May be the AGE Late Cretaceous; late Maastrichtian.
direct ancestor of T. bernardi. DISTRIBUTION AND FORMATIONS Antarctic Peninsula;
Lopez de Bertodano.
HABITAT Continental coastal, polar.
NOTES Shared its habitat with Morturneria.
168

NOTES Shared its habitat with Polycotylus latipinnis,

PLIOPLATECARPINES
Protostega.
MEDIUM-SIZED RUSSELLOSAURANS FROM
THE LATE CRETACEOUS OF NORTH AMERICA Gavialimimus almaghribensis
AND AFRICA 5 m (16 ft) TL, 350 kg (750 lb)
FOSSIL REMAINS Majority of skull, badly crushed.
ANATOMICAL CHARACTERISTICS Heads medium sized, ANATOMICAL CHARACTERISTICS Head elongated,
teeth medium sized. Tail vertebrae at strong downward snout long and narrow, teeth large, stout, widely spaced,
flexure supported a fairly large upper tail fin. Transitional not numerous.
anguilliform-carangiform swimmers. AGE Late Cretaceous; late Maastrichtian.
HABITAT Coastal to deep oceans. DISTRIBUTION AND FORMATIONS Morocco; upper
HABITS Swimming performance very good to high. Couche III.
Ambush and especially pursuit fishers and hunters of small HABITAT Continental shelf.
to big game.
NOTES Absence from at least some other seas probably Ectenosaurus clidastoides
reflects lack of sufficient sampling. This group appears to 6 m (20 ft) TL, 600 kg (1,300 lb)
have evolved high-speed swimming independently from FOSSIL REMAINS Complete skull with partial skeleton,
Mosasaurinians. severely flattened, skin, internal fibers.
ANATOMICAL CHARACTERISTICS Head large, fairly
Angolasaurus bocagei narrow, snout long, teeth medium sized. Scales small, 2 ×
4 m (13 ft) TL, 200 kg (450 lb) 3 mm.
FOSSIL REMAINS Partial skull. AGE Late Cretaceous; early Santonian.
ANATOMICAL CHARACTERISTICS Insufficient DISTRIBUTION AND FORMATIONS Kansas, Texas; lower
information. Niobrara.
AGE Late Cretaceous; middle or late Turonian. HABITAT Interior seaway was at maximum breadth and
DISTRIBUTION AND FORMATIONS Angola; Tadi. depth.
HABITAT Continental shelf. NOTES Original specimen in a German museum lost
during World War II. Shared its habitat with Selmasaurus?
Selmasaurus? johnsoni johnsoni, Plesioplatecarpus, Chelosphargis.
FOSSIL REMAINS Partial skull and skeleton, possibly Plesioplatecarpus planifrons
immature. 5 m (16 ft) TL, 350 kg (750 lb)
ANATOMICAL CHARACTERISTICS Teeth large, long FOSSIL REMAINS A number of skulls and skeletons of
spikes. varying completeness, often severely flattened top to
AGE Late Cretaceous; early Santonian. bottom.
DISTRIBUTION AND FORMATIONS Kansas; lower ANATOMICAL CHARACTERISTICS Snout rather short,
Niobrara. head broad.
HABITAT Interior seaway was at maximum breadth and AGE Late Cretaceous; middle Coniacian to early
depth. Santonian.
NOTES Placement in the much later genus Selmasaurus is DISTRIBUTION AND FORMATIONS Kansas, Alabama;
uncertain. Shared its habitat with Ectenosaurus, lower Niobrara, upper Eutaw.
Plesioplatecarpus, Chelosphargis. HABITAT Interior seaway was at maximum breadth and
depth, continental shelf.
Selmasaurus russelli NOTES Shared its habitat with Selmasaurus? johnsoni,
Adult size uncertain Ectenosaurus, Chelosphargis.
FOSSIL REMAINS Minority of skull and skeleton, possibly
immature. Latoplatecarpus nichollsae
ANATOMICAL CHARACTERISTICS Skull appears to have Adult size uncertain
been akinetic. FOSSIL REMAINS Two skulls complete and partial,
AGE Late Cretaceous; early Campanian. severely crushed, two partial skeletons, possibly immature.
DISTRIBUTION AND FORMATIONS Alabama; lower ANATOMICAL CHARACTERISTICS Quadrate and
Mooreville Chalk. tympanic circle large, teeth large.
HABITAT Continental shelf. AGE Late Cretaceous; early Campanian.
169

AGE Late Cretaceous; middle Santonian to earliest

Latoplatecarpus Campanian.
nichollsae DISTRIBUTION AND FORMATIONS Kansas; middle to
uppermost Niobrara.
HABITAT Interior seaway was shifting from maximum to
less broad and deep.
DISTRIBUTION AND FORMATIONS Manitoba; lower NOTES Shared its habitat with Styxosaurus snowii,
Pierre Shale. Polycotylus latipinnis, Dolichorhynchops osborni, Eonatator
HABITAT Interior seaway was continuing to become less sternbergii, Clidastes propython, Ctenochelys stenoporus,
broad and deep. Protostega gigas. May be the direct ancestor of
NOTES Probably includes L. willistoni. Shared its habitat Plioplatecarpus? primaevus.
with Styxosaurus browni, Elasmosaurus, Dolichorhynchops
bonneri, Tylosaurus proriger, Globidens? dakotensis, Plioplatecarpus? primaevus
Toxochelys latiremus. 3.9 m (12.5 ft) TL, 170 kg (375 lb)
FOSSIL REMAINS A number of partial skulls and
Platecarpus tympanicus skeletons, mostly immature.
5.4 m (17.5 ft) TL, 440 kg (1,000 lb) ANATOMICAL CHARACTERISTICS Head short, orbits
FOSSIL REMAINS Skull and numerous other remains. large. Foreflipper much larger than aft.
ANATOMICAL CHARACTERISTICS Snout not elongated, AGE Late Cretaceous; early to late Campanian or to early
teeth medium sized. Fore and aft flippers similar in size. Maastrichtian.
Scales fairly large, 3.5 × 4.5 mm.
Platecarpus tympanicus
Plioplatecarpus? primaevus
170

DISTRIBUTION AND FORMATIONS Dakotas, Kansas, HABITS Swimming performance very good. Ambush and
Saskatchewan; lower to middle Pierre Shale, upper pursuit fishers and hunters of small to big game.
Bearpaw Shale.
HABITAT Interior seaway was becoming very narrow. Halisaurus platyspondylus
HABITS Deep water if adult orbits large. 4 m (13 ft) TL, 200 kg (450 lb)
NOTES May include P. peckensis. Shared its habitat with FOSSIL REMAINS Partial skulls and skeleton.
Nakonanectes, Terminonatator, Tylosaurus pembinensis, T. ANATOMICAL CHARACTERISTICS Insufficient
saskatchewanensis. information.
AGE Late Cretaceous; middle to late Maastrichtian.
Plioplatecarpus? houzeaui DISTRIBUTION AND FORMATIONS New Jersey,
Adult size uncertain Maryland; New Egypt, Severn.
FOSSIL REMAINS Crushed skull and partial skeleton, HABITAT Continental coastal.
probably immature, possibly other remains.
ANATOMICAL CHARACTERISTICS Teeth fairly large. Halisaurus arambourgi
AGE Late Cretaceous; middle Maastrichtian. 5 m (16 ft) TL, 350 kg (750 lb)
DISTRIBUTION AND FORMATIONS Belgium; Brown FOSSIL REMAINS Two partial skulls and minority of
Phosphate Chalk. skeleton.
HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Snout not long, teeth
HABITS Deep water. widely spaced, medium sized.
NOTES May be the direct ancestor of P. marshi. AGE Late Cretaceous; late Maastrichtian.
DISTRIBUTION AND FORMATIONS Morocco; unnamed.
Plioplatecarpus marshi HABITAT Continental coastal.
5 m (16 ft) TL, 350 kg (750 lb)
FOSSIL REMAINS Partial skull and several partial
skeletons.
ANATOMICAL CHARACTERISTICS Insufficient Halisaurus
information. arambourgi
DISTRIBUTION AND FORMATIONS Belgium; upper
Gulpen.
HABITS Deep water.
NOTES May be the direct descendant of P. houzeaui. Eonatator sternbergii
Whether any other known species belong to this genus is 3.5 m (11 ft) TL, 100 kg (220 lb)
at best uncertain. FOSSIL REMAINS Partial skull and majority and minority
of two skeletons.
MOSASAURANS ANATOMICAL CHARACTERISTICS Head fairly large,
fairly narrow, snout elongated. Flippers medium and
SMALL TO GIGANTIC MOSASAURIDS FROM similar in size.
THE LATE CRETACEOUS, GLOBAL AGE Late Cretaceous; earliest Campanian.
DISTRIBUTION AND FORMATIONS Kansas; uppermost
ANATOMICAL CHARACTERISTICS Anguilliform to Niobrara.
transitional anguilliform-carangiform swimmers. HABITAT Interior seaway was continuing to become less
broad and deep.
HALISAURINES NOTES Shared its habitat with Styxosaurus snowii,
Polycotylus latipinnis, Dolichorhynchops osborni, Platecarpus
SMALL TO MEDIUM-SIZED MOSASAURANS tympanicus, Clidastes propython, Ctenochelys stenoporus,
FROM THE LATE CRETACEOUS, GLOBAL Protostega gigas.
ANATOMICAL CHARACTERISTICS Heads not very large. Eonatator? coellensis

Tail vertebrae at modest downward flexure supported a 2.8 m (9 ft) TL, 50 kg (110 lb)
modest upper tail fin. Anguilliform swimmers. FOSSIL REMAINS Skull and majority of skeleton with soft
HABITAT Continental coastal. tissues.
171

ANATOMICAL CHARACTERISTICS Head fairly large, MOSASAURINE MISCELLANEA

fairly narrow, snout elongated. Flippers medium and
similar in size.
AGE Late Cretaceous; Campanian. Dallasaurus turneri
DISTRIBUTION AND FORMATIONS Colombia; upper 1 m (3.3 ft) TL, 2 kg (4.5 lb)
Lydite. FOSSIL REMAINS Partial skull and two partial skeletons.
HABITAT Continental coastal. ANATOMICAL CHARACTERISTICS Flippers not fully
developed.
Phosphorosaurus ponpetelegans AGE Late Cretaceous; middle Turonian.
Adult size not certain DISTRIBUTION AND FORMATIONS Texas; lower Arcadia
FOSSIL REMAINS Majority of skull and minority of Park Shale.
skeleton, possibly immature. HABITAT Interior seaway.
ANATOMICAL CHARACTERISTICS Orbits face somewhat NOTES May be a more basal mosasauroid. Shared its
forward and may have had overlapping fields of vision. habitat with Russellosaurus.
AGE Late Cretaceous; earliest Maastrichtian.
DISTRIBUTION AND FORMATIONS Japan; Hakobuchi. CLIDASTINIANS
SMALL TO GIGANTIC MOSASAURANS
Phosphorosaurus ortliebi FROM THE LATE CRETACEOUS, GLOBAL
Adult size not certain
FOSSIL REMAINS Partial skull, possibly immature. ANATOMICAL CHARACTERISTICS Teeth robust. Trunk
ANATOMICAL CHARACTERISTICS Insufficient elongated. Tall tail vertebrae at modest downward flexure
information. supported a modest upper tail fin, tail including aft distal
AGE Late Cretaceous; early Maastrichtian. downward-flexed segment rather short. Anguilliform
DISTRIBUTION AND FORMATIONS Belgium; Ciply swimmers.
Phosphatic Chalk. HABITAT Coastal.
HABITAT Island archipelago shallows. HABITS Swimming performance very good. Ambush and
NOTES Shared its habitat with Tylosaurus or Hainosaurus pursuit fishers and hunters of small to big game, some
bernardi, Prognathodon solvayi, Mosasaurus lemonnieri. crushers.
Clidastes liodontus
MOSASAURINES 3 m (10 ft) TL, 55 kg (120 lb)
SMALL TO GIGANTIC MOSASAURANS FROM FOSSIL REMAINS Several specimens.
THE LATE CRETACEOUS, GLOBAL ANATOMICAL CHARACTERISTICS Head medium sized,
fairly narrow, teeth medium sized, stout. Flippers small,
ANATOMICAL CHARACTERISTICS Tail vertebrae at fore larger than aft.
modest downward flexure supported a modest upper tail AGE Late Cretaceous; late Coniacian to late Santonian.
fin. Anguilliform swimmers. DISTRIBUTION AND FORMATIONS Kansas, Texas; lower
HABITAT Coastal. to middle Niobrara.
HABITS Swimming performance very good. Ambush and HABITAT Interior seaway was shifting from maximum to
especially pursuit fishers and hunters of small to big game. less broad and deep.
NOTES Assigned time span probably too long for a single
species. May be direct ancestor of C. propython.
Clidastes liodontus
172

Clidastes liodontus
Clidastes propython Prognathodon? overtoni

3 m (10 ft) TL, 55 kg (120 lb) 8 m (25 ft) TL, 1 tonne
FOSSIL REMAINS Several specimens. FOSSIL REMAINS Skulls and skeletal remains.
ANATOMICAL CHARACTERISTICS Head shallow, teeth ANATOMICAL CHARACTERISTICS Head large, fairly
rather small, stout. broad, heavily constructed, snout deep, teeth large, stout
AGE Late Cretaceous; middle Santonian to earliest and blunted. Flippers medium and similar in size.
Campanian. AGE Late Cretaceous; late Campanian.
DISTRIBUTION AND FORMATIONS Kansas, South DISTRIBUTION AND FORMATIONS South Dakota,
Dakota, Colorado; middle to uppermost Niobrara. southern Alberta; upper Pierre Shale, lower Bearpaw Shale.
HABITAT Interior seaway was shifting from maximum to HABITAT Interior seaway was becoming much narrower.
less broad and deep. HABITS Specialized for crushing hard-shelled prey.
NOTES Shared its habitat with Styxosaurus snowii, NOTES Shared its habitat with Albertonectes,
Polycotylus latipinnis, Dolichorhynchops osborni, Platecarpus Dolichorhynchops herschelensis, Mosasaurus? missouriensis,
tympanicus, Eonatator sternbergii, Ctenochelys stenoporus, Archelon.
Protostega gigas.
Prognathodon? currii
FOSSIL REMAINS Skull and minority of skeleton.
ANATOMICAL CHARACTERISTICS Head large, fairly
Clidastes broad, heavily constructed, snout deep, teeth large, stout,
propython and blunted.
DISTRIBUTION AND FORMATIONS Israel; upper
Mishash.
Prognathodon? overtoni
173

Unnamed genus and species

5 m (16 ft) TL, 250 kg (550 lb)
FOSSIL REMAINS Majority of skull.
Prognathodon? ANATOMICAL CHARACTERISTICS Head fairly heavily
currii constructed, snout shallow, teeth medium sized, stout.
AGE Late Cretaceous; Campanian or Maastrichtian.
DISTRIBUTION AND FORMATIONS Northern New
Zealand; Maungataniwha Sandstone.
NOTES Placement within Prognathodon overtoni incorrect.
HABITAT Continental shelf. Globidens alabamaensis

HABITS Specialized for crushing hard-shelled prey. 4.5 m (15 ft) TL, 200 kg (450 lb)
FOSSIL REMAINS Minority of skull and skeleton.
Prognathodon solvayi ANATOMICAL CHARACTERISTICS Skull heavily
Adult size uncertain constructed, teeth short, blunt, aft set fairly large knobs.
FOSSIL REMAINS Adult partial skull, immature skull and AGE Late Cretaceous; middle Campanian.
partial skeleton. DISTRIBUTION AND FORMATIONS Alabama; upper
ANATOMICAL CHARACTERISTICS Teeth large. Mooreville Chalk.
AGE Late Cretaceous; early Maastrichtian. HABITAT Continental shelf.
DISTRIBUTION AND FORMATIONS Belgium; Ciply HABITS Highly specialized for crushing hard-shelled prey.
Phosphatic Chalk. NOTES Shared its habitat with Toxochelys moorevillensis,
HABITAT Island archipelago shallows. Corsochelys, Ctenochelys acris, Prionochelys.
HABITS Specialized for crushing hard-shelled prey.
NOTES P. giganteus may be the adult of this species. The Globidens? dakotensis
assignment of at least some other species to this genus is 6 m (20 ft) TL, 450 kg (1,000 lb)
uncertain. Shared its habitat with Tylosaurus or FOSSIL REMAINS Majority of skull.
Hainosaurus bernardi, Phosphorosaurus, Mosasaurus ANATOMICAL CHARACTERISTICS Head fairly broad,
lemonnieri. heavily constructed, especially lower jaw, snout rather
short, teeth short, blunt, aft set fairly large knobs.
Tenerasaurus (or Prognathodon?) hashimi AGE Late Cretaceous; early Campanian.
Adult size uncertain DISTRIBUTION AND FORMATIONS South Dakota; lower
FOSSIL REMAINS Majority of skeleton, with soft tissues, Pierre Shale.
probably immature. HABITAT Interior seaway was continuing to become less
ANATOMICAL CHARACTERISTICS Insufficient broad and deep.
information. HABITS Highly specialized for crushing hard-shelled prey.
AGE Late Cretaceous; late Maastrichtian. NOTES Whether this is the same genus as incomplete G.
DISTRIBUTION AND FORMATIONS Jordan; upper alabamaensis is not certain. Additional specimens from
Muwaqqar Chalk. other locations may or may not represent the species.
HABITAT Continental shelf. Shared its habitat with Styxosaurus browni, Elasmosaurus,
NOTES Whether this is a distinct genus or within Dolichorhynchops bonneri, Tylosaurus proriger,
Prognathodon is not certain. Latoplatecarpus, Toxochelys latiremus.
Unnamed genus and

Globidens?
species (Maungataniwha
dakotensis
Sandstone)
174

Globidens? simplex Eremiasaurus heterodontus

5.5 m (18 ft) TL, 350 kg (750 lb) 5 m (16 ft) TL, 300 kg (650 lb)
FOSSIL REMAINS Partial skull, other remains. FOSSIL REMAINS Majority of two skulls and skeletons,
ANATOMICAL CHARACTERISTICS Skull heavily badly damaged.
constructed, teeth short, blunt knobs, especially aft. ANATOMICAL CHARACTERISTICS Head fairly broad,
AGE Late Cretaceous; late Maastrichtian. snout fairly deep, teeth large, increasingly blunted
DISTRIBUTION AND FORMATIONS Angola; unnamed. progressing aft.
HABITAT Continental shelf. AGE Late Cretaceous; late Maastrichtian.
HABITS Highly specialized for crushing hard-shelled prey. DISTRIBUTION AND FORMATIONS Morocco; unnamed.
NOTES That this species belongs to the same genus as the HABITS Continental shelf.
much earlier Globidens is problematic.
Plesiotylosaurus crassidens
6.5 m (21 ft) TL, 700 kg (1,500 lb)
MOSASAURINIANS
FOSSIL REMAINS Skull remains, badly damaged.
LARGE TO GIGANTIC MOSASAURANS ANATOMICAL CHARACTERISTICS Skull heavily
FROM THE LATE CRETACEOUS, GLOBAL constructed, teeth small.
AGE Late Cretaceous; middle Maastrichtian.
ANATOMICAL CHARACTERISTICS Head medium sized, DISTRIBUTION AND FORMATIONS Central California;
teeth sharp. Tall tail vertebrae at strong downward flexure lower Moreno.
supported a fairly large upper tail fin. Transitional HABITAT Continental shelf.
anguilliform-carangiform swimmers. NOTES Shared its habitat with Plotosaurus.
HABITAT Coastal to deep oceans.
HABITS Swimming performance very good to high. Mosasaurus? missouriensis
Pursuit fishers and hunters of small to big game. 6.5 m (21 ft) TL, 700 kg (1,500 lb)
NOTES This group appears to have evolved high-speed FOSSIL REMAINS Skull and partial skeleton, other remains.
swimming independently from plioplatecarpines. ANATOMICAL CHARACTERISTICS Head somewhat
deep, teeth medium sized.
Moanasaurus mangahouangae AGE Late Cretaceous; late Campanian.
12 m (40 ft) TL, 4 tonnes DISTRIBUTION AND FORMATIONS South Dakota,
FOSSIL REMAINS Majority of skull, minority of skull and Montana, Alberta; upper Pierre Shale, middle Bearpaw
skeleton. Shale.
ANATOMICAL CHARACTERISTICS Teeth medium sized. HABITAT Interior seaway was becoming much narrower.
AGE Late Cretaceous; Campanian and/or Maastrichtian. NOTES Shared its habitat with Prognathodon? overtoni,
DISTRIBUTION AND FORMATIONS Northern New Nichollsemys, Archelon.
Zealand; unnamed.
HABITAT Island coastal and probably deep ocean.
NOTES Probably includes Rikisaurus tehoensis.
Mosasaurus?
missouriensis
Moanasaurus
mangahouangae or
Rikisaurus tehoensis
Mosasaurus? conodon
7 m (23 ft) TL, 900 kg (2,000 lb)
FOSSIL REMAINS Skull and other partial specimens.
Eremiasaurus broad, shallow, teeth fairly large.
heterodontus AGE Late Cretaceous; late Campanian and/or early
Maastrichtian.
DISTRIBUTION AND FORMATIONS New Jersey,
Maryland? Alabama? Arkansas? Montana? Colorado?
South Dakota?; Navesink, Severn? Demopolis Chalk?
175

Marlbrook Chalk? upper Pierre Shale? lower Bearpaw

Shale? Mosasaurus
NOTES Whether this species belongs in Mosasaurus is hoffmanni
uncertain. It is possible that the species is limited to the
original poorly preserved specimen from New Jersey.
Shared that habitat with Cimoliasaurus.
DISTRIBUTION AND FORMATIONS Netherlands, etc.?;
Maastricht, etc.?
NOTES A classic mosasaur. Assignment of many specimens
Mosasaurus?
from a number of continents to this species and to the
conodon
genus is uncertain, those not from the late Maastrichtian
being especially problematic. May include a Russian lower
jaw that may be the largest known specimen of the taxon.
Mosasaurus? beaugei
8.5 m (28 ft) TL, 1.5 tonnes
Mosasaurus FOSSIL REMAINS Two partial skulls, other remains.

lemonnieri ANATOMICAL CHARACTERISTICS Insufficient
information.
DISTRIBUTION AND FORMATIONS Morocco; unnamed.
Mosasaurus lemonnieri NOTES Whether this species belongs in Mosasaurus is
12 m (40 ft) TL, 4.5 tonnes uncertain.
FOSSIL REMAINS Skulls and skeletal remains, juvenile
to adult. Plotosaurus bennisoni
ANATOMICAL CHARACTERISTICS Head not deep, 13 m (43 ft) TL, 5.9 tonnes
teeth rather small. FOSSIL REMAINS Skull and skeletal remains, adult and
AGE Late Cretaceous; early Maastrichtian. immature.
DISTRIBUTION AND FORMATIONS Belgium; Ciply ANATOMICAL CHARACTERISTICS Head somewhat
Phosphatic Chalk. shallow, moderately broad, teeth fairly numerous,
HABITAT Island archipelago shallows. medium sized. Chest fairly deep. Fins fairly long, narrow,
NOTES Shared its habitat with Tylosaurus or Hainosaurus similar in size fore and aft.
bernardi, Prognathodon solvayi, Phosphorosaurus. May be AGE Late Cretaceous; middle Maastrichtian.
the direct ancestor of M. hoffmanni. DISTRIBUTION AND FORMATIONS Central California;
lower Moreno.
Mosasaurus hoffmanni HABITAT Continental shelf and possibly deep ocean.
13 m (43 ft) TL, 5.5 tonnes HABITS High-speed pursuit predator.
FOSSIL REMAINS Skulls and skeletal remains. NOTES P. tuckeri may be adult of this species. Shared its
ANATOMICAL CHARACTERISTICS Head not deep, habitat with Plesiotylosaurus. The fastest-swimming known
teeth large. mosasaur.
Plotosaurus
bennisoni
176

Plotosaurus
bennisoni
OPHIDIOMORPHS NOTES Absence from at least some other seas probably

reflects lack of sufficient sampling. May be the lizards
SMALL TOXICOFERANS FROM THE LATE closest to snakes, which may have had an aquatic origin;
CRETACEOUS TO THE MODERN ERA, GLOBAL this is uncertain because snakes may have evolved from
burrowing lizards.
ANATOMICAL CHARACTERISTICS Fairly diverse. Heads
neither large nor heavily constructed. Vertebral series Dolichosaurus longicollis
elongated and made extraflexible via a high number of 1.5 m (5 ft) TL, 3.5 kg (8 lb)
vertebrae (140 or more) and extra articulations between FOSSIL REMAINS Several partial skulls and skeletons.
them. Aft tails straight. Limbs at least somewhat reduced, ANATOMICAL CHARACTERISTICS Head moderate in
not full flippers. Hydrodynamically streamlined, size, neck elongated.
anguilliform swimmers. AGE Late Cretaceous; early Cenomanian.
HABITAT Coastal and brackish shorelines, lagoons, reefs, DISTRIBUTION AND FORMATIONS Southeastern
estuaries, probably accessed freshwater frequently, England; Chalk Marl.
including rainwater atop salt water. HABITAT Island nearshore.
HABITS Swimming performance good. Shallow-water NOTES May have shared its habitat with Coniasaurus.
ambush and pursuit fishers of small to medium-sized
game. Possibly bred and nested on beaches. Coniasaurus cressidus
0.5 m (1.6 ft) TL, 0.15 kg (0.35 lb)
DOLICHOSAURIDS FOSSIL REMAINS A few partial specimens.

ANATOMICAL CHARACTERISTICS Teeth large and
SMALL OPHIDIOMORPHS FROM THE LATE robust, blunt tipped, especially aft.
CRETACEOUS OF EUROPE AND THE MIDDLE AGE Late Cretaceous; early or middle Cenomanian.
EAST DISTRIBUTION AND FORMATIONS Southeastern
England; Chalk Marl.
ANATOMICAL CHARACTERISTICS Uniform. Heads HABITAT Island nearshore.
moderate in size, subtriangular, teeth stout, subconical,
gently recurved. Trunks and tails very elongated and tails Aphanizocnemus libanensis
especially slender. Ribs heavily constructed. Fingers and 0.3 m (1 ft) TL, 0.28 kg (0.6 lb)
toes usually present. FOSSIL REMAINS Skull and skeleton.
HABITAT Coastal and brackish shorelines, lagoons, reefs, ANATOMICAL CHARACTERISTICS Snout rather small,
estuaries. coronoid process moderate in height. Neck moderately
HABITS Swimming performance good. Shallow-water long. Arms and legs fairly well developed, lower leg bones
ambush and pursuit fishers of small game. Possibly bred very short and broadened.
and nested on beaches. AGE Late Cretaceous; middle Cenomanian.
Aphanizocnemus libanensis
177

Aphanizocnemus libanensis
DISTRIBUTION AND FORMATIONS Lebanon; Hakel or long. Arms fairly well developed.
Sannine. AGE Late Cretaceous; late Cenomanian.
HABITAT Continental nearshore. DISTRIBUTION AND FORMATIONS Croatia; unnamed.
Pontosaurus kornhuberi NOTES May have shared its habitat with Aigialosaurus.
1 m (3.3 ft) TL, 1.1 kg (2.5 lb)
FOSSIL REMAINS Skull and skeleton, latter severely Adriosaurus suessi
flattened top to bottom, with soft tissues. 0.3 m (1 ft) TL, 0.28 kg (0.6 lb)
ANATOMICAL CHARACTERISTICS Snout pointed and FOSSIL REMAINS Majority of skull and skeleton, severely
rather small, teeth small. Neck moderately long. Arms and flattened top to bottom.
legs fairly well developed. ANATOMICAL CHARACTERISTICS Snout moderate in
size. Arms and legs small.
AGE Late Cretaceous; late Cenomanian.
DISTRIBUTION AND FORMATIONS Slovenia; upper
Povir.
Pontosaurus NOTES It is not certain whether the members of this
kornhuberi genus are variations of one species, or different genera.
Shared its habitat with A.? skrbinensis, A.? microbrachis,
Acteosaurus, Eidolosaurus, Mesoleptos, Komensaurus,
Carsosaurus.
Unnamed genus or Adriosaurus skrbinensis

0.4 m (1.3 ft) TL, 0.7 kg (0.15 lb)
AGE Late Cretaceous; middle Cenomanian. FOSSIL REMAINS Partial skull and majority of skeleton,
DISTRIBUTION AND FORMATIONS Lebanon; Sannine. severely flattened top to bottom.
HABITAT Continental nearshore. ANATOMICAL CHARACTERISTICS Arms much shorter
NOTES Some lizard-type scales are preserved on skull and than legs.
body. Shared its habitat with Eupodophis, possibly with AGE Late Cretaceous; late Cenomanian.
Aphanizocnemus. DISTRIBUTION AND FORMATIONS Slovenia; upper
Povir.
Pontosaurus lesinensis HABITAT Island nearshore.
1.2 m (4 ft) TL, 2 kg (4.5 lb) NOTES Not certain belongs in larger-limbed genus
FOSSIL REMAINS Skull and partial skeleton. Adriosaurus.
ANATOMICAL CHARACTERISTICS Snout pointed,
coronoid process short, teeth small. Neck moderately Unnamed genus or Adriosaurus microbrachis
0.5 m (1.6 ft) TL, 0.15 kg (0.35 lb)
FOSSIL REMAINS Majority of skeleton, severely flattened
top to bottom.
ANATOMICAL CHARACTERISTICS Arms reduced to
humerus splinter, legs strongly reduced.
Pontosaurus AGE Late Cretaceous; late Cenomanian.
lesinensis DISTRIBUTION AND FORMATIONS Slovenia; upper
Povir.
NOTES Not certain belongs in much larger-limbed genus
Adriosaurus.
178

Acteosaurus tommasinii
0.5 m (1.6 ft) TL, 0.15 kg (0.35 lb)
OPHIDIANS (SNAKES)
FOSSIL REMAINS Majority of skeleton, severely flattened SMALL OPHIDIOMORPHS FROM THE
top to bottom. LATE CRETACEOUS TO THE MODERN ERA,
ANATOMICAL CHARACTERISTICS Arms much shorter GLOBAL
than legs.
AGE Late Cretaceous; late Cenomanian. ANATOMICAL CHARACTERISTICS Fairly uniform. Heads
DISTRIBUTION AND FORMATIONS Slovenia; upper small, subrectangular in side view, lightly constructed,
Povir. upper temporal bar incomplete, very akinetic, snout not
HABITAT Continental nearshore. large, teeth bladed, recurved with an S-curved, very sharp
tip. Neck moderately long, trunk extremely elongated,
Eidolosaurus trauthi slender, and hyperflexible, tail moderate in length or
Size uncertain short, total vertebrae over 160. Limbs if present vestigial
FOSSIL REMAINS Partial remains. and not functional in locomotion, which is entirely
ANATOMICAL CHARACTERISTICS Insufficient serpentine. Highly hydrodynamically streamlined, entirely
information. anguilliform swimmers.
AGE Late Cretaceous; late Cenomanian. HABITAT Nearshore reefs, reefs, mangroves, lagoons.
DISTRIBUTION AND FORMATIONS Slovenia; upper HABITS Swimming performance good. Ambush and
Povir. crevice fishers of small game. Deposit eggs ashore or give
HABITAT Island nearshore. live birth in the water.
Mesoleptos zendrinii
1 m (3.3 ft) TL, 1 kg (2 lb)
SIMOLIOPHIIDS (SEA SNAKES)
FOSSIL REMAINS Two partial skeletons. SMALL OPHIDIANS FROM THE LATE CRETACEOUS
ANATOMICAL CHARACTERISTICS Neck rather long. OF EUROPE AND THE MIDDLE EAST
DISTRIBUTION AND FORMATIONS Slovenia; upper ANATOMICAL CHARACTERISTICS Uniform. Highly
Povir. aquatic. Arms absent, if legs present they are very
HABITAT Island nearshore. reduced. Nonvenomous.
NOTES A specimen from Palestine referred to this species NOTES Absence from at least some other seas probably
may be a different taxon. reflects lack of sufficient sampling. May be basal snakes
that evolved from marine, dolichosaur-type lizards, or
Judeasaurus tchernovi snakes that evolved from land ancestors and invaded the
1 m (3.3 ft) TL, 1.1 kg (2.5 lb) seas.
FOSSIL REMAINS Majority of skull and minority of
skeleton. Eupodophis descouensi
ANATOMICAL CHARACTERISTICS Snout large, coronoid 0.9 m (3 ft) TL, 0.7 kg (1.5 lb)
process tall, teeth fairly large. FOSSIL REMAINS Skull and skeleton, severely flattened
AGE Late Cretaceous; late Cenomanian or early Turonian. top to bottom.
DISTRIBUTION AND FORMATIONS Israel or West Bank; ANATOMICAL CHARACTERISTICS Skull subtriangular
upper Kefar Sha’ul or Bina. in top view. Tail very short. Foot present but very
HABITAT Continental nearshore. reduced.
AGE Late Cretaceous; middle Cenomanian.
DISTRIBUTION AND FORMATIONS Lebanon;
Sannine.
NOTES Shared its habitat with Pontosaurus kornhuberi,
Judeasaurus possibly with Aphanizocnemus.
tchernovi
Haasiophis terrasanctus
FOSSIL REMAINS Skull and skeleton, probably juvenile,
severely flattened top to bottom.
179

ANATOMICAL CHARACTERISTICS Head subrectangular Pachyrhachis problematicus

in top view, coronoid process short. Tail fairly long. Foot 1.9 m (6 ft) TL, 7 kg (15 lb)
present but very reduced. FOSSIL REMAINS Skull and majority of two skeletons.
AGE Late Cretaceous; early or middle ANATOMICAL CHARACTERISTICS Skull subrectangular
Cenomanian. in top view, coronoid process tall. Tail probably not long.
DISTRIBUTION AND FORMATIONS West Bank; Foot absent.
Amminadav or Bet-Meir. AGE Late Cretaceous; early Cenomanian.
HABITAT Continental nearshore. DISTRIBUTION AND FORMATIONS West Bank;
NOTES Shared its habitat with or may be a juvenile of Bet-Meir.
Pachyrhachis. HABITAT Continental nearshore.
Pachyrhachis problematicus
Pachyrhachis problematicus
180

T E S T U D I N E S ( T U R T L E S )
Pachyophis woodwardi CHELONIOIDEANS

0.6 m (2 ft) TL, 0.2 kg (0.5 lb) (SEA TURTLES)
FOSSIL REMAINS Minority of skull and complete
skeleton, two partial skeletons, severely flattened top to SMALL TO GIGANTIC TESTUDINES FROM
bottom. THE EARLY CRETACEOUS TO THE MODERN
ANATOMICAL CHARACTERISTICS Skeleton thick boned. ERA, GLOBAL
Legs absent.
AGE Late Cretaceous; late Cenomanian. Bouliachelys
DISTRIBUTION AND FORMATIONS Bosnia and (Chelonioideans)
Herzegovina; unnamed.
NOTES Shared its habitat with or may be the adult of
Mesophis.
Mesophis nopscai
0.3 m (1 ft) TL, 0.025 kg (0.06 lb)
FOSSIL REMAINS Majority of skeleton, severely flattened
top to bottom, possibly juvenile.
ANATOMICAL CHARACTERISTICS Skeleton not heavily
built. Legs absent.
DISTRIBUTION AND FORMATIONS Bosnia and
Herzegovina; unnamed.
HABITAT Island nearshore. ANATOMICAL CHARACTERISTICS Fairly uniform.
Heads at least fairly broad, subtriangular in top view.
Necks nonretractable. Trunk enclosed in a very broad,
shallow carapace that is flatter on bottom than on more-
TESTUDINES domed top, never fully plated, dorsal midline row of plates
(TURTLES) are paralleled by a solid row of side plates, outer struts are
separated by lateral spaces inside outer rim of plates, four
SMALL TO GIGANTIC NEODIAPSIDS FROM irregular interlocking plates cover part of ventral carapace.
THE EARLY JURASSIC TO THE MODERN ERA, Tails nonretractable. Limbs nonretractable full flippers,
GLOBAL leading two fingers and sometimes first toe clawed,
humerus points forward and rest of arm arcs out sideways,
ANATOMICAL CHARACTERISTICS Diverse. Fully foreflippers always large and markedly longer and more
terrestrial to fully marine. Skulls akinetic, unsplit hydrofoil shaped than aft flippers. Moderately
nostrils usually at front of snout, snout usually very hydrodynamically streamlined, propulsion provided entirely
short and orbits placed well forward, temporal openings by foreflippers, aft flippers for stability and maneuvering.
closed, parietal eye absent, deep paired embayments HABITAT Coastal to deep oceans.
on rear of skull separated by long midline plate, beaked HABITS Swimming performance good. Herbivorous to
and toothless. Necks short. Trunk enclosed in a omnivorous with small game fishing, prey often including
carapace, most ribs not swept backward. Tail short. jellyfish. Move onto beaches and use clawed foreflippers
Shoulder girdle within carapace, limbs sprawl out to dig nests for their soft-shelled eggs, no parental care.
to sides. NOTES Differing phylogenetic studies do not allow
HABITAT Fully terrestrial to marine. further subdivision into families at this time.
HABITS Herbivorous to omnivorous.
NOTES Relationships to other reptiles uncertain. Desmatochelys? padillai
2 m (6.5 ft) TL, 400 kg (900 lb)
FOSSIL REMAINS Four skulls, majority of skeleton, two
partial skeletons.
ANATOMICAL CHARACTERISTICS Head deep, snout
short. Lateral dorsal spaces large, separated by thin struts,
carapace rim plates narrow. Foreflippers large.
181

AGE Early Cretaceous; late Barremian and/or lower Aptian. AGE Late Cretaceous; late Coniacian? to early Campanian.
DISTRIBUTION AND FORMATIONS Colombia; upper DISTRIBUTION AND FORMATIONS Nebraska, South
Paja. Dakota, Kansas; lower to upper Niobrara?, lower Pierre
HABITAT Continental shelf. Shale.
NOTES That this species is inthe much later and HABITAT Interior seaway was continuing to become less
distinctive genus Desmatochelys is problematic. broad and deep.
NOTES Assigned time span probably too long for this
single species, which could be limited to late Santonian
and/or early Campanian. Shared its habitat with
Styxosaurus browni, Elasmosaurus, Dolichorhynchops
bonneri, Tylosaurus proriger, Latoplatecarpus, Globidens?
dakotensis.
Desmatochelys? Toxochelys moorevillensis

padillai 2 m (6.5 ft) TL, 400 kg (900 lb)
FOSSIL REMAINS Majority of skeleton.
ANATOMICAL CHARACTERISTICS Carapace nearly
circular, lateral dorsal spaces very small.
DISTRIBUTION AND FORMATIONS Alabama; upper
Mooreville Chalk.
NOTES Shared its habitat with Globidens alabamaensis,
Ctenochelys acris, Corsochelys, Prionochelys.
Mexichelys coahuilaensis
0.6 m (2 ft) TL, 12 kg (25 lb)
FOSSIL REMAINS Several skulls and partial skeletons.
Desmatochelys
ANATOMICAL CHARACTERISTICS Head very broad,
lowi
snout short, lower jaw shallow.
DISTRIBUTION AND FORMATIONS Northeastern
Mexico; Cerro del Pueblo.
Desmatochelys lowi
1.9 m (6 ft) TL, 250 kg (550 lb)
ANATOMICAL CHARACTERISTICS Head not deep, snout
medium sized. Carapace heart shaped, lateral dorsal spaces
large, separated by thin struts, carapace rim plates narrow.
Foreflippers large.
AGE Late Cretaceous; late Cenomanian. Mexichelys
DISTRIBUTION AND FORMATIONS Nebraska; middle coahuilaensis
Greenhorn Limestone.
HABITS Interior seaway approaching its greatest maximum.
Toxochelys latiremis
2 m (6.5 ft) TL, 400 kg (900 lb)
FOSSIL REMAINS Skulls, majority of skeleton.
ANATOMICAL CHARACTERISTICS Carapace heart
shaped, lateral dorsal spaces medium sized.
182

Nichollsemys baieri DISTRIBUTION AND FORMATIONS Alabama;

Adult size uncertain upper Mooreville Chalk.
FOSSIL REMAINS Four complete and partial skulls, HABITAT Continental shelf.
possibly immature.
ANATOMICAL CHARACTERISTICS Insufficient Prionochelys matutina
information. 0.7 m (2.3 ft) TL, 15 kg (30 lb)
AGE Late Cretaceous; late Campanian. FOSSIL REMAINS Minority of skull and skeletal remains,
DISTRIBUTION AND FORMATIONS Montana; middle some juvenile.
Bearpaw Shale. ANATOMICAL CHARACTERISTICS Carapace heart
HABITAT Interior seaway was becoming much narrower. shaped, lateral dorsal openings large, separated by thin
NOTES Shared its habitat with Prognathodon? overtoni. struts.
Ctenochelys stenoporus DISTRIBUTION AND FORMATIONS Alabama;
1.3 m (4.3 ft) TL, 65 kg (150 lb) upper Mooreville Chalk.
FOSSIL REMAINS Partial skulls, majority of skeletons. HABITAT Continental shelf.
ANATOMICAL CHARACTERISTICS Carapace oval,
lateral dorsal spaces large, separated by thin struts, Allopleuron hofmanni
carapace rim plates small. 2.3 m (7.5 ft) TL, 700 kg (1,500 lb)
AGE Late Cretaceous; earliest Campanian. FOSSIL REMAINS Skulls and skeletons.
DISTRIBUTION AND FORMATIONS Kansas; ANATOMICAL CHARACTERISTICS Head rather small,
uppermost Niobrara. moderately broad. Carapace heart shaped, lateral dorsal
HABITAT Interior seaway was continuing to become openings large, separated by thin struts, carapace rim plates
less broad and deep. medium sized, bottom plates reduced. Aft foot permanently
NOTES Shared its habitat with Styxosaurus snowii, flexed sideways, aftmost toe short and divergent.
Polycotylus latipinnis, Dolichorhynchops osborni, AGE Late Cretaceous; late Maastrichtian.
Platecarpus tympanicus, Eonatator sternbergii, DISTRIBUTION AND FORMATIONS Belgium; Maastricht.
Clidastes propython, Protostega gigas.
Ctenochelys acris
1 m (3.3 ft) TL, 35 kg (80 lb)
FOSSIL REMAINS Partial skulls and skeletons,
some juvenile.
ANATOMICAL CHARACTERISTICS Head
Allopleuron hofmanni
broad, snout very short. Carapace heart shaped,
carapace rim plates medium sized.
DISTRIBUTION AND FORMATIONS
Alabama; upper Mooreville Chalk.
NOTES Shared its habitat with
Globidens alabamaensis, Toxochelys
moorevillensis, Prionochelys, Corsochelys.
Corsochelys halinches
1.5 m (5 ft) TL, 120 kg (250 lb)
FOSSIL REMAINS Minority of skull and
majority of skeleton.
ANATOMICAL CHARACTERISTICS Head
small. Carapace heart shaped, lateral spaces
large and separated by narrow struts, carapace
rim plates narrow.
183

HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Carapace oval, lateral

NOTES The presence of later members of this genus in dorsal spaces rather small, carapace rim plates large.
the Cenozoic shows that this species survived the K/Pg AGE Late Cretaceous; early Maastrichtian.
extinction. DISTRIBUTION AND FORMATIONS Northern Japan;
Hakobuchi.
Euclastes wielandi HABITAT Island coastal.
Adult size uncertain NOTES Later specimens probably do not belong to this
FOSSIL REMAINS Skulls, complete and partial, possibly taxon.
juvenile.
ANATOMICAL CHARACTERISTICS Head broad. Cratochelone berneyi
Carapace oval, lateral dorsal spaces small. 4? m (13? ft) TL, 3? tonnes
AGE Late Cretaceous; latest Maastrichtian and early FOSSIL REMAINS Minority of skeleton.
Paleocene. ANATOMICAL CHARACTERISTICS Insufficient
DISTRIBUTION AND FORMATIONS New Jersey, information.
Maryland; Hornerstown, Brightseat. AGE Early Cretaceous; middle Albian.
HABITAT Continental shelf. DISTRIBUTION AND FORMATIONS Northeastern
NOTES This species appears to have survived the K/Pg Australia; Toolebuc.
extinction; other species of this genus were present in the HABITAT Interior seaway.
Cenozoic. NOTES Although fragmentary remains make it hard to be
certain, may rival later Archelon as largest known marine
Alienochelys selloumi turtle. Shared its habitat with ?Kronosaurus,
2.5 m (8 ft) TL, 800 kg (1,800 lb) Eromangasaurus, Longirostra, Bouliachelys, and the much
FOSSIL REMAINS Complete skull. more common Notochelone.
ANATOMICAL CHARACTERISTICS Snout very broad and
squared off so head is subrectangular in top view, nostrils Notochelone costata
between orbits, temporal region fairly long, lower jaw 2 m (6.5 ft) TL, 400 kg (900 lb)
fairly deep. FOSSIL REMAINS Skull and partial skeletons.
AGE Late Cretaceous; latest Maastrichtian. ANATOMICAL CHARACTERISTICS Snout short, temporal
DISTRIBUTION AND FORMATIONS Morocco; unnamed. region fairly long, head not deep, moderately broad.
HABITAT Continental shelf. Lateral dorsal spaces moderately large.
Mesodermochelys undulatus DISTRIBUTION AND FORMATIONS Northeastern
2 m (6.5 lb) TL, 400 kg (900 lb) Australia; Toolebuc.
FOSSIL REMAINS Minority of skull and over a dozen HABITAT Interior seaway.
partial skeletons.
Alienochelys
Notochelone
selloumi
costata
184

Bouliachelys
suteri
Bouliachelys suteri
0.5 m (1.5 ft) TL, 5 kg (10 lb)
FOSSIL REMAINS Four skulls, majority of skeleton,
two partial skeletons.
deep and broad.
DISTRIBUTION AND FORMATIONS Northeastern
Australia; Toolebuc.
HABITAT Interior seaway.
Santanachelys gaffneyi
0.2 m (0.6 ft) TL, 0.4 kg (0.9 lb)
FOSSIL REMAINS Skull and skeleton.
Head rather small, snout very short
and temporal region elongated.
Carapace oval, lateral dorsal spaces medium sized, Santanachelys gaffneyi
separated by narrow struts, carapace rim plates narrow.
Foreflipper medium sized, fingers and toes poorly
ossified, no clawed toe present.
AGE Early Cretaceous; late Aptian
and/or early Albian.
Northeastern Brazil; Romualdo.
Chelosphargis advena
0.6 m (2 ft) TL, 6 kg (13 lb)
FOSSIL REMAINS Partial skull and skeletons.
ANATOMICAL CHARACTERISTICS Carapace heart
shaped, lateral dorsal spaces small, carapace rim plates
large.
AGE Late Cretaceous; late Coniacian.
DISTRIBUTION AND FORMATIONS Kansas; lower Rhinochelys amaberti
Niobrara. Adult size not certain
HABITAT Continental shelf. FOSSIL REMAINS Skulls, possibly juvenile.
NOTES Shared its habitat with Selmasaurus? johnsoni, ANATOMICAL CHARACTERISTICS Head deep,
Ectenosaurus, Plesioplatecarpus. quite broad, snout fairly broad.
185

AGE Early Cretaceous; late Albian.

Eastern France; unnamed.
Rhinochelys Rhinochelys pulchriceps

pulchriceps 0.4 m (1.3 ft) TL, 3.5 kg (8 lb)
FOSSIL REMAINS A few skulls.
Head very deep, snout quite broad and
very short.
AGE Late Cretaceous; early
Cenomanian.
Eastern England; lower West Melbury
Marly Chalk.
Rhinochelys? nammourensis
0.85 m (2.75 ft) TL, 35 kg (80 lb)
FOSSIL REMAINS Majority of skull and
skeleton.
Head very large, snout not broad,
quite short, and temporal region long.
Carapace heart shaped, lateral dorsal
spaces small, separated by short robust
struts, carapace rim plates large.
Foreflipper medium sized, aft flipper
Rhinochelys? small, no clawed toe present.
nammourensis AGE Late Cretaceous; middle
Cenomanian.
Lebanon; unnamed.
Protostega gigas
2.2 m (7.2 ft) TL, 650 kg (1,400 lb)
FOSSIL REMAINS A number of skulls
Protostega and skeletons of varying completeness.

gigas ANATOMICAL CHARACTERISTICS
Head very large, beak parrot-like.
Carapace roundish, lateral dorsal spaces
large, separated by long narrow struts,
carapace rim plates large. Foreflipper
large, aft flipper fairly large.
AGE Late Cretaceous; earliest
Campanian.
Kansas, Alabama; uppermost Niobrara,
lower Mooreville Chalk.
HABITAT Interior seaway was
186

continuing to become less broad and

deep, continental shelf.
NOTES Shared its habitat with
Styxosaurus snowii, Polycotylus
latipinnis, Dolichorhynchops osborni,
Selmasaurus russelli, Platecarpus
tympanicus, Eonatator sternbergii,
Clidastes propython, Ctenochelys
stenoporus. Second largest known
marine turtle. May be a direct ancestor
of Archelon.
Protostega gigas
Archelon ischyros
FOSSIL REMAINS Five skulls and/or
skeletons of varying completeness.
Head large, beak parrot-like. Carapace
somewhat squarish, lateral dorsal
spaces very large, separated by long
narrow struts, carapace rim plates
small. Foreflipper very large, aft flipper
fairly large.
South Dakota, Wyoming; upper Pierre
Shale.
Archelon ischyros
187

Archelon
ischyros
matched by earlier, less well-preserved

Cratochelone berneyi, second largest of
all known turtles.
Archelon ischyros Terlinguachelys fischbecki

2.5 m (8 ft) TL, 450 kg (1,000 lb)
broad. Carapace somewhat elongated.
AGE Late Cretaceous; middle Campanian.
DISTRIBUTION AND FORMATIONS Texas; middle Aguja.
HABITAT Interior seaway was becoming much narrower. HABITAT Interior seaway.
HABITS Large size indicates may have preferred deeper
waters, but shallowing seaway indicates opposite. Strongly Ocepechelon bouyai
hooked beak may have been used for predation, grubbing 3.5 m (11 ft) TL, 2 tonnes
on seabeds, intraspecific combat, scavenging, FOSSIL REMAINS Majority of skull.
defense against predators. ANATOMICAL CHARACTERISTICS Head shallow, very
NOTES Shared its habitat with Prognathodon? overtoni, broad aft, tip of snout tubular, snout elongated, nostrils
Mosasaurus? missouriensis. The classic Mesozoic marine well back between orbits that face strongly upward,
turtle. Largest known sea turtle, although may be temporal region expanded well aft of jaw joint.
188

A R C H O S A U R O M O R P H S
May have been only archosaurs to bear live young aside

from metriorhynchid crocs.
Dinocephalosaurus orientalis
Ocepechelon 3 m (11 ft) TL
bouyai FOSSIL REMAINS Skull and two partial skeletons, fetus,
severely flattened from top to bottom.
ANATOMICAL CHARACTERISTICS Head somewhat
elongated and subrectangular in top view because
squared-off snout tip is fairly broad, large nostrils set a
little aft of snout tip, orbits set well aft and temporal
AGE Late Cretaceous; latest Maastrichtian. region not large, lower jaws shallow, a few upper and
DISTRIBUTION AND FORMATIONS Morocco; unnamed. lower teeth large. Neck extremely long primarily because
HABITAT Continental shallows. of elongation of vertebrae.
HABITS Apparent suction feeder. AGE Middle Triassic; middle Anisian.
NOTES Shared its habitat with Zarafasaura. DISTRIBUTION AND FORMATIONS Southern China;
middle Guanling.
ARCHOSAURO- HABITS Ambush fisher of small game.
MORPHS NOTES Not sufficiently well known to estimate mass.

Shared its habitat with Atopodentatus, Dianopachysaurus.
SMALL TO GIGANTIC NEODIAPSIDS FROM
THE MIDDLE PERMIAN TO THE MODERN
ERA, GLOBAL ARCHOSAURIFORMES
ANATOMICAL CHARACTERISTICS Extremely diverse. SMALL TO GIGANTIC ARCHOSAUROMORPHS
Cervical ribs often overlap one another. FROM THE LATE PERMIAN TO THE MODERN
HABITAT AND HABITS Extremely variable, marine to ERA, GLOBAL
fully terrestrial to highly aerial, highly herbivorous to
archpredatory. ANATOMICAL CHARACTERISTICS Extremely diverse.
NOTES Includes modern crocodilians and birds, which Preorbital opening in front of orbits and opening in
are living dinosaurs. middle of lower jaws, one or both sometimes closed off,
parietal eye very reduced or absent, teeth when present
set in sockets.
TANYSTROPHEIDS HABITAT AND HABITS Extremely variable, marine to
fully terrestrial to highly aerial, highly herbivorous to
SMALL TO MEDIUM-SIZED NEODIAPSIDS archpredatory.
FROM THE TRIASSIC OF THE NORTHERN
HEMISPHERE CRUROTARSI OR
PSEUDOSUCHIANS
ANATOMICAL CHARACTERISTICS Diverse. Fully
terrestrial to aquatic. Skulls akinetic, orbits face strongly SMALL TO GIGANTIC ARCHOSAURIFORMES
upward, scleral rings present, teeth conical, some on FROM THE EARLY TRIASSIC TO THE MODERN
mouth roof. Necks at least somewhat elongated, slender ERA, GLOBAL
cervical rib aft rods elongated and overlapping. Trunks
fairly elongated. Digits poorly streamlined, aquatic ANATOMICAL CHARACTERISTICS Diverse. Skulls
propulsion mainly from limbs. akinetic, parietal eye absent. Necks including cervical ribs
HABITAT Terrestrial and/or freshwaters to coastal short. Trunks fairly long, not broad, gastralia slender.
nearshore. Tails moderately long. Ankle a complex joint with a large
HABITS Swimming performance poor to mediocre. At calcaneal heel tuber. Usually armored to some degree.
least some probably gave live birth. HABITAT Fully terrestrial to highly marine.
NOTES Aquatic nature of other members of this group, HABITS Herbivorous to archpredatory.
including hyperlong-necked Tanystropheus, is uncertain.
189

CROCODYLIFORMES THALATTOSUCHIANS
SMALL TO GIGANTIC PSEUDOSUCHIANS SMALL TO LARGE MESOEUCROCODYLIANS
FROM THE LATE TRIASSIC TO THE MODERN FROM THE EARLY JURASSIC TO EARLY
ERA, GLOBAL CRETACEOUS, GLOBAL
ANATOMICAL CHARACTERISTICS Diverse. Heads ANATOMICAL CHARACTERISTICS Somewhat diverse.

heavily constructed, akinetic, often shallow with roof Highly aquatic. Upper temporal openings large. Aft tail
flattened, nostrils at tip of snout, preorbital opening flattened. Arms strongly reduced both absolutely and
reduced or closed off, temporal openings often reduced compared to legs, latter are paddles, inner fingers and toes
or closed off, squamosal at upper rear corner of skull most robust, toes increasingly long progressing outward.
massive, teeth usually have bulbous roots, no teeth on Moderately hydrodynamically streamlined, primarily axial
mouth roof. Cervical rib overlap modest when present. undulators of anguilliform grade, limbs primarily for
Gastralia limited to aft portion of abdomen, rib-free stability and maneuvering, pressed tight to body when
lumbar region present. Clavicles absent, coracoid hydrocruising.
elongated. A pair of carpals elongated. Liver pump HABITAT Freshwaters to perhaps deep oceans.
respiratory system partially present. HABITS Swimming performance fair to very good.
HABITAT Fully terrestrial to highly marine. Ambush and/or pursuit fishers and hunters of small to
HABITS Herbivorous to archpredatory. big game. Probably able to move onshore and breed on
beaches.
MESOEUCROCODYLIANS NOTES Restriction of nostrils to tip of snout may have
inhibited evolution of group as marine creatures.
SMALL TO GIGANTIC CROCODYLIFORMES
FROM THE EARLY JURASSIC TO THE MODERN TELEOSAURIDS
ERA, GLOBAL
SMALL TO LARGE THALATTOSUCHIANS
ANATOMICAL CHARACTERISTICS Fairly diverse. Partial FROM THE EARLY TO LATE JURASSIC
secondary mouth roof present. Pelvis at least somewhat OF THE EASTERN HEMISPHERE
reduced, mobile pubis part of the liver-pump respiratory
system. Toes increasingly long progressing outward. ANATOMICAL CHARACTERISTICS Uniform. Heads
HABITAT Fully terrestrial to highly marine. shallow, snouts elongated, tips a little expanded laterally,
HABITS Herbivorous to archpredatory. snouts much narrower than temporal region, preorbital
NOTES Includes modern crocodilians. opening closed, lower jaws shallow, teeth numerous, not
large, procumbent. Aft tail straight. Hands very small, not
Pelagosaurus (Thalattosuchians) hydrodynamic. Parallel single rows of armor atop midline
of neck, trunk, and most of tail, multiple rows of armor
along midline of aft abdomen and first half of tail.
HABITAT Coastal freshwaters to perhaps deep oceans.
HABITS Swimming performance fair. Ambush and pursuit
fishers and hunters of small to medium-sized game.
NOTES The once major multispecies genus Steneosaurus
has been replaced by a number of new genera.
Macrospondylus muscle study
190

ANATOMICAL CHARACTERISTICS Snout very long and

Plagiophthalmosuchus narrow, tip laterally expanded.
gracilirostris AGE Late Jurassic; late Kimmeridgian.
southwestern France; lower Kimmeridge Clay, Ancholme
Group.
Plagiophthalmosuchus gracilirostris HABITAT Island archipelago shallows.
3 m (10) TL, 60 kg (130 lb) HABITS Preferred small prey.
FOSSIL REMAINS Skull and skeleton severely flattened NOTES Shared its habitat with Pliosaurus brachydeirus,
top to bottom, skull. Colymbosaurus megadeirus, Torvoneustes, Plesiosuchus,
ANATOMICAL CHARACTERISTICS Head long, snout Neosteneosaurus, Lemmysuchus.
very long and very narrow, aft head much shorter, teeth
very numerous and slender. Mystriosaurus laurillardi
AGE Early Jurassic; early Toarcian. 4 m (13 ft) TL, 140 kg (300 lb)
DISTRIBUTION AND FORMATIONS Northeastern FOSSIL REMAINS Majority of skull, partial skull.
England, Luxembourg; lower Whitby Mudstone, unnamed. ANATOMICAL CHARACTERISTICS Snout elongated and
HABITAT Island shallows. narrow, robust, teeth medium sized and fairly stout.
HABITS Preferred small prey. AGE Early Jurassic; early Toarcian.
NOTES Shared its habitat with Rhomaleosaurus, DISTRIBUTION AND FORMATIONS Southern Germany,
Hauffiosaurus, Eurhinosaurus, Mystriosaurus, northeastern England; Posidonienschiefer, lower Whitby
Macrospondylus. Mudstone.
Mycterosuchus nasutus NOTES Shared its habitat with Meyerasaurus,
5 m (16 ft) TL, 250 kg (550 lb) Rhomaleosaurus, Hauffiosaurus, Seeleyosaurus, Hydrorion,
FOSSIL REMAINS Skulls complete and partial, skeleton Hauffiopteryx, unnamed genus trigonodon, Suevoleviathan,
severely flattened top to bottom. Eurhinosaurus, Stenopterygius, Plagiophthalmosuchus,
ANATOMICAL CHARACTERISTICS Snout very long and Macrospondylus, Platysuchus.
very narrow, aft head much shorter, teeth very numerous
and small. Platysuchus multiscrobiculatus
AGE Early Jurassic; late middle Callovian. 2.75 m (9 ft) TL, 45 kg (100 lb)
DISTRIBUTION AND FORMATIONS Central England; FOSSIL REMAINS Skull and skeleton, severely flattened
upper Oxford Clay. top to bottom.
HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Head long, snout
HABITS Preferred small prey. very long and very narrow, aft head much shorter, teeth
very numerous and slender.
Aeolodon priscus AGE Early Jurassic; early Toarcian.
3.5 m (11 ft) TL, 90 kg (200 lb) DISTRIBUTION AND FORMATIONS Southern Germany;
FOSSIL REMAINS Two complete skulls and skeletons, Posidonienschiefer.
severely flattened top to bottom. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Head long, snout HABITS Preferred small prey.
very numerous and small. Teleosaurus cadomensis
AGE Late Jurassic; early Tithonian. 3 m (10) TL, 60 kg (130 lb)
DISTRIBUTION AND FORMATIONS Southern Germany, FOSSIL REMAINS Several specimens.
southeastern France; Mörnsheim, unnamed. ANATOMICAL CHARACTERISTICS Snout extremely
HABITAT Island archipelago shallows. long and very narrow, aft head much shorter, teeth
HABITS Preferred small prey. very numerous and slender.
NOTES Shared its habitat with Geosaurus giganteus, AGE Middle Jurassic; Bathonian.
Dakosaurus maximus, Cricosaurus elegans, Rhacheosaurus.
Bathysuchus megarhinus
4 m (13 ft) TL, 140 kg (300 lb) Teleosaurus
cadomensis
FOSSIL REMAINS Minority of two skulls.
191

DISTRIBUTION AND FORMATIONS Northern France; DISTRIBUTION AND FORMATIONS Central China;
unnamed. Ziliujing.
HABITAT Island archipelago shallows. HABITAT Coastal shallows.
HABITS Preferred small prey. NOTES Was previously assigned to poorly preserved and
NOTES Most original remains destroyedby Allied different Peipehsuchus teleorhinus. Shared its habitat with
bombardment in World War II. Probably includes Bishanopliosaurus youngi.
T. geoffroyi.
Indosinosuchus potamosiamensis
MACHIMOSAURIDS
4 m (13 ft) TL, 140 kg (300 lb) SMALL TO LARGE THALATTOSUCHIANS FROM
FOSSIL REMAINS Complete and partial skulls. THE EARLY JURASSIC TO EARLY CRETACEOUS
ANATOMICAL CHARACTERISTICS Snout moderately OF THE EASTERN HEMISPHERE
elongated, aft head about as elongated, teeth moderately
numerous, medium sized, fairly stout. ANATOMICAL CHARACTERISTICS Uniform. Heads
AGE Late Jurassic; Tithonian? shallow, snouts elongated, tips a little expanded laterally,
DISTRIBUTION AND FORMATIONS Thailand; lower snouts much narrower than temporal region, preorbital
Phu Kradung. opening closed, lower jaws shallow, teeth numerous,
HABITAT Coastal fresh watercourses. procumbent. Aft tail straight. Hands very small, not
NOTES May include I. kalasinensis. Appears to show hydrodynamic. Parallel single rows of armor atop midline
that at least some marine crocodilians entered freshwater. of neck, trunk, and most of tail, multiple rows of armor
Age poorly known. along midline of aft abdomen and first half of tail.
HABITAT Coastal freshwaters to perhaps deep oceans.
Unnamed genus and species HABITS Swimming performance fair. Ambush and pursuit
4 m (13 ft) TL, 140 kg (300 lb) fishers and hunters of small to medium-sized game.
FOSSIL REMAINS Majority of skull, severely flattened top NOTES The once major multispecies genus Steneosaurus
to bottom. has been replaced by a number of new genera.
ANATOMICAL CHARACTERISTICS Snout very long and
very narrow, aft head much shorter, teeth very numerous Macrospondylus bollensis
and small. 5 m (16.5 ft) TL, 250 kg (550 lb)
AGE Early Jurassic; Toarcian. FOSSIL REMAINS Numerous skulls and skeletons, many
complete.
ANATOMICAL CHARACTERISTICS Head long, snout
Indosinosuchus very numerous and rather slender.
potamosiamensis
Macrospondylus bollensis
192

Macrospondylus bollensis
AGE Early Jurassic; early Toarcian. HABITAT Island archipelago shallows.

DISTRIBUTION AND FORMATIONS Southern Germany, HABITS Preferred small prey.
Luxembourg, northeastern England; Posidonienschiefer, NOTES Shared its habitat with Yvridiosuchus.
lower Whitby Mudstone.
HABITAT Island archipelago shallows. Charitomenosuchus leedsi
HABITS Preferred small prey. 3 m (10) TL, 60 kg (130 lb)
NOTES Shared its habitat with Meyerasaurus, FOSSIL REMAINS A complete and partial skulls, severely
Rhomaleosaurus, Hauffiosaurus, Seeleyosaurus, Hydrorion, flattened top to bottom.
Hauffiopteryx, unnamed genus trigonodon, ANATOMICAL CHARACTERISTICS Snout extremely long
Suevoleviathan, Eurhinosaurus, Stenopterygius, and very narrow, aft head much shorter, teeth very
Plagiophthalmosuchus, Mystriosaurus, Platysuchus. The numerous and slender.
classic marine crocodilian, long placed within Steneosaurus. AGE Middle Jurassic; middle Callovian.
Clovesuurdameredeor stephani northern France; middle Oxford Clay, Marnes de Dives.
5 m (16.5 ft) TL, 250 kg (550 lb) HABITAT Island archipelago shallows.
FOSSIL REMAINS Partial skull. HABITS Preferred small prey.
ANATOMICAL CHARACTERISTICS Insufficient NOTES Shared its habitat with Neosteneosaurus,
information. Lemmysuchus, Thalattosuchus.
AGE Middle Jurassic; Bathonian.
DISTRIBUTION AND FORMATIONS Southern England; Proexochokefalos heberti
Cornbrash. 5.5 m (18 ft) TL, 600 kg (1,300 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS Skull.
HABITS Preferred small prey. ANATOMICAL CHARACTERISTICS Snout moderately
elongated, aft head almost as elongated, teeth limited to
Seldsienean megistorhynchus front half of jaws, moderately numerous, small, fairly
8 m (26 ft) TL, 1 tonne stout.
FOSSIL REMAINS Partial skulls. AGE Middle Jurassic; late Callovian.
ANATOMICAL CHARACTERISTICS Snout very long and DISTRIBUTION AND FORMATIONS Northwestern
narrow, tip laterally expanded. France; Marnes de Dives.
AGE Middle Jurassic; Bathonian. HABITAT Island archipelago shallows.
DISTRIBUTION AND FORMATIONS Southern England, NOTES Shared its habitat with Charitomenosuchus.
France; Cornbrash, unnamed.
Proexochokefalos
HABITS Preferred small prey.
heberti
Deslongchampsina larteti
3 m (10) TL, 60 kg (130 lb)
FOSSIL REMAINS Several complete and partial skulls. Proexochokefalos bouchardi
ANATOMICAL CHARACTERISTICS Snout very long and 5 m (16.5 ft) TL, 250 kg (550 lb)
very narrow, aft head much shorter, teeth moderately FOSSIL REMAINS Majority of skull.
numerous, fairly large and slender. ANATOMICAL CHARACTERISTICS Snout moderately
AGE Middle Jurassic; late Bathonian. elongated, aft head almost as elongated, teeth limited to
DISTRIBUTION AND FORMATIONS Central England; front half of jaws, moderately numerous, small, fairly
Cornbrash. stout.
193

AGE Middle Jurassic; Kimmeridgian.

DISTRIBUTION AND FORMATIONS Northwestern Yvridiosuchus
France; Reuchenette. boutilieri
Neosteneosaurus edwardsi
5 m (16.5 ft) TL, 250 kg (550 lb)
FOSSIL REMAINS Complete and partial remains. HABITS Preferred small prey.
ANATOMICAL CHARACTERISTICS Snout elongated and NOTES Shared its habitat with Deslongchampsina.
narrow.
AGE Middle Jurassic; middle Callovian. Unnamed genus buffetauti
DISTRIBUTION AND FORMATIONS Southern England, 5 m (16.5 ft) TL, 350 kg (800 lb)
northern France; middle Oxford Clay, Ancholme Group. FOSSIL REMAINS Two skulls and partial skeleton.
HABITAT Island archipelago shallows. ANATOMICAL CHARACTERISTICS Snout elongated,
HABITS Preferred small prey. robust, temporal region very large in length and breadth,
NOTES Shared its habitat with Bathysuchus, Lemmysuchus, teeth medium sized.
Charitomenosuchus. AGE Late Jurassic; early Kimmeridgian.
DISTRIBUTION AND FORMATIONS Southern and
Andrianavoay baroni northern Germany, northern France; Lacunosamergal,
5 m (16.5 ft) TL, 250 kg (550 lb) Langenberg, Calcaires Coguilliers.
FOSSIL REMAINS Partial skull. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Snout elongated and HABITS Preferred larger prey.
narrow. NOTES Placement in Machimosaurus very dubious
AGE Middle Jurassic; Bathonian. because original genus and species M. hugii is based on
DISTRIBUTION AND FORMATIONS Madagascar; poorly preserved material.
uncertain.
Lemmysuchus obtusidens Unnamed genus

5 m (16.5 ft) TL, 250 kg (550 lb) buffetauti
FOSSIL REMAINS Several specimens, complete to partial.
ANATOMICAL CHARACTERISTICS Snout moderately
elongated, aft head almost as elongated.
DISTRIBUTION AND FORMATIONS Eastern England,
northwestern France; middle Oxford Clay, Ancholme Unnamed genus mosae
Group. 5 m (16.5 ft) TL, 350 kg (800 lb)
HABITAT Island archipelago shallows. FOSSIL REMAINS Majority of two skulls and skeletons.
NOTES Shared its habitat with Bathysuchus, ANATOMICAL CHARACTERISTICS Snout elongated,
Neosteneosaurus, Charitomenosuchus. robust, temporal region very large in length and breadth,
teeth medium sized.
Yvridiosuchus boutilieri AGE Late Jurassic; latest Kimmeridgian and/or lowest
4 m (13 ft) TL, 140 kg (300 lb) Tithonian.
FOSSIL REMAINS Two partial skulls. DISTRIBUTION AND FORMATIONS Eastern France;
ANATOMICAL CHARACTERISTICS Snout elongated and unnamed, Argiles de Châtillon.
narrow, robust, aft head almost as elongated, shallow, HABITAT Island archipelago shallows.
teeth moderately numerous, medium sized, and fairly HABITS Preferred larger prey.
stout. NOTES Placement in Machimosaurus very dubious
AGE Middle Jurassic; late Bathonian. because original genus and species M. hugii is based on
DISTRIBUTION AND FORMATIONS Central England, poorly preserved material. One specimen was destroyed
northern France; Cornbrash, unnamed. by Allied bombardment in World War II.
194

Unnamed genus rex

7.5 (25 ft) TL, 1 tonne
PELAGOSAURIDS
FOSSIL REMAINS Partial skull and minority of skeleton, MEDIUM-SIZED TO LARGE METRIORHYNCHOIDS
severely flattened top to bottom. FROM THE EARLY TO LATE JURASSIC OF
ANATOMICAL CHARACTERISTICS Snout robust, EUROPE
temporal region very large in length and breadth.
AGE Early Cretaceous; late Hauterivian and/or early ANATOMICAL CHARACTERISTICS Uniform. Heads
Barremian. shallow, snouts elongated, tips a little expanded laterally,
DISTRIBUTION AND FORMATIONS Tunisia; lower snouts much narrower than temporal region, preorbital
Douiret. opening closed, lower jaws shallow, teeth numerous, not
HABITAT Continental shallows. large. Aft tail straight. Hands very small, not
HABITS Preferred larger prey. hydrodynamic. Parallel single rows of armor along midline
NOTES Original placement in much earlier of neck, trunk, and most of tail, twin rows of armor along
Machimosaurus highly problematic. Indicates that midline of aft abdomen and first half of tail.
teleosaurids persisted into Early Cretaceous. HABITAT Coastal.
HABITS Swimming performance good. Ambush and
METRIORHYNCHOIDS pursuit fishers and hunters of small to medium-sized game.
SMALL TO LARGE THALATTOSUCHIANS FROM Pelagosaurus typus

THE EARLY JURASSIC TO EARLY CRETACEOUS 3 m (10 ft) TL, 60 kg (130 lb)
OF EUROPE AND THE AMERICAS FOSSIL REMAINS Numerous specimens, some nearly
complete.
ANATOMICAL CHARACTERISTICS Orbits face more ANATOMICAL CHARACTERISTICS Head very long,
sideways. Armor at least a little reduced. snout extremely long and very narrow, aft head much
HABITAT Coastal shallows to perhaps deep oceans. shorter, teeth moderate in number and slender. Tail
HABITS Swimming performance good to very good. moderately long.
Ambush and/or pursuit fishers and hunters of small to AGE Early Jurassic; Toarcian.
big game. DISTRIBUTION AND FORMATIONS Southern England,
NOTES Absence from at least some other seas probably northern France; lower Upper Lias.
reflects lack of sufficient sampling. HABITAT Island archipelago shallows.
Pelagosaurus typus
195

Magyarosuchus fitosi
4.5 m (15 ft) TL, 200 kg (450 lb)
METRIORHYNCHIDS
FOSSIL REMAINS Partial remains. SMALL TO LARGE METRIORHYNCHOIDS
ANATOMICAL CHARACTERISTICS Insufficient FROM THE LATE JURASSIC TO THE EARLY
information. CRETACEOUS, GLOBAL
AGE Late Jurassic; late Tithonian.
DISTRIBUTION AND FORMATIONS Hungary; Kisgerecse ANATOMICAL CHARACTERISTICS Fairly uniform. Orbits
Marl. face largely sideways. Sharply downward-flexed aft tail
HABITAT Island archipelago shallows. vertebrae supported large soft upper fin that was dorsal
half of semilunate tail flukes. Forelimbs a small flipper.
Teleidosaurus calvadosii Pelvis reduced. Unarmored.
3 m (10 ft) TL, 70 kg (150 lb) HABITAT Coastal shallows to perhaps deep oceans.
FOSSIL REMAINS Skull, other remains. HABITS Swimming performance very good. Ambush and
ANATOMICAL CHARACTERISTICS Snout moderately especially pursuit fishers and hunters of small to big game.
elongated, robust, teeth moderate in number, short
and stout. GEOSAURINES
AGE Middle Jurassic; middle Bathonian.
DISTRIBUTION AND FORMATIONS Northern France; SMALL TO LARGE METRIORHYNCHIDS FROM
Calcaire de Caen. THE MIDDLE JURASSIC OF EUROPE AND
HABITAT Island archipelago shallows. SOUTH AMERICA
NOTES Some remains destroyed by Allied bombardment
in World War II. ANATOMICAL CHARACTERISTICS Snouts not extremely
elongated, preorbital opening closed.
HABITAT Coastal shallows to perhaps deep oceans.
HABITS Fishers and hunters of small to big game.
Teleidosaurus NOTES Absence from at least some other seas probably
calvadosii reflects lack of sufficient sampling.
Purranisaurus casamiquelai
3 m (10 ft) TL, 80 (180 lb)
FOSSIL REMAINS Two skulls, complete and partial.
Eoneustes gaudryi ANATOMICAL CHARACTERISTICS Teeth limited tofront
3 m (10 ft) TL, 70 kg (150 lb) portions of jaws, fairly large and robust.
FOSSIL REMAINS Partial skulls and skeletons. AGE Middle Jurassic; middle Callovian.
ANATOMICAL CHARACTERISTICS Insufficient DISTRIBUTION AND FORMATIONS Northern Chile;
information. unnamed.
AGE Middle Jurassic; late Bajocian to middle Bathonian. HABITAT Continental shelf.
DISTRIBUTION AND FORMATIONS Northern France; NOTES Probably includes P. westermanni.
Upper Lias.
HABITAT Island archipelago shallows. Purranisaurus potens
NOTES May include Eoneustes bathonicus. Some remains 2.5 m (8 ft) TL, 50 kg (110 lb)
destroyed by Allied bombardment in World War II. FOSSIL REMAINS Majority of skull.
ANATOMICAL CHARACTERISTICS Teeth not large.
Zoneait nargorum AGE Late Jurassic and/or Early Cretaceous; late
5 m (16 ft) TL, 350 kg (750 lb) Tithonian and/or early Berriasian.
FOSSIL REMAINS Several partial skulls and minority of
skeleton.
ANATOMICAL CHARACTERISTICS Standard for group.
AGE Middle Jurassic; latest Aalenian and/or earliest
Bajocian. Purranisaurus
DISTRIBUTION AND FORMATIONS Oregon; lower potens
Snowshoe.
196

DISTRIBUTION AND FORMATIONS Central Argentina; NOTES May include Geosaurus grandis. Shared its habitat
upper Vaca Muerta. with Aeolodon, Dakosaurus maximus, Rhacheosaurus,
HABITAT Continental shelf. Cricosaurus elegans.
NOTES Shared its habitat with Cricosaurus puelchorum.
Suchodus brachyrhynchus
Neptunidraco ammoniticus 3 m (10 ft) TL, 80 kg (180 lb)
2.5 m (8 ft) TL, 50 kg (110 lb) FOSSIL REMAINS Majority of skull, severely flattened top
FOSSIL REMAINS Partial skull and minority of skeleton. to bottom.
ANATOMICAL CHARACTERISTICS Insufficient ANATOMICAL CHARACTERISTICS Snout not elongated.
information. AGE Middle Jurassic; middle Callovian.
AGE Middle Jurassic; late Bajocian and/or earliest DISTRIBUTION AND FORMATIONS Southern England;
Bathonian. lower Oxford Clay.
DISTRIBUTION AND FORMATIONS Northern Italy; HABITAT Island archipelago shallows.
lower Rosso Ammonitico Veronese. NOTES Shared its habitat with Peloneustes,
HABITAT Island archipelago shallows. Pachycostasaurus, Simolestes, Liopleurodon, Cryptoclidus,
Muraenosaurus, Tricleidus, Ophthalmosaurus,
Torvoneustes carpenteri Tyrannoneustes, Gracilineustes.
4.5 m (15 ft) TL, 275 kg (600 lb)
FOSSIL REMAINS Two partial skulls. Plesiosuchus manselii
ANATOMICAL CHARACTERISTICS Snout not elongated, 7 m (23 ft) TL, 1 tonne
teeth fairly large. FOSSIL REMAINS Partial skull.
AGE Late Jurassic; late Kimmeridgian. ANATOMICAL CHARACTERISTICS Snout not elongated.
DISTRIBUTION AND FORMATIONS Southern England; AGE Late Jurassic; late Kimmeridgian.
lower Kimmeridge Clay. DISTRIBUTION AND FORMATIONS Southern England;
HABITAT Island archipelago shallows. lower Kimmeridge Clay.
NOTES May include T. coryphaeus. Shared its habitat with HABITAT Island archipelago shallows.
Pliosaurus brachydeirus, Colymbosaurus megadeirus, NOTES Shared its habitat with Pliosaurus brachydeirus,
Bathysuchus, Plesiosuchus. Colymbosaurus megadeirus, Bathysuchus, Torvoneustes.
Tyrannoneustes lythrodectikos
2.5 m (8 ft) TL, 50 kg (110 lb)
DAKOSAURINES
FOSSIL REMAINS Partial skull and skeletal remains. MEDIUM-SIZED METRIORHYNCHIDS FROM
ANATOMICAL CHARACTERISTICS Teeth stout. THE MIDDLE TO LATE JURASSIC OF EUROPE
AGE Middle Jurassic; middle Callovian. AND SOUTH AMERICA
lower Oxford Clay. ANATOMICAL CHARACTERISTICS Heads deep along
HABITAT Island archipelago shallows. most of their length, moderately broad, preorbital
NOTES Shared its habitat with Peloneustes, opening small, bar on upper orbital rim shaded eyes, teeth
Pachycostasaurus, Simolestes, Liopleurodon, Cryptoclidus, moderate in number and quite large. Trunk shallow.
Muraenosaurus, Tricleidus, Ophthalmosaurus, Suchodus, Foreflipper subcircular, clawless.
Gracilineustes. HABITAT Coastal shallows to perhaps deep oceans.
HABITS Pursuit fishers and hunters of small to especially
Geosaurus giganteus big game.
3 m (10 ft) TL, 80 (180 lb) NOTES Absence from at least some other seas probably
FOSSIL REMAINS Four skulls, severely flattened or partial. reflects lack of sufficient sampling.
ANATOMICAL CHARACTERISTICS Snout not elongated,
teeth large, intermeshing. Dakosaurus unnamed species
AGE Late Jurassic; late early Tithonian. 4 m (13 ft) TL, 140 kg (300 lb)
DISTRIBUTION AND FORMATIONS Southern Germany; FOSSIL REMAINS Complete skull and skeleton.
Mörnsheim. ANATOMICAL CHARACTERISTICS Head very large,
HABITAT Island archipelago shallows. subtriangular.
AGE Late Jurassic; late Kimmeridgian.
197

Dakosaurus unnamed species
DISTRIBUTION AND FORMATIONS Southern Germany; AGE Late Jurassic; late Tithonian.
Torleite. DISTRIBUTION AND FORMATIONS Central Argentina;
HABITAT Island archipelago shallows. middle Vaca Muerta.
NOTES May be direct ancestor of D. maximus. HABITAT Continental shelf.
NOTES Shared its habitat with Arthropterygius?
Dakosaurus maximus thalassonotus, unnamed genus and species, Sumpalla?
4.5 m (15 ft) TL, 200 kg (450 lb)
FOSSIL REMAINS Complete and partial skulls and METRIORHYNCHINES
skeleton.
ANATOMICAL CHARACTERISTICS Head very large, MEDIUM-SIZED METRIORHYNCHIDS FROM
subtriangular. THE MIDDLE TO LATE JURASSIC OF EUROPE
AGE Late Jurassic; late early Tithonian.
DISTRIBUTION AND FORMATIONS Southern Germany; ANATOMICAL CHARACTERISTICS Snouts highly
Mörnsheim. elongated, fairly narrow, preorbital opening closed, lower
HABITAT Island archipelago shallows. jaws shallow, teeth medium sized, robust. Foreflipper
NOTES Shared its habitat with Aeolodon, Geosaurus clawed.
giganteus, Rhacheosaurus, Cricosaurus elegans. HABITAT Coastal.
HABITS Fishers of small to medium-sized game.
Dakosaurus andiniensis NOTES The once major multispecies genus Metriorhynchus
5 m (16.5 ft) TL, 275 kg (600 lb) has been replaced by new genera. Absence from at least
FOSSIL REMAINS Complete and partial skulls, minority of some other seas may reflect lack of sufficient sampling.
skeleton.
ANATOMICAL CHARACTERISTICS Head more Thalattosuchus superciliosus
rectangular in side and top views, teeth more robust, 3 m (10 ft) TL, 115 kg (255 lb)
lower jaw deeper. FOSSIL REMAINS Skull and skeletal remains.
ANATOMICAL CHARACTERISTICS Coronoid process
prominent, teeth numerous.
AGE Middle Jurassic; late Callovian.
Dakosaurus DISTRIBUTION AND FORMATIONS Northern France;
andiniensis Marnes de Dives.
NOTES May include Metriorhynchus moreli. Shared its
habitat with Proexochokefalos.
Thalattosuchus
superciliosus
198

Thalattosuchus superciliosus
199

Thalattosuchus
superciliosus
Maledictosuchus riclaensis
4 m (13 ft) TL, 275 kg (600 lb)
RHACHEOSAURINES
FOSSIL REMAINS Majority of skull and minority of SMALL TO MEDIUM-SIZED METRIORHYNCHIDS
skeleton. FROM THE MIDDLE TO EARLY CRETACEOUS
ANATOMICAL CHARACTERISTICS Teeth not numerous. OF EUROPE AND SOUTH AMERICA
DISTRIBUTION AND FORMATIONS Northeastern ANATOMICAL CHARACTERISTICS Snouts long and fairly
Spain; Agreda. narrow, preorbital opening closed, lower jaws shallow,
HABITAT Continental shelf. teeth moderate in number. Foreflipper not clawed.
HABITAT Coastal.
HABITS Fishers of small to medium-sized game.
Maledictosuchus lack of sufficient sampling.
riclaensis
Rhacheosaurus gracilis
1.5 m (5 ft) TL, 10 kg (22 lb)
FOSSIL REMAINS A few skulls and skeleton.
ANATOMICAL CHARACTERISTICS Teeth rather small.
Gracilineustes leedsi AGE Late Jurassic; late early Tithonian.
4 m (13 ft) TL, 140 kg (300 lb) DISTRIBUTION AND FORMATIONS Southern Germany;
FOSSIL REMAINS Complete and partial skull, minority of Mörnsheim.
skeleton, severely flattened top to bottom. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Standard for group. NOTES Shared its habitat with Aeolodon, Geosaurus
AGE Middle Jurassic; middle Callovian. giganteus, Dakosaurus maximus, Cricosaurus elegans.
lower Oxford Clay.
NOTES Shared its habitat with Peloneustes, Rhacheosaurus
Pachycostasaurus, Simolestes, Liopleurodon, Cryptoclidus, gracilis
Muraenosaurus, Tricleidus, Ophthalmosaurus,
Tyrannoneustes, Suchodus.
200

Cricosaurus bambergensis AGE Late Jurassic; late Kimmeridgian.

1.6 m (5.2 ft) TL, 12 kg (25 lb) DISTRIBUTION AND FORMATIONS Southern Germany;
FOSSIL REMAINS Skull and skeleton, severely flattened upper Torleite.
top to bottom. HABITAT Island archipelago shallows.
ANATOMICAL CHARACTERISTICS Teeth fairly large. NOTES Although this species and C. suevicus lived at same
AGE Late Jurassic; early late Kimmeridgian. time and in same region, they have not been found
DISTRIBUTION AND FORMATIONS Southern Germany; together in same location.
lower Torleite.
HABITAT Island archipelago shallows. Cricosaurus suevicus
NOTES May the direct ancestor of one of the German 2 m (6.5 ft) TL, 22 kg (50 lb)
species below. FOSSIL REMAINS Skull and skeleton.
ANATOMICAL CHARACTERISTICS Teeth rather small.
Cricosaurus albersdoerferi Trunk slender. Tail and lower tail lobe fairly long.
2.1 m (7 ft) TL, 28 kg (60 lb) AGE Late Jurassic; late Kimmeridgian.
FOSSIL REMAINS Skull and skeleton. DISTRIBUTION AND FORMATIONS Southern Germany;
ANATOMICAL CHARACTERISTICS Teeth moderate in Nusplingen.
size. Trunk not as slender. Tail and lower lobe not as HABITAT Island archipelago shallows.
long.
Cricosaurus albersdoerferi
Cricosaurus suevicus
201

Cricosaurus elegans Cricosaurus vignaudi

3 m (10 ft) TL, 80 kg (180 lb) 3 m (10 ft) TL, 80 kg (180 lb)
FOSSIL REMAINS Majority of skull, partial skull. FOSSIL REMAINS Majority of skull and minority of
ANATOMICAL CHARACTERISTICS Teeth fairly large. skeleton.
AGE Late Jurassic; late early Tithonian. ANATOMICAL CHARACTERISTICS Insufficient
DISTRIBUTION AND FORMATIONS Southern Germany; information.
Mörnsheim. AGE Late Jurassic; middle Tithonian.
HABITAT Island archipelago shallows. DISTRIBUTION AND FORMATIONS East-central Mexico;
NOTES C. rauhuti may be adult of this species. Shared its upper Pimienta.
habitat with Aeolodon priscus, Geosaurus giganteus, HABITAT Continental shallows.
Dakosaurus maximus, Rhacheosaurus.
Cricosaurus? puelchorum
1.5 m (5 ft) TL, 10 kg (22 lb)
Cricosaurus ANATOMICAL CHARACTERISTICS Teeth fairly large.
araucanensis AGE Early Cretaceous; lower Berriasian.
upper Vaca Muerta.
NOTES Placement of poorly preserved specimen in earlier
Cricosaurus araucanensis Cricosaurus is not certain. Shared its habitat with
3 m (10 ft) TL, 80 kg (180 lb) Purranisaurus potens.
FOSSIL REMAINS Skulls and partial skulls and skeleton.
ANATOMICAL CHARACTERISTICS Snout elongated, Enaliosuchus schroederi
teeth fairly large. 1.5 m (5 ft) TL, 10 kg (22 lb)
AGE Late Jurassic; early Tithonian. FOSSIL REMAINS Majority of skull and minority of
DISTRIBUTION AND FORMATIONS Western Argentina; skeleton.
lower Vaca Muerta. ANATOMICAL CHARACTERISTICS Teeth fairly large.
HABITAT Continental shallows. AGE Early Cretaceous; early Valanginian.
NOTES May include C. lithographicus. Shared its habitat DISTRIBUTION AND FORMATIONS Southern Germany;
with Pliosaurus? patagonicus, Caypullisaurus. Stadthagen.
NOTES Shared its habitat with Lagenanectes, Enaliosuchus.
202

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D I A P S I DS

SMALL TO GIGANTIC TETRAPODS FROM THE

ANATOMICAL CHARACTERISTICS Extremely variable. Two temporal

02 MMR – Field Guide pp066-208.indd 67 16/05/2022 14:13

HABITAT AND HABITS Extremely variable, marine to fully terrestrial

ANATOMICAL CHARACTERISTICS Fairly uniform. Skulls ANATOMICAL CHARACTERISTICS Necks moderate in

ANATOMICAL CHARACTERISTICS Heads subtriangular,

02 MMR – Field Guide pp066-208.indd 68 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 69 16/05/2022 14:13

Anshunsaurus wushaensis DISTRIBUTION AND FORMATIONS Southeast China;

02 MMR – Field Guide pp066-208.indd 70 16/05/2022 14:13

ANATOMICAL CHARACTERISTICS Heads short and Xinpusaurus? xingyiensis

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Concavispina biseridens Agkistrognathus campbelli

02 MMR – Field Guide pp066-208.indd 72 16/05/2022 14:13

AGE Early or Middle Triassic. Clarazia schinzi

02 MMR – Field Guide pp066-208.indd 73 16/05/2022 14:13

ANATOMICAL CHARACTERISTICS Highly variable.

ATOPODENTATIANS ANATOMICAL CHARACTERISTICS Highly variable.

ANATOMICAL CHARACTERISTICS Heads small, snout

02 MMR – Field Guide pp066-208.indd 74 16/05/2022 14:13

Paraplacodus muscle study

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02 MMR – Field Guide pp066-208.indd 76 16/05/2022 14:13

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Cyamodus rostratus Cyamodus kuhnschnyderi

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Shared its habitat with Anshunsaurus huangguoshuensis, PLACOCHELYIDS

Unnamed genus? orientalis

02 MMR – Field Guide pp066-208.indd 79 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 80 16/05/2022 14:13

HABITS Poor locomotion both in water and onshore.

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HABITAT Continental nearshore. Neusticosaurus pusillus

Neusticosaurus (or Pachypleurosaurus) edwardsi

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Neusticosaurus peyeri ANATOMICAL CHARACTERISTICS Head small, fairly

02 MMR – Field Guide pp066-208.indd 85 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 86 16/05/2022 14:13

AGE Middle Triassic; middle Anisian. SIMOSAURIDS

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PAXPLESIOSAURS DISTRIBUTION AND FORMATIONS Nevada; upper Favret.

PAXPLESIOSAUR MISCELLANEA ANATOMICAL CHARACTERISTICS Head medium sized,

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SMALL TO GIGANTIC PAXPLESIOSAURS FROM Pistosaurus longaevus

Microcleidus muscle study

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02 MMR – Field Guide pp066-208.indd 92 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 93 16/05/2022 14:13

DISTRIBUTION AND FORMATIONS Northwestern Macroplata tenuiceps

02 MMR – Field Guide pp066-208.indd 94 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 95 16/05/2022 14:13

02 MMR – Field Guide pp066-208.indd 96 16/05/2022 14:13

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Gallardosaurus iturraldei Simolestes? keileni

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Simolestes vorax NOTES Shared its habitat with Peloneustes, Pachycostasaurus,

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02 MMR – Field Guide pp066-208.indd 101 16/05/2022 14:13