Palaeont. afr., 21, 143-159 (1978) 

PERMO-TRIASSIC "LIZARDS" FROM THE KAROO SYSTEM 

PART II 

A GLIDING REPTILE FROM THE UPPER PERMIAN OF MADAGASCAR 

by 

Robert L. Carroll 

Redpalh Museum, McGill University, Montreal, P.Q, Canada, HJA 2K6. 

ABSTRACT 
DaedaLosaurus madagascariemis, gen. et sp. nov. from the Upper Permian of Madagascar is a 

small reptile in which the trunk ribs are greatly elongated to support a gliding "membrane" 
similar to those in the Upper Triassic lizards Kuehneosaurus and lcarosaurus, and the living agamid 
Draco. The membrane is supported by 21 pairs of ribs compared with five to seven in Draco, ten 
in l carmauTUs and II in Kuehneosaurus. The total body mass is estimated as 250 grams, the area of 
the membrane nearly 200 cm2, with a wing loading of approximately 1,25 glcm2. A second 
species in the fauna, belonging to the same family, Coelurosauravus elivemis Piveteau, has a very 
similar appendicular skeleton, but ribs of normal proportions. The maxillary dentition of Coelu­
rosauravus is acrodont, that of DaedaLosaurus pleurodont. In neither genus is the temporal region 
of the skull adequately known, although the configuration of the jugal in Daedalosaurus suggests 
that the lower temporal bar may be reduced. The primitive nature of the appendicular skeleton, 
with little evidence or the specializations seen in contemporary lizards, suggests that these genera 
should not be classified among the Squamata, but among the Eosuchia. 

CONTENTS 
Page 

INTRODUCTION ........................................................................................................................................... 143 
COELUROSAURAVUS ELIVENSIS PIVETEAU .................................... .... ....... ...................... ..... ....................... 144 
DAEDALOSAURUS MADAGASCARIENSIS, gen. et sp. nov .................................................................................... 149 

Skull .. ...... .. ... ...... .. ........ .. .... ....... .. ..... .. ..... ....... ..... .. ....... ........ .. ................ ... ....... ..................................... ...... 151 
Vertebrae ..................................................................................................................................................... 151 
Ribs .............................................................................................................................................................. 154 
Appendicular skeleton ................................................................................................................................... 155 
Flight characteristics and comparison with other gliding genera ....................................................................... 156 

THE TAXONOMIC POSITION OF COELUROSAURAVUS AND DAEDALOSAURUS ............................................ 158 
ACKNOWLEDGEMENTS ................................................................................................................................ 159 
REFERENCES .................. ... .................. ...... ...... .. .... ... .... .. ... .. ............ .... ..... ........ ........... ........ ... .. ..................... 159 

143 

INTRODUCTION 
Knowledge of Late Permian and Early Triassic 

reptiles has come primarily from the Karoo age beds 
of South Africa and Russia. The fauna from these 
deposits consists almost entirely of therapsids, with 
diapsids as relatively rare elements. Reptiles of this 
age have also been described from Madagascar (Pi­
veteau, 1926). Because of the difficulty of prepara­
tion and the lack of comparative material then 
available, the significance of this fauna has not been 
generally appreciated. Development of new tech­
niques of preparation and casting reveals an exten­
sive and varied fauna, quite different from that 
known elsewhere, consisting almost entirely of diap­
sids. 

Karoo age reptiles are known primarily from de­
posits exposed as a narrow band running north and 
south along the west side of the crystalline massif 
that makes up much of the eastern two-thirds of the 
island (fig. 1). Most of the material described by Pi­
veteau, including all the specimens discussed in this 
paper, came from the upper course of the Sakamena 
River, near Mount Eliva in the southwestern part of 
the island. All the vertebrates described from this 
area occur in nodules, and are typically found 
weathered free from the surrounding matrix. 

In view of the significance of the fauna, made up 
of many genera as yet known only from Madagascar, 
it is important to establish the age of the beds with 
as much assurance as possible. Piveteau (1926) attri-



144 

Figure I. Map showing approximate extent of the Lower Sa­
kamena Formation in south-western Madagascar 
and localities /Tom which Upper Permian reptiles 
have been described. 

buted the reptilian fauna of Mount Eliva to the 
Upper Permian, primarily on the basis of assumed 
relationships to South African genera, and relative 
levels of evolution. Although concepts of the rela­
tionships of early diapsids have changed consider­
ably since his determination, stratigraphic work and 
comparison of other fossils tend to support his esti­
mate of the age of the beds. Stratigraphic work re­
ported by Besairie (1972) places the beds which 
contain these vertebrates in the Lower Sakamena 
Formation. Farther north, this formation is overlain 
by beds containing the same genera as those of the 
extremely well-known Eotriassic fish fauna in the 
north of the island, e.g. Boreosomus and Birgeria 
(Lehman et ai., 1959). 

Recent palynological work by Goubin (1965) 
on numerous localities in Madagascar divides the 
Lower Sakamena into three zones, all of which are 

indicated as Upper Permian. The terrestrial verte­
brates occur in the uppermost of the three zones rec­
ognized by Goubin, and so at the very top of the 
Permian. Hart (969) correlates the entire Lower Sa­
kamena, typified by members of the Striatiti pollen 
zone, with the Russian Tatarian. 

In addition to the terrestrial vertebrates of the 
Mount Eliva fauna, Priem (924) described several 
fish specimens as members of a new species, Atherston­
ia colcanapi. According to recent study of this ma­
terial by Gardner (pers. comm.) the fish accord well 
with other species of Atherstonia from the Upper Per­
mian; members of this genus are not known in later 
horizons. Although perhaps less important as strati­
graphic indicators, the mega-plant material from 
these beds should be cited as well. Beneath the verte­
brate bed is a rich flora, dominated by Thinnjeldia 
and Glossopteris. Within the vertebrate bearing nod­
ules are also impressions of Glossopteris indica (Car­
pentier, 1936). 

Most of the specimens from the area of Mount 
Eliva have been attributed to the aquatic eosuchian 
genera Tangasaurus and Hovasaurus. There are also 
three more or less complete skeletons of small li ­
zard-like forms, well ossified, with slender limbs and 
without evident aquatic adaptations. Two of these 
were illustrated by Piveteau, and described as a new 
genus and species of coelurosaur, Coelurosauravus eli­
vensis. The third is of the same general size and limb 
proportions, but has greatly elongate ribs, resem­
bling those of the Upper Triassic gliding lizards 
Kuehneosaurus (Robinson, 1962) and Icarosaurus (Col­
bert, 1970). In none of the Madagascar specimens 
is the skull well preserved, but the dentition is very 
different in the two genera. Coelurosauravus has a 
distinctly acrodont attachment of the posterior max­
illary teeth, which are few in number and widely 
spaced. The form with long ribs has a subpleur­
odont implantation, with a large number of slender 
teeth. Although the two forms clearly belong to dis­
tinct genera, the similarity of the appendicular ske­
leton suggests that they may belong to the same 
family. Both are clearly distinct from most other 
Late Permian or Early Triassic reptiles including the 
primitive lizards from the South African Karoo beds 
(Carroll, 1975). The apparent close relationship of 
the two Madagascan genera makes it useful to con­
sider them together. 

COELUROSAURA VUS ELIVENSIS PIVETEAU 

The two specimens of Coelurosauravus described by 
Piveteau are numbered 1908-11-21a (the type, fig. 
2 and 1908-11-22a, fig. 3) in the collection of the 
Institute of Palaeontology, National Museum of 
Natural History, Paris. He recognized as belonging 
to the same genus a disarticulated maxilla and other 
skull bones in the same block as a skeleton of the 
genus with long ribs 0908-5-2; fig. 5). The first two 
specimens show the remains of almost complete ske­
letons, preserved in nodules exhibiting a clearly 



concentric structure in cross-section. The bone had 
entirely weathered out prior to collection. The sur­
face has since been cleaned by washing and picking 
out mud from the deeper cavities with a needle. The 
matrix is rather coarse-grained and some areas 
badly weathered, so that surface detail tends to be 

11 

/~ 
./ :.::\.'. 

145 

poor. The specimens have been studied through the 
use of high fidelity latex casts. 

In none of the specimens is the cranium ade­
quately preserved. All three show the maxilla, but the 
remainder of the skull appears to have been much 
less stoutly constructed, and much had disintegrated 
prior to preservation. The only bones of the skull 
roof to be seen clearly are the parietal and frontal, 
visible in the counterpart blocks of 1908-5-2, 
adjacent to the maxilla. What appears at first glance 
to be a very elongate parietal is apparently the leti: 
frontal overlapping the right parietal, judging from 
the configuration of these bones in the primitive li­
zard Paliguana (Carroll, 1975). The frontal is an ob-

1cm 

Figure 2. Type of CoelUTOsauravUJ elivemis. Institut de Palt~ontologie, Paris, no. 1908-11-21a. Numbers indicate approximate posi­
tion of presacral vertebrae and distal tarsals. Roman numerals indicate the number of metatarsals. a, astragalus; cal, cal­
caneum; cen, centrale; ec f, ectepicondylar foramen; en f, entepicondylar foramen; F, femur; Fi, fibula; H, humerus; m, 
maxilla; pm, premaxilla; R radius; T, tibia; U, ulna. X 2. 



146 

long bone, with a strong longitudinal ridge 
separating the orbital margin from the dorsolateral 
margin of the braincase. A groove on the anterola­
teral margin of the frontal may be interpreted as the 
position of prefrontal attachment. The area where 
the postfrontal might be expected to have attached is 
obscured where the frontal overlaps the antero­
medial margin of the parietal. 

The parietal resembles those of both Youngina 
(Cow, 1975) and Paliguana in being short, with a 
prominent lateral process extending behind the dor­
sal temporal opening, and a large parietal foramen 
midway along the medial margin. The postero­
lateral margin is recessed in the area of the tabular 
and postparietal. The ventral surface is strongly con­
cave. The configuration of the frontal and parietal in 
younginids and paliguanids is unfortunately so sim­
ilar that it is not possible, on the basis of these 
bones, to identiry to which group Coelurosauravus 
might be allied. 

Much of the skull may originally have been pre­
sent in 1908-11-21a, but only the tooth-bearing el­
ements are identifiable. The premaxilla shows a 
large margin for the external nares anteriorly (as the 
bone is oriented) with the posterior margin slanting 
posterodorsally. It is possible that this indicates an 
anteromedial position for the external nares as in 
Kuehneosaurus. Alternatively, this bone may have 
been reversed prior to fossilization. It shows three or 
four small, peg-like teeth, apparently quite unlike 
those of the maxilla. Maxillae are preserved in all 
three specimens. They appear to be very massive, 
and are peculiar in several respects. In most early li­
zard-like reptiles, the maxilla is very narrow posteri­
orly, but in these specimens, the posterior extremity 
is thick and ends abruptly in what appears to be a 
large surface for medial attachment, presumably 
with the ectopterygoid. The bones appear relatively 
short, although it is not certain whether any show 
the anterior margin. Each maxilla bears a small 
number of widely spaced triangular acrodont teeth. 
Seven is the maximum number preserved. The most 
anterior tooth in the type specimen appears to be set 
in a socket. The combination of peg-like premaxil­
lary teeth, a socketed anterior maxillary tooth and 
acrodont teeth in the remainder of the maxilla is 
strikingly close to the pattern common among living 
agamid lizards. One apparent difference is that in 
agamids, the jugal extends far anteriorly, bracing 
the medial surface of the maxilla nearly to the front 
of the orbit. In Coelurosauravus, the maxilla appears 
very high posteriorly, and apparently forms most of 
the ventral margin of the orbit. This does not neces­
sarily preclude a parallel anterior extension of the 
jugal, but there is no direct evidence for it. 

Oddly, no bone can assuredly be identified from 
the lower jaw. A splint-shaped element in 
1908-11-22a, associated with a bone which some­
what resembles a quadrate, might be an angular, 
sheathing a long retroarticular process. No further 
cranial elements can be recognized, and the specifi-

cation of this form as a "lizard" or eosuchian cannot 
be based on evidence of the temporal region. 

In 1908-11-22a most of the vertebral column is in 
place, extending from behind the skull in a tight cir­
cle, crossing over itself in the cervical region. Unfor­
tunately, the most anterior vertebrae are obscured in 
the area of the occiput, precluding positive identifi­
cation of the atlas or axis. Twenty-three vertebrae 
can be identified, almost certainly not including the 
sacrals. The first visible centrum is quite long, and 
so might be the axis but not the atlas, to judge by the 
relative length of these elements in most primitive 
reptiles. The last visible vertebra is hence probably 
the 24th presacral. In the type, the trunk vertebrae 
are less well displayed. The cervicals and anterior 
trunk vertebrae are too badly obscured to estimate 
their numbers. Several in the mid-region of the 
trunk are badly disarticulated. Only the posterior 
trunk, sacral and anterior caudal vertebrae are in 
articulation and relatively clearly exposed. Because 
of regional variation in the length of the centra, the 
relative position along the column may be corre­
lated in the two specimens. The vertebrae which may 
thus be identified as the 25th in the type bears a 
short slender rib. Posterior to it are five or six addi­
tional vertebrae anterior to the first haemal arch. 
None are well enough preserved to determine which 
are the sacral vertebrae. There were apparently be­
tween 25 and 27 presacral vertebrae. Judging by the 
pattern in nearly all primitive reptiles, there were 
presumably two sacral vertebrae. Nine to eleven 
caudal vertebrae are preserved. The tail was almost 
certainly considerably longer in the living animal. 

Throughout the column, the centra are longer 
relative to their width than in most other primitive 
reptiles. Equally striking is the change in vertebral 
length throughout the column. In Saurostemon and 
Palaeagama from the Upper Permian and Lower Tri­
assic of South Africa (Carroll, 1977 ) and the better­
known Jurassic lizards, the cervical vertebrae are rel­
atively short, but the length remains essentially con­
stant throughout the remainder of the column. In 
Coelurosauravus, the cervical vertebrae are not parti­
cularly short, but those in the posterior trunk region 
are considerably longer. This pattern may be seen in 
the gliding lizard lcarosaurus as well. The immedi­
ately presacral vertebrae are shorter, as are the sac­
rals and the most anterior caudals. More posteriorly, 
however, the caudals lengthen to approach the 
length of the longer trunk vertebrae. 

In the region of the carpus of the long-ribbed 
genus as preserved, are paired atlas arches and an 
axis which may pertain to Coelurosauravus . The 
paired nature of the atlas arch compares with eosu­
chians and sphenodontids, but contrasts with the 
median nature of this element in lizards, including 
the Lower Triassic genus Palaeagama (Carroll, 1975). 
Well-defined transverse processes suggest the pres­
ence of atlas ribs, absent in early lizards. The most 
posterior cervical and anterior trunk vertebrae have 
relatively high, rectangular neural spines; not elong-



147 

Figure 3. CoeluTOsauravUJ elivensis, Institut de Paleonto10gie, Paris, no. 1908-11-22a. Numbers indicate the approximate position ot 
presacral vertebrae. ang, angular; Cl, clavicle; Clei, cleithrum; en f, entepicondylar foramen; H, humerus; m, maxilla; R, 
radius; S, scapulocoracoid; U, ulna. X 2. 

ate anteroposteriorly as are those of lcaroJaurus, but 
clearly much more prominent than in most small 
primitive lizards. lbe spines become more triangular 
in shape posteriorly. 

The transverse processes of the fourth, fifth and 
sixth vertebrae extend laterally to an appreciable ex­
tent and have a long anteroventrally sloping artic­
ulating surface. As well as can be seen, those of the 

anterior seven or eight vertebrae are not signifi­
cantly different from those of other early lizard-like 
forms. Most posteriorly, the articulating surfaces 
appear to be much shorter, and to shift posteriorly 
toward the middle of the elongate pedicle. Just an­
terior to the sacrum, they revert to a more anterior 
position. 

In the caudal series, the elongate anterior trans-



148 

verse processes common to eosuchians and lizards 
are not visible. Normal haemal arches are evident in 
the anterior portion of the tail. Small crescentic in­
tercentra can be seen anterior to vertebrae four and 
five in 1908-11-22a. 

Ribs are present throughout most of the trunk 
region in 1908-11-22a. Little can be seen of those 
that would have been attached to the cervical verte­
brae, although the size of the transverse processes 
indicates that the heads would have been substan­
tial. It is possible that the short ribs opposite verte­
brae seven through nine are displaced from the 
cervical region. In 1908-11-21a, ribs in the anterior 
trunk are much longer and appear wide. Ribs asso­
ciated with vertebrae ten through 16 are more or less 
in position in 1908-11-22a. Except for being some­
what expanded towards the distal end (perhaps as a 
result of crushing), they are of typical size and confi­
guration seen in other small reptiles of this age. Ribs 
associated with the more posterior trunk vertebrae 
appear considerably smaller. In general, the ribs as­
sociated with this specimen can be said to be normal 
for a primitive reptile. One much longer rib accom­
panies 1908-11-21a. It is approximately the length 
of the femur, it is not in articulation with any verte­
bra, and is therefore possibly from another spec­
imen. The loss of several trunk vertebrae and most 
of the ribs in the central portion of the trunk makes 
it difficult, however, to rule out the presence of 
longer ribs in this region. 

The shoulder girdle is barely visible in the type. In 
1908-11-22a, however, much of the structure can be 
made out. The scapulocoracoid appears as a single 
ossification, with a large screw-shaped glenoid of 
generally primitive configuration. The posterior 
articulating surface is oriented vertically, suggesting 
that the humerus had more freedom to angle ven­
trally than in captorhinomorphs and pelycosaurs. 
The scapula appears tall and narrow, although the 
anterior margin cannot be determined. The inter­
clavicle is not evident. The narrow blades of both 
clavicles can be seen, but not the extent of the stems. 
A bone adjacent to the blade of the scapula may be a 
remnant of the cleithrum. If so, this bone is rela­
tively massive. There is no evidence of an ossified 
sternum. 

The humerus, exposed in both skeletons, is a long 
narrow bone, with the ends twisted in a primitive 
fashion. The proximal portion shows a well-defined 
narrow strap-shaped articulating surface. Except for 
proportional differences related to the relative size 
of the head of the humerus, it resembles the pattern 
seen in pelycosaurs and captorhinomorphs 
(Holmes, 1977) and does not show evidence of the 
inception of a lacertoid pattern, such as is noted in 
the contemporary South African lizard Saurostemon 
(Carroll, 1977). The articulating surface is somewhat 
bulbous, however, indicating less restriction of move­
ment at this joint. The shaft is long and slim. The 
distal extremity is flattened. At the very posterior 
margin of the bone is a narrow slit of the entepicon-

A 

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Pi 

Figure 4. Limb elements or CoelurosauravUJ and DaedalosauTUJ. 
A, Ventral view of humerus o f Daedalosaurus mada­
galcariensiJ. B, Ventral view o f hand and associated 
ulna and radius or Daedalosaurus madOf!,ascariensis. C, 
Ventral view or lower rear limb of CoelurosauravUJ eli ­
vensis. All X 2. 1-5, distal carpals and tarsa ls ; a, as­
tragal us; ca l, calcaneum; cap , capitellum ; cen, 
centrale; ec C ectepicondylar foramen ; en f, entepi­
condylar 10 r<ll11('n ; I, il1lC'rmedium ; Lc, la teral cen­
trale; Mc, medial centrale ; Pi, pisiform; Ra, radiale; 
II', trochlea ; Ul, ulnarC'. 

dylar foramen. Anteriorly, there appears to be a 
small ectepicondylar foramen. Dorsally, approxi­
mately midway in the width of the bone is a slightly 
raised area for articulating with the olecranon. The 
surface of the humerus is distinctly recessed proxi­
mally and posterior to this area. The ventral surface 
of the distal extremity is not exposed. 



The radius is a narrow cylindrical bone, 62 per 
cent the length of the humerus. Distally, the bone is 
somewhat compressed in the plane of the carpus. 
The articulating surface is not well exposed. The 
total length of the ulna is 73 per cent that of the hu­
merus, with a very well-developed olecranon, the tip 
of which shows a distinct notch for the insertion of 
the triceps. The proximal end of the bone is 
markedly compressed in an anteroposterior direc­
tion; the distal end of the shaft narrows considerably 
above the area of distal expansion. The distal end of 
the bone does not appear to be elongated as a 
specialized epiphyseal articulating surface as in Sau­
rostemon. 

The carpus is not exposed. Metacarpals and pha­
langes are preserved in 1908-11-22a but the hand is 
too incomplete to be restored without reference to 
other primitive reptiles. One digit shows three quite 
long phalanges. Other phalanges and possibly meta­
carpals can be seen in the type specimen, beyond 
the distal end of the humerus. 

149 

angular. Proximally, it appears to be overriden by 
the lateral margin of the astragalus. Just beneath this 
area is apparently the margin of the perforating fo­
ramen. The astragalus has a prominent articulating 
surface on the lateral margin for the tibia. The cen­
trale is a small bone largely obscured by overlying 
elements. The area of the fifth distal tarsal is poorly 
exposed, but there is no reason to think that this el­
ement is missing. Distal tarsals two, three and four 
are relatively well exposed, but the third and fourth 
appear broken, so that their original outline is diffi­
cult to restore exactly. The first distal tarsal is not 
exposed, but was probably bent under the other el­
ements. Distal tarsals two and three are unusually 
long relative to their width. The metatarsals are all 
in place, although the proximal end of the first is 
obscured, and the second is underneath the third 
and fourth . The fifth, as in more primitive reptiles 
but in contrast with paliguanids, is nearly as long as 
the fourth, and shows no evidence of even incipient 
hooking. It does, however, articulate with the fourth 

TABLE 1 
Dimension of appendicular skeleton of CoeluTOsauravus and Daedalosaurus (in cm) 

Humerus Ulna Ulna Radius Femur Tibia Fibula 

(total) (minus olecranon) 
C oelurosauravU.l 
Type 1908-11-21a 29 - -

1908-11-22a 31,5 23 20 

DaedaloJaurus 29,5 22 19,5 

In 1908-11-21a, an ill-defined area probably rep­
resenting the puboischiadic plate can be seen near 
the head of the femur, adjacent to the area of tran­
sition between the trunk and caudal vertebrae. No 
details can be seen. Most of the hind limb can be 
seen in this sp~cimen. The femur is approximately 
three per cent longer than the humerus, and has a 
somewhat thicker shaft. Details are obscured, but 
the bone can be interpreted as rather lizard-like in 
appearance, with a slightly sigmoidal curvature of 
the shaft and with the distal surface twisted antero­
ventrally. There appears to be a prominent internal 
trochanter arising a short distance distal to the head. 
There is certainly no trace of the adductor crest that 
distinguishes the ventral surface of the femur in 
many primitive tetrapods. The tibia and fibula are of 
nearly equal length, 66 per cent that of the femur. 
Both bones are lightly built with particularly narrow 
shafts. The distal end of the fibula is not greatly ex­
panded. 

The foot (fig. 4) is exposed in ventral view, except 
for the third metatarsal which has rolled over to ex­
pose its dorsal surface. The definition of the individ­
ual elements is not especially good, the nature of the 
articulating surfaces poorly defined . The tarsus ap­
pears considerably more primitive than that of Sau­
_rostemon, with the ankle as a whole quite long 
relative to its width. The calcaneum is vaguely rect-

- 31 20 20± 
20± - - -

19,5 31 20 -

as well as the fifth distal tarsal. One or more pha­
langes from each of the digits are preserved. There is 
only a single terminal phalanx, displaced but associ­
ated with the first digit because of its short distance 
from the tarsus. It is short, sharply pointed and re­
curved. Three phalanges of the fifth digit are in 
articulation and two from the second, the last of 
which are the longest. A phalangeal count of two, 
three, four, five, four can be confidently restored. 

Few, if any, ventral dermal scales are evident in 
either of the skeletons. 

DAEDALOSAURUS gen. nov. 

Type speCIes, Daedalosaurus madagascariensis 

Diagnosis - Primitive reptile, apparently classifiable 
within the order Eosuchia, with trunk ribs nine 
through 26 greatly elongate to support a gliding 
membrane. Approximately 29 presacral vertebrae, 
transverse processes not greatly elongate. Ribs four 
through eight supporting thoracic cavity. Dentition 
pleurodont. Tall narrow scapula, not fenestrate an­
teriorly. Large cleithrum retaining dorsal blade. 
Sternum apparently unossified. Iliac blade ex­
panded anteriorly to form a triangular plate. No 
evidence of ossified epiphyses. 



150 

Figure 5. Daedalosaurus madagascariensis, type, n. gen. et n. sp. Institut de Pah~ontologie, no. 1908-5-2. Photograph of specimen in 
which the bone is represented largely as a natural cast. The portion including the skull and end of the tail is from the 
counterpart of the remainder of the specimen and is printed in reverse. Approximately X 1. 





DAEDALOSAURUS MADAGASCARIENSIS sp. nov. 

Diagnosis - same as for genus. 
Type specimen - Nearly complete skeleton preserved 
in contiguous blocks numbered 1908-5-2 and 1908-
11-21 and unnumbered, partial counterpart. Insti­
tut de Paleontologie, Museum National d'Histoire 
Naturelle, Paris. 
HonLOn - Upper portion of the Lower Sakamena 
Formation, uppermost Permian. 
Locality - Vicinity of Mount Eliva, upper reaches of 
the Sakamena River Valley, South-western Mada­
gascar. 
Etymology - Generic name in reference to Dae­
dalus; according to Greek legend, the inventor of 
wings and the father of Icarus. The specific name 
refers to the island on which the fossil was found. 

Skull 
The skull can be seen on a small slab that was ap­

parently the only portion to be collected of the coun­
terpart of the block in which most of the skeleton is 
visible (figs. 5 and 7.) The orientation can be deter­
mined by the caudal vertebrae and the ends of three 
rit-s that are preserved in both blocks. Skull el­
ements of Coelurosauravus are also preserved as coun­
terparts in both blocks. The skull of Daedalosaurus 
appears to be displaced only slightly from its normal 
position relative to the cervical vertebrae. The pos­
terior portion was unfortunately broken off, appar­
ently at the time of collection to judge by the relative 
freshness of the break. The central and anterior part 
of the skull roof is visible in ventral view, together 
with disarticulated elements of the palate and cheek. 
Unfortunately the surface of the block was weath­
ered so that details of the bone surface are not clear; 
identity of the disarticulated elements is thus uncer­
tain. 

The right pre- and postfrontals, parietal, frontal 
and possibly a portion of the nasal are displayed. 
The upper temporal opening appears to be very 
lateral in position relative to the midline established 
by the frontal. This may be partially the result of 
flattening of the originally highly arched parietal 
into the plane of the rock, but probably also from 
lateral displacement of the bone relative to the fron­
tal. The left maxilla is exposed medially. It resem­
bles that of primitive captorhinomorph reptiles. The 
bone is incomplete anteriorly. The portion preserved 
bears 17 peg-like teeth, with space for two others. 
The teeth are set in a strongly developed medial 
ridge. Like those of most primitive reptiles, they are 
subpleurodont, with the lateral surface of the max­
illa extending a short distance across their bases. 
Posteriorly the bone is notched for the jugal, which 
does not appear to have overlapped the bone exten­
sively. As the skull is tentatively reassembled (fig. 8), 
there appears to be a considerable gap between the 
maxilla and prefrontal, indicating the presence of an 
extensive lachrymal. A triradiate bone in the area of 
the left cheek is almost certainly the postorbital, of 
the shape common to paliguanids and younginids. 

BP - K 

151 

Beyond it is a thin bone which appears to be ~e 
right jugal in medial view and the adjacent portIon 
of the pterygoid and ectopterygoid, reversed back to 
front. The jugal shows no posterior ramus, but the 
manner of preservation is such that it might origi­
nally have been present and yet not be exposed. Me­
dially, the jugal shows a process for attachment to 
the ectopterygoid. The ectopterygoid is a stout, 
rounded bone, demonstrating the presence of a 
large suborbital fenestra in contrast to the plate-like 
ectopterygoid of captorhinomorphs. Farther away 
from the skull is a bone with limited exposure show­
ing a small area of unfinished surface which may 
represent the region of the pterygoid that had artic­
ulated with the parasphenoid. Extending ventrally 
from the parietal is a thin plate of bone which is 
probably the dorsal portion of the prootic but not 
enough is exposed for description. 

Although the skull roof looks reasonably com­
plete and some of the cheek elements are present, 
reliable reconstruction of the skull is not possible. 
There is nothing to compensate for the absence of 
the squamosal and quadrate in determining the na­
ture of the temporal region. The pattern could re­
semble that of such taxonomically distinct forms as 
Youngina, Pro lacerta or Paliguana. 

Vertebrae 
Most of the vertebral column is obscured by the 

ribs as preserved (recalling that the specimen is pre­
served largely as a natural cast, these elements were 
lost with the missing portion of the counterpart 
block). In the restoration, the trunk vertebrae are 
drawn on the basis of those of Coelurosauravus since 
the features of the appendicular skeleton are very 
similar in the two genera. It may be significant that a 
similar elongation of the mid-trunk vertebrae also 
occurs in Icarosaurus and may be associated with 
flight in that genus. 

Only the cervical, thoracic, sacral and caudal ver­
tebrae can be seen in Daedalosaurus. The main block 
is broken within the cervical series. Judging by its 
greater length, the first vertebra exposed is probably 
the axis. Nothing resembling an atlas is visible. The 
ribs of the first, second and third vertebra are short 
and single headed. This, together with their position 
anterior to the shoulder girdle, warrants designation 
of these vertebrae as cervicals. The next five verte­
brae bear longer, presumably ventrally directed, ribs 
that supported the walls of the thoracic cavity. Behind 
them were at least 18 vertebrae bearing elongate ribs 
extending laterally from the column. The total pre­
sacral vertebral count appears to be 29. There are 
two sacral vertebrae, and 29 caudals are preserved, 
although the original length of the tail was probably 
considerably greater. The vertebrae are all amphi­
coelous. 

Vertebrae two through five are relatively well pre­
served in lateral view. Their general proportions 
resemble those of Coelurosauravus. The centra are 
relatively long and the neural spines tall and square 



152 

prf 

1cm 

~; po 
~{., 

~j'" 
-.~ . 

.. • ., - pt 

~ . 

." ...... .! 

/'''. r.art f / ' .• ~ 
. .' " .. ~.~: "~ 
" ~..:.~J 

pt 

A 

15 20 25 

25 20 

B 
Figure 7. Daedalosaurus TlUUiagascariensis. A, Drawing of latex cast of the counterpart of the block illustrated in Figure 6. B, End of tail 

illustrated in Figure 6. Numbers indicate position of caudal vertebrae; art f, facet of pterygoid for articulation with para­
sphenoid; ect, ectopterygoid; f, frontal; j, jugal (medial view); m, maxilla; n, nasal; p, parietal; pf, postfrontal; po, post­
orbital; prf, prefrontal; pro, prootic; pt, pterygoid. Maxilla with acrodont dentition, frontal and parietal below long ribs 
attributed to CoeluTosauravus. X 2. 



in outline. The transverse process of the second ver­
tebra is not well preserved. The dorsal margin of the 
articulating surface of the third is oriented hori­
zontally, at the end of a narrow, laterally directed 
process that extends beyond the level of the 
zygapophyses. The zygapophyses of this and the 
other anterior vertebrae are large, with the articulat­
ing surfaces flat and tilted at approximately 15 de­
grees from the horizontal. Vertebrae four through 
seven show a pattern of buttressing and excavation 
which can be attributed to selective forces acting to 
maximize the strength and minimize the mass. The 
zygapophyses are connected by a lateral ridge above 
the level of the transverse processes. The processes 
themselves are supported by three converging 
ridges; two extend horizontally beneath the ridge 
connecting the zygapophyses, and a third extends 
obliquely anteroventrally. The articulating surface is 
restricted to a sma1l oval at the end of the process. 
The transverse process of the eighth vertebra has a 
more normal configuration for a primitive reptile. 
The articulating surface is more extensive, and 
angles anteroventrally. It is thickest posterodorsally, 
and thins anteriorly. All that is visible of the more 
posterior trunk vertebrae is a series of transverse 
processes protruding between the ribs. Those that 
may be attributed to vertebrae 13, 14 and 15 show 
an articulating surface that appears horizontal dor­
sally. It is thick anteriorly, and tapers to the rear. 
Apparently the transverse processes of the trunk ver­
tebrae were no longer than those more anterior in 
position, rather than being elongate as are those of 
Kuehneosaurus and [carosaurus. If the transverse pro­
cesses were long, it is likely that they would have 
been oriented in the same plane as the remainder of 
the skeleton, rather than at right angles to it, and 
hence would have been preserved. 

Three poorly-exposed vertebrae can be seen just 
anterior to the sacrum. Their centra are short, as are 
those of the vertebrae just anterior to the sacrum of 
Coelurosauravus and [carosaurus. The two sacral verte­
brae are incompletely exposed at the margin of the 
ilium, partially obscured by trunk ribs. The centra 
are short, and the neural spines low and rounded. 

Twenty-nine caudal vertebrae can be seen in se­
quence behind the sacrum. The first three are short, 
as are the sacrals, the centrum of the second approx-

\ 
I 
I 

" 

153 

Figure 8. Restoration of DaedaLosauruJ madagascariensis skull 1I1 

lateral view, based on disarticulated elements. X 2. 

imately 3,7 mm in length., The length increases 
over the next three, after which the length is nearly 
constant (centra approximately 5 mm in length) to 
the end of the preserved series. The neural arches 
gradually decrease in height and width. The rre­
served portion of the tail is nearly the length 0 the 
presacral column, but its original length may have 
been much greater. A striking feature of the tail is 
the apparent absence of long ribs fused to the trans­
verse processes of the anterior vertebrae, a feature of 
both eosuchians and most lizards. This may be a re­
sult of the relatively poor preservation and difficulty 
of preparation in this area, but if these h4d been 
prominent features, it would be expected that they 
would be preserved in the plane of the block. The 
vertebrae in this area are, however, visible in lateral 
view, rather than dorsoventrally. Caudal vertebrae 
two and three show small, ventrolaterally directed 
articulating surfaces, low down on the anterior sur­
face of the centrum, which may have accommodated 
unfused ribs, although none are visible. More pos­
teriorly, there appear to be neither transverse pro­
cesses nor ribs, although the vertebrae are well ex­
posed. Unfused caudal ribs are characteristic of 
captorhinomorphs but not eosuchians, suggesting 
that Daedalosaurus is primitive in this regard. This 
feature may, in contrast, be a specialization, for the 
anterior transverse processes in Draco are extremely 
short. This may be associated with the great flexibil­
ity of the tail in relationship with flight control (Col­
bert, 1967). [carosaurus, however, has more normal 
transverse processes in this area. 

TABLE 2 
Length of ribs supporting gliding membrane in DaedaLosauruJ (in em) 

Rib no . Length Rib no. Length Rib no. Length 

9 1,6 16 15,4 23 9,1 
10 2,0 17 16,0 24 7,5 
II 2,9 18 16,2 25 3,9 (6,1 ) 

12 4,2 19 15,8 26 4,7 
13 6,2 20 14,7 27 1,4 + 
14 8,8 21 12,0 28 1,1 
15 10,7 (II) 22 9,7 29 0,9 

Numbers in parentheses indicate probable original length of incompletely preserved ribs. 

BP - L 



154 

For most of the column the presence or absence 
of trunk intercentra cannot be determined. It is pos­
sible that normal crescentic intercentra are present 
in the cervical region, but preservation is not good 
enough to determine this for certain. For much of 
the caudal region, intercentral ossifications are ex­
posed. Only a few elongate haemal arches are ob­
served. Like the apparent reduction of the caudal 
ribs this may contribute to a more efficient tail shape 
correlated with gliding. 

Ribs 
Ribs may be associated with all of the presacral 

vertebrae. Short, single-headed ribs are visible adja­
cent to the vertebrae recognized as the second and 
third cervicals, and one more anterior rib is proba­
bly from the missing atlas. These ribs are approxi­
mately three-fourths the length of the centra and 
were apparently oriented almost straight laterally to 
judge by the configuration of the rib head and the 
articulating surface of the transverse process. 

Ribs lying just behind the shoulder girdle have 
been identified as belonging with vertebrae four and 
five. They are much longer, have cylindrical or nar­
rowly spatulate shafts, and the two heads are clearly 
separate. On the right side, ribs six, seven and eight 
are in articulation with their respective vertebrae. 
The lengths of the flattened shafts appear to increase 
progressively, although they are somewhat obscured 
by overlying elements. The width of the heads seems 
to narrow somewhat. It is probable that ribs four 
through eight extended ventrally to surround the 
chest cavity as in most primitive tetrapods. They are 
relatively long, to give a deep thoracic cavity. It is 
probable that they were continued ventrally in carti­
lage, and attached to a cartilagenous sternum, struc­
tures that are preserved in the contemporary 
(Permo-Triassic) paliguanid lizards. 

The orientation and configuration of the ribs 
changes dramatically behind the thoracic series, as 
in the Upper Triassic lizards Kuehneosaurus and Icaro­
saurus and the living agamid Draco; the trunk ribs 
were almost certainly elaborated to support a large 
gliding surface or "wing". 

From the ninth through 26th vertebra, the ribs of 
both left and right sides are, for the most part, ex­
tended laterally to the left of the column, those from 
the right side having been flipped over to expose 
their ventral surface. The ribs from the right side, 
for most of their length, overlie those from the left 
and appear to retain almost their normal position 
relative to one another, although the distance by 
which they are separated from each other in the fos­
sil is almost certainly less than it was in life. Recon­
struction of the original position and determination 
of the relative length of the ribs is extemely impor­
tant in this form, but complicated by several factors. 
Where the ribs from both sides overlap one another, 
it is sometimes difficult to determine whether a par­
ticular segment is from the left or right side. Several 
of the upper ribs as seen in the specimen are incom-

pletely exposed where they cross over the lower ribs 
and the sacral region, so that it is not always possible 
to associate proximal and distal portions of the 
same shaft. Ventral scales are plentiful in the spec­
imen, and are of similar configuration to the smaller 
ribs and the terminal portions of the larger ribs. 
Nevertheless, the preservation of the right ribs is 
sufficiently complete that there was relatively little 
difficulty in assembling the "wing". 

Proximally, ribs nine through 26 from the right 
side all appear to be in their natural order, al­
though a short gap separates numbers 20 and 21. The 
only serious difficulty in establishing the number 
and configuration lies with those most posterior in 
position. If it is assumed that all ribs lying more or 
less parallel to one another are from the right side, 
the count reaches 27, the most posterior of which is 
still very long. One of the posterior ribs, succeeding 
number 23, is angled slightly forward and beneath 
the others, suggesting that it may be from the left 
side. This suggests that the last long right rib present 
is the 26th. If three pairs of short presacral ribs (as 
seen on the left side) had originally been present, the 
presacral count would reach 29. That of Kuehneosau­
rus (Colbert, 1970, Table 1) is 28. 

In order to confirm the count on the right side, an 
attempt was made to determine the number present 
on the left, although they are much less well associ­
ated. None of the cervicals can be identified. Five 
disarticulated ribs, scattered through the trunk area 
(two of which have widely separated heads) may be 
from the thoracic region. None that are preserved 
can be identified with those making up the anterior 
margin of the right gliding surface: the ninth, tenth 
and 11 tho The 12th is only tentatively recognized. At 
least 15 other long ribs can be accounted for, al­
though their specific location cannot be ascertained. 
One group of seven parallels the longest ribs from 
the right side, numbers 16 through 20. The prox­
imal ends of these ribs are not exposed, however, so 
their total length cannot be established nor their 
original position specified. Two other ribs terminate 
with this group, but are displaced proximally. A 
second group of six originates at the level of the 
bases of ribs 19-21 on the right side. They extend 
anteriorly at an angle of nearly 90 degrees to the first 
group. They appear complete proximally. The three 
longest ribs are only slightly shorter than the longest 
from the right side. 

From the established pattern of the anterior ribs 
on the right side, and the number of ribs accounted 
for on the left side, it would appear that the last long 
left rib is associated with the 26th vertebra. Three 
ribs, the last two of which are definitely short, and 
the first of which is incomplete distally, lie .iust an­
terior to the pubis, at right angles to the other trunk 
ribs. Since they underlie surrounding elements, they 
may be from the left side. This would account for a 
total of 29 ribs. 

Except for ribs 21, 22 and 26 from the right side, 
the proximal ends appear complete. There is little if 





any differentiation of the head rrom the shaft and 
the articulating surfaces appear to be nearly flat. 
This surface appears to have been roughly oval in 
outline but this is difficult to ascertain due to crush­
ing. The lateral extremities of the ribs are cylindrical 
and gradually diminish to a point. Proximally, the 
shaft is crushed, obscuring the original configura­
tion. It is probable that this portion was originally a 
flattened oval, either hollow or filled with extremely 
cancellous bone. Crushing has resulted in the for­
mation of a uniform linear groove, extending for 
much of the length of the shaft. 

Most of the ribs appear essentially straight. Ribs 
nine through 12 are curved slightly at the tip. It is 
difficult to determine whether the curvature was pri­
marily posterior or ventral in the living animal. If 
ventral, it would have served to form a thick leading 
edge to the gliding surface, at least near the body. 
Similar curvature is seen in the 16th rib. Ribs 19 
through 26 appear to show gentle curvature 
throughout their length, although the tip remains 
straight. 

The tips of ribs 15 and 25 on the right side, and 27 
(from the left) are missing or obscured by other el­
ements. 

When the ribs making up the wing are assembled 
in their natural order a plausible pattern is estab­
lished, based on evidence from both sides. The 
length of ribs nine through 12 increases gradually. 
The length increases more rapidly from the 13th to 
16th. The 18th is apparently the longest of all, with a 
length of approximately 16,2 cm. The 19th and 20th 
gradually diminish in length. The 21st is notably 
shorter, the remainder progressively so. The length 
of the 27th rib is uncertain. The last two are much 
shorter, but appear to be oriented laterally, extend­
ing beyond the expected limit of the trunk. The 29th 
is curved slightly posteriorly. 

Appendicular skeleton 
In contrast to the South African paliguanids, the 

shoulder girdle of Daedalosaurus shows no speci­
fically lacertoid characteristics. It is surprisingly 
primitive in retaining a very large cleithrum. This 
bone is visible on both sides and shows a flattened 
dorsal portion extended posterodorsally over the 
margin of the scapula, as in primitive pelycosaurs. 
The shaft is a robust cylindrical rod, extending 
nearly to the level of the glenoid as preserved. The 
great length of the thoracic ribs, number four 
through eight, indicates the presence of a cartilage­
nous extrascapula, suggesting that the cleithrum 
had occupied a more dorsal position in life. The 
clavicle has a shorter stem, but with a somewhat 
widened blade that curved ventrally around the 
trunk. The interclavicle is not evident. The left half 
of the endochondral girdle is partially exposed in 
medial view beneath the anterior vertebrae. It ap­
pears to be ossified as a single unit. The scapular 
blade is very high and narrow, resembling that of 
primitive pelycosaurs. The coracoid area, seen 10 

155 

dorso-medial view on the left side, extends well be­
hind the scapula, suggesting a relatively long gle­
noid, although little of this structure can be seen in 
this specimen. None of the foramina characteristic 
of the scapulocoracoid in other early reptiles can be 
recognized. The anterior margin of the left side of 
the pectoral girdle is at the level of the anterior end 
of the third centrum; that on the right, approxi­
mately one segment more posterior in position. 
Judging by the configuration of the "wing" and the 
relative position of these structures in Draco, the gle­
noid was probably at the level of the fifth and sixth 
centra. 

There is no evidence of an ossified sternum;· a 
cartilaginous structure was probably present, however. 

The humerus is a long narrow bone, very similar 
to that of Coelurosauravus, with the ends twisted in a 
primitive fashion. The proximal articulating surface 
is long, narrow and clearly defined. It follows the 
pattern of primitive captorhinomorphs and pelyco­
saurs (Holmes, 1977) except for changes in propor­
tions. There is no evidence of the lacertoid 
specialization seen in Paliguana (Carroll, 1977). The 
great width of the glenoid and humeral articulation 
suggests a retention of the sprawled posture of 
primitive reptiles, rather than the raised forelimb 
common to lizards. The shaft is long and slim. At 
the posterior margin is a narrow entepicondylar fo­
ramen. Anteriorly, the area of the ectepicondylar 
foramen is somewhat damaged, but this opening 
appears to be as in other early diapsids. Articulating 
surfaces for the ulna and radius are extremely well 
defined. Ventrally, the capitellum appears as a 
hemisphere, somewhat elongated in the direction of 
the long axis of the humerus. Dorsally, the distal 
surface resembles that of Coelurosauravus. 

In as far as proportions can be determined and 
details of structure seen, the ulna and radius of Dae­
dalosaurus are identical with those of Coelurosauravus. 

The carpus of the right limb can be seen in ventral 
view. The elements are somewhat jumbled, but 
knowledge of the pattern common to captorhinids, 
eosuchians and primitive lizards allows identifica­
tion of all the bones. Their specific orientation and 
configuration is more difficult to establish. The ul­
nare is especially poorly exposed. The intermedium, 
radiale and ulnare have considerable areas of fin­
ished bone. The other elements show primarily un­
finished surface, making determination of the origin­
al orientation particularly difficult to determine. 
Judging from other primitive reptiles and modem 
lizards, the rounded appearance of the carpals is 
probably a result of incomplete ossification with the 
adult condition expected to show the pattern of a 
close fitting mosaic. The carpus appears much ad­
vanced from the configuration seen in the primitive 
diapsids Petrolacosaurus and Galesphyrus (Carroll, 
1976), in its relatively limited extent, both proximo­
distally and mediolaterally, but the specific configu­
ration of the bones does not approach the condition 
seen in lizards (Carroll, 1977). The proximal ends of 



156 

all four metacarpals are present. They overlap one 
another laterally when the carpus is restored to its 
natural width. The remainder of the hand is not pre­
served. In the restoration, it is shown with the pha­
langes of Coelurosauravus. 

The right half of the pelvic girdle is exposed in 
lateral view. Unlike that of most primitive lizards, 
but rather as in lcarosaurus, the iliac blade is broadly 
expanded above the acetabulum, especially anteri­
orly. This is surprising, in view of the limited degree 
of expansion of the sacral ribs. The pubis in lateral 
view appears very lacertoid, with the lateral expres­
sion of the pubic tubercle. It is tempting to restore a 
lizard-like thyroid fenestration, but this cannot be 
substantiated from the angle that the girdle is pre­
served. The pubic buttress of the acetabulum ex­
tends sharply laterally so as to limit the anterior 
swing of the femur. The ischium is largely obscured 
by the femur. 

The two femora are superposed and extend di­
rectly posteriorly from the girdle. Little can be seen, 
beyond the gross dimensions. Their length exceeds 
that of the humerus by only five per cent, in contrast 
to the relatively much longer femur in lcarosaurus 
and Kuehneosaurus. They are succeeded by slim tibiae 
65 per cent the length of the femur. Bone is present 
in the area of the tarsus, but the individual elements 
cannot be distinguished. Numerous phalanges and 
?metatarsals can be seen, some in articulation with 
each other and others scattered about. They are sim­
ilar to comparable elements in Coelurosauravus. Two 
articulated series compare with the fourth and fifth 
digits in that genus, although the individual el­
ements are thinner. 

No impressions of squamate-like epidermal scales 
are evident on the block, although they are pre­
served with some of the eosuchian specimens from 
this horizon. Many ventral dermal scales are present. 
In comparison with those of most primitive reptiles, 
they appear very long. It is probable that they served 
to reinforce the abdominal wall in the absence of 
normal ventrally directed ribs. Here, the outdated 
term "ventral ribs" seems quite appropriate. 

Flight characteristics and comparison with other gliding genera 
A total of 21 pairs of ribs support the gliding sur­

face in Daedalosaurus, a far greater number than that 
observed in other gliding lizards: five to seven in 
Draco, ten in Icarosaurus, eleven in Kuehneosaurus. The 
outline of the wing of Daedalosaurus appears inter­
mediate between that of lcarosaurus and Draco. Its 
lateral extent, relative to other body proportions 
resembles that of lcarosaurus, but it has a greater an­
teroposterior extent, as does that of Draco. Ribs are 
present on all the presacral vertebrae in Daedalosau­
rus, but the posterior portion of the wing in Draco, 
Icarosaurus and Kuehneosaurus is without bony sup­
port. 

The ribs appear to differ markedly from those of 
Draco in their stiffness or lack of inherent curvature. 

In the modem genus, the ligaments attached to the 
ends of the ribs are very important in influencing the 
shape and overall configuration of the wing. In Dae­
dalosaurus, as in Kuehneosaurus and lcarosaurus, the 
ribs appear to be the primary factor in defining the 
outline of the wing membrane. Restorations of the 
wing in all the gliding genera is shown in Figure 10. 

A major uncertainty in restoring the total area of 
the wing, and also the mass of the body, lies in the 
lack of direct information conceming the length of 
the posterior trunk vertebrae. Taking those of Coelu­
rosauravus as a guide, the total snout-vent length is 
approximately 20 cm and the wing area is just less 
than 200 cm2. Mass can be estimated on the basis 
of a graph given by Pough (1973), comparing snout­
vent length and body mass in modem lizards, as­
suming normal body proportions. On this basis, 
Daedalosaurus would have had a mass of approxi­
mately 250 grams. This does not take into consider­
ation the mass of the wing itself, but this may have 
been compensated for by lightening of other parts 
of the skeleton, as is suggested by the configuration 
of the cervical vertebrae. As in Draco, the flight sur­
face may have been augmented by other extensions 
of tissue, e.g. folds running along the side of the 
neck. Despite the much greater lateral extent of the 
membrane in Daedalosaurus, the wing load is very 
much higher, approximately 1,25 gr/cm2 compared 
with 0,23 gm/cm2 calculated for Icarosaurus and a 
range of 0,33 gm/cm2 to 0,76 gm/cm2 in Draco (Col­
bert, 1970). 

The greater wing load in Daedalosaurus does not in 
itself indicate that this genus was any less capable of 
gliding than Draco, since other factors greatly dimin­
ish the aerodynamic qualities in animals with very 
small wing surface. Colbert also cites an extreme 
amount of variability in relative wing size in Draco, 
suggesting that there appears to be no very critical 
mass to wing-area relationship. Wing load differ­
ences of up to 44 per cent are reported in animals of 
similar mass (Colbert, 1970). 

In Draco, the anterior margin of the wing is at­
tached ventral to the general surface of the mem­
brane, along the flank behind the forearm, thus 
providing a thick leading edge for the airfoil, at le~st 
near the body. The distal curvature of the first four 
ribs in Daedalosaurus, if ventrally oriented, may have 
served a similar purpose. Posteriorly, the wing in 
Draco attaches to the under surface of the thigh. The 
much longer posterior ribs in Daedalosaurus, if cap­
able of folding or bending backwards, would have 
seriously interfered with movement of the rear limb 
if similarly attached. The posterior margin is here 
restored as ending on the dorsal surface of the thigh. 

For Draco, folding the wings is relatively simple 
since they are not very long, and the posterior mar­
gin is rounded so that none of the ribs are long 
enough to extend beyond the base of the rear limb. _ 
If the ribs of Daedalosaurus, or for that matter, Icaro­
saurus or Kuehneosaurus, are folded posteriorly, they 



157 

Figure 10. Restoration of gliding reptiles. A, Kuehneosaurus, Upper Triassic, modified !Tom Robinson, printed in Romer (1966 ). B, 
lcaro.lauruJ, Upper Triassic, !i'om Colbert (1970). C, Daeda!o.IGurus, Upper Permian. D, Draco, living genus, Iwm Colbert 
(1967). All drawn to the same scale. 

pass above the rear limbs, well beyond the base of 
the tail. Colbert has suggested that in Icarosaurus, the 
close association of the bases of the ribs would have 
restricted posterior folding and necessitated some 
degree of dorsal flexure. In Daedalosaurus, the trans­
verse processes are all apparently short, as in Draco, 
and widely separated. The heads of the ribs are nar­
row but the nature of the articulating surfaces be­
tween the bones would have made folding in a 
posterior direction difficult. The transverse pro­
cesses of vertebrae in the mid trunk region are flat 
dorsally, with the articulating surface thickest anteri­
orly and sloping to a rounded point posteriorly. 
This shape is only slightly modifiea from the pattern 

in other primitive reptiles and does not seem well 
suited to a posterior hinging of the ribs, if the wing 
is folded back in the manner of Draco. One might 
suggest from the nature of the transverse process 
and rib articulation in Daedalosaurus that the rib 
hinged dorsally. There is, however, no likely epaxial 
musculature that would have made such movement 
possible, and the antagonistic action of rib depres­
sors would imply a totally improbable flapping 
flight. In Draco, the transverse processes, while not 
as long as those in Kuehneosaurus and lcarosaurus, ex­
tend well out from the arch and terminate in a 
rounded, evenly convex surface to which attached 
the end of the rib. Folding of the wings appears to 



158 

occur through both hinging at this joint and bend­
ing throughout the shaft of the ribs through the ac­
tion of the ligaments. Because of the great width of 
the wing in Daedalosaurus, it is almost impossible to 
conceive of its being held rigidly from the body 
when not in use as a gliding membrane, but this 
genus may have had only limited ability to change 
its configuration. 

Although it is difficult to see beyond the flight 
characteristics of the "wings" in these genera, an ex­
tensive vascularized area of tissue such as this could 
not fail to have a striking effect on heat regulation as 
well. The living Draco is reported to use the brightly 
coloured surface of the wings in courtship displays 
(Colbert, 1970). 

THE TAXONOMIC POSITION OF 
COELUROSAURA VUS 

AND DAEDALOSAURUS 

Despite the clearly-defined differences in the den­
tition and the striking specialization of the ribs in 
Daedalosaurus, this genus and Coelurosauravus may be 
allocated to the same family on the basis of close 
similarities in the appendicular skeleton. In size and 
proportions, the elements of the forelimb are nearly 
indistinguishable and quite different from those de­
scribed in other contemporary reptiles. To the ex­
tent that they are preserved in both genera, the 
pectoral girdle, vertebrae and rear limb are also 
similar. 

Romer (1968, p. 114) suggested the possibility 
that Coelurosauravus might be considered, along with 
Araeoscelis, as an early euryapsid, but admitted that 
adequate knowledge of the anatomy was lacking. 
Coelurosauravus differs from Araeoscelis in having a 
short rather than elongate neck, and in having acro­
dont rather than pleurodont teeth. There are other­
wise no anatomical features of Coelurosauravus that 
suggest close affinity with nothosaurs or other, more 
specialized, euryapsids. A review of other members 
of the Permian Madagascar fauna demonstrates the 
presence of several genera that are plausible ances­
tors of nothosaurs and plesiosaurs. They are clearly 
diapsid derivatives, but show no important similari­
ties to Coelurosauravus. 

The general configuration of the skeleton of 
Coelurosauravus and Daedalosaurus can be termed "li­
zard-like" in that they are small, with light bodies 
and long limbs. The skulls are poorly known but 
may well show the lacertoid pattern, with an upper 
temporal opening and a streptostylic quadrate. The 
pattern of the dentition in Coelurosauravus and the 
ribs in Daedalosaurus also emphasize lizard affinities, 
for both specializations may be seen in living mem­
bers of the Lacertilia. 

Comparison with other contemporary Permo­
Triassic lizards, the paliguanids (Carroll, 1975 and 
1977), demonstrates the absence of other structural 
and developmental aspects that appear basic to li­
zards as a group, however. Notably, Coelurosauravus 

and Daedalosaurus lack ossified epiphyses and 
specialized joint surfaces of the limbs. The sternum 
is unossified, and fenestration of the scapulocora­
coid is not developed. The retention of a large 
c1eithrum also seems a very primitive feature. On 
the basis of our current understanding of primitive 
lizards and the possible origin of this group, these 
genera seem more conveniently placed among the 
Eosuchia - which remains an ill-defined assem­
blage of primitive lineages. It is reasonable to think 
that Coelurosauravus and Daedalosaurus are among the 
descendants of a more primitive assemblage that 
also gave rise to the Paliguanidae and hence to 
modem lizards. In a broad adaptive sense, the lizard 
habitus may be thought of as beginning with the eosu­
chian or even captorhinomorph level of structural 
sophistication. Hence we see in these forms an es­
sentially modem type of dental and locomotor 
specialization together with very primitive features 
of the remainder of the skeleton. 

A further factor which should be considered in 
evaluatjng the taxonomic position of Daedalosaurus 
and Coelurosauravus is total body size. The esti­
mated body mass for both is in the range of 200 to 
300 grams. According to work by Pough (1973), 
primitive lizards rarely exceed a body mass of 
150 grams unless they evolve the ability to feed pri­
marily on plant material. Consequently most primi­
tive lizards, fossil and living, are small insectivores. 
Members of more advanced lizard groups have de­
veloped structural or metabolic specializations that 
permit active predation on animals other than in­
sects, and may reach a much larger body size. The 
relatively large body size of Daedalosaurus and Coelu­
rosauravus is unexpected among primitive lizards, 
and suggests a different feeding and metabolic strat­
egy. In common with most eosuchians which have a 
similar or even greater body size, they may have had 
a significantly lower metabolic rate, and correspon­
dingly smaller food requirements. The living Tua­
tara (Farlow, 1975) may provide a useful analogy of 
a non-squamate lepidosaur, with low food require­
ments. 

On the basis of the known material of Daedalosau­
rus and Coelurosauravus, it may be assumed that the 
similarities of the appendicular skeleton are the in­
heritance from a close common ancestor. It seems 
probable that the gliding membrane evolved ra­
pidly, with relatively little change in the remainder 
of the skeleton. All the trunk ribs are modified, in 
contrast with the greater regional specialization in 
the later genera, with fewer elongate ribs and the 
posterior trunk vertebrae without ribs at all. The 
vertebrae are excavated laterally, but the transverse 
processes are not elongate. 

It is possible that Coelurosauravus was adapted to 
an arboreal way of life, possibly including jumping 
from trees, that provided a structural pre-adapta­
tion to the pattern of Daedalosaurus. The elongation 
of the vertebral column and the deep thoracic 
region may fall into this category. 



It is natural to consider the possibility of relation­
ship between DaedalosauTUS and the late Triassic 
kuehneosaurids. Unfortunately, this is made diffi­
cult by the lack of good cranial material of Daedalo­
sauTUS and the limited information regarding the 
appendicular skeleton of Kuehneosaurus. On the basis 
of the available information, DaedalosaurUs seems 
more appropriately classified among the eosuchi­
ans. The skull of kuehneosaurids, on the other 
hand, unequivocally places them in the Lacertilia. 
The described elements of the appendicular skele­
ton, however, are generally more primitive than 
those of the Permo-Triassic paliguanid lizards. 
Some of the apparently primitive features, as the 
configuration of the endochondral shoulder girdle, 
may be specializations related to gliding. It is very 
difficult to evaluate whether such shared features are 
indicative of close relationship or to similarity in 
adaptation. Of the anatomical features known in all 
three genera, there are none that would, of them­
selves, preclude DaedalosauTUs having given rise to 
Kuehneosaurus and lcarosaurus . Such a derivation 
would imply, however, that most of the lacertoid 
features of the kuehneosaurids had evolved separ­
ately from those of the paliguanids and that the la­
certoid grade of evolution had been achieved 
independently in the two families . It seems slightly 
more likely that the gliding adaptations were devel­
oped separately in the kuehneosaurids and the 
coelurosauravids in the light of the presence of both 
gliding and non-gliding genera in the latter family. 

Except for the extreme specialization of the ribs in 
DaedalosauTUs and the specialization of the dentition 
in Coelurosauravus, these genera may be included 
among the eosuchians on the basis of the generally 
primitive pattern of the limbs and pectoral girdle. If 
the paliguanids are classified among the Squamata, 

159 

and the Prolactertiformes are no longer considered 
to be associated with the ancestry of lizards, the 
coelurosauravids may be the most lizard-like of the 
eosuchians. The presence of more specialized lacertoid 
features among the Upper Permian and Lower Tri­
assic paliguanids (Carroll, 1977) indicates that this 
group must have diverged from primitive eosuchi­
ans by the early or middle Permian. Consequently, 
the known coelurosauravids are certainly too late to 
be considered ancestral to lizards. They appear, 
however, to retain primitive features which help us 
to visualize the nature of the ancestral group. 

ACKNOWLEDGEMENTS 
I wish to thank Professor Piveteau for encouraging me to 

study the Permian reptiles of Madagascar. His pioneering work 
in this fauna revealed an otherwise unexpected diversity among 
early diapsids. Professor Lehman has been very helpful in pro­
viding space and other facilities at the Institut de Paleontologie. 
Mr. D. Goujet deserves special thanks for locating specimens 
and arranging lor technical assistance. The photograph was 
taken by Mr. R. Kandarown and Mr. D. Serrette. Pamela Gas­
kill's painstaking illustration of the specimens is greatly appreci­
ated . This study was an outgrowth of work on Karoo reptiles 
initiated in South Africa in 1972-73. The assistance of Drs. 
Cruickshank and Gow at the Bernard Price Institute and Drs. 
Barry and Cluver at the South African Museum is greatly appre­
ciated. This work was supported by grants from the Faculty of 
Graduate Studies and Research, McGill University, the Merrill 
Trust, and the National Research Council of Canada. 

ADDENDUM 
Since completing this paper, my attention has been drawn to 

a paper by Schaumberg (1976 ) describing two newly discovered 
specimens from the Upper Permian of Germany. They are very 
poorly preserved, but clearly show the presence of ribs similar 
to those of DaedalosauTUl. Schaumberg suggests similarities with 
WeigeltisauTUl, originally described by Weigelt as Palaeochamaeleo. 
None of these specimens are sufficiently well-preserved or thor­
oughly enough described for detailed comparison, but all may 
have been closely related. 

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