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Ann Pal$ontol (2001) 87, 1,61-73
0 2001 Editions sclentlfiques et midlcales Elsevler SAS Tous drolts r6servCs
Draconyx loureiroi, a new camptosauridae
(Dinosauria, Omithopoda) from the Late Jurassic
of LourinhQ, Portugal
Octho MATEUSb* ‘*, Miguel Telles ANTUNESa9 b9 ’
a Academia das Cu?nclas de Llsboa
b Centro de Estudos Geoldglcos, Faculdade de C&was e Tecnologra da Umversrdade Nova
de Llsboa, Qumta da Terre, 2825-114 Capanca, Portugal
’ GEAL - Museu da Lounnh& 2530 Laurmhi?, Portugal
(Received 6 October 2000, accepted after revlslon 20 December 2000)
Abstract - A new omlthopod dinosaur IS described here under the name of Draconyx
lourewol n gen , n sp on teeth, caudal vertebrae, forelimb, hmdhmb, and foot material that
were found m assoclatlon m the Late Jurassic-Tlth?man of LourmhB, Portugal Draconyx IS a
Camptosaumlae related to Camptosaurus 0 2001 Editions sclentlfiques et m&kales Elsevler
SAS
Dinosaurs / Ornithopoda / Draconyx / Portugal /Jurassic
R&m6 - Draconyx lourewoz, un nouveau camptosaundae (Dinosauria, Omithopoda)
du Jurassique suphieur de hwinh5, Portugal. Un nouveau dmosaure omithopode est ~1
dkcnt sous le nom de Draconyx lounwol n gen , n sp Des dents, des vert&bres caudales, une
partle des membres antkneur et posttneur, et le mat&e1 d’un pled ont t?tk trouvks assocks
dans le Jurasslque termmal-Tlthomen de ~ourmhl, Portugal Draconyx est un Campto-
saundae apparent6 .?I Camptosaums 0 2001 Editions sclentlfiques et mkdlcales Elsevler SAS
Dinosaures / Omitbopoda / Druconyx / Portugal / Jurassique
* Correspondence and reprmts
omateus@Qdmosaurs corn
mta@fct unl pt
-61-
0 MATEUS, M T ANTUNES
INTRODUCTION
Remnants of Late Jurassic ormthopods from Portugal have been described by
Sauvage [ 131, Thulborn [ 151, and Galton [3] The last one ascribed some hmb mate-
nal to the genus Camptosaurus
In 199 1, Carlos Anuncla@io (member of the Museum of Lourmhii) found some of
the foot and limb matenal of an omlthopod at Vale Frades, Lourmhii (figure 1)
Two omlthopod footprmts found at Port0 Escada m 1999 (unpublished data)
might be made by mdlvlduals of the species described here
SY STEMATICS
ORNITHISCHIA Seeley, 1888
ORNITHOPODA Marsh, 187 1
Iguanodontla Dollo, 1888
Ankylopollexla Sereno, 1986
Draconyx n gen
Etymology: Draco, latm word for dragon, and onyx, greek word for claw, m
recogmtlon of the claw matenal
Diagnosis: As for the species
Draconyx lowetrot n sp
Etymology: Zourezror, after JoPo de Lourelro (1717 - 1791), Portuguese Jesuit,
pioneer m Palaeontology m Portugal, also an excellent botanist, astronomer and med-
ical doctor, well-known for his “Flora Cochmchmensu” (he spent a large part of his
life m Southeast Asia)
Horizon: Late Jurassic - Tlthoman, “Bombarral” Unit according to Manuppella
(1986)
Locality: Vale Frades, LourmhB, western Portugal (figure I)
Material: Remains from one mdlvldual from Vale Frades (holotype) two maxll-
lary teeth, three mid-antenor caudal centra, one chevron, distal eplphysls of nght
humerus, one manual phalanx, three manual ungual phalanges, distal eplphysls of
nght femur, eplphyses of tibia and fibula, astragalus, calcaneum, three tarsals (II-V),
-62-
DRACONYX LOUREIROI, A NEW CAMPTOSAURIDAE
n Cenozoic
0 Tthoncan 1
Late
m Klmtnendglan Jurassic
n Lias ijF
0
!i
4
2
Figure 1 Slmphfied geologlcal map based on Manuppella [6] The followmg sites are shown from north
to south Palmogo (theropod nest and embryos [7, S]), Vale Frades (holotype of Draconyx gen et sp
n , this study), Peralta (holotype of Lmrmhanosaurus [9], Port0 das Barcas (holotype of Brachm-
saurus arala~~s, [5]), Port0 Dmheuo (holotype of Dmhetrosaurus [l]). and Valmltao (Ceruro-
saurus sp , [lo])
Figure 1. Carte geologlque slmphtiee d’apres Manuppella [6] Les sites smvants sont figuris du nord au
sud Palmogo (md avec des embryons de theropode [7, S]), Vale Frades (locahte type de Draconyx,
cette etude), Peralta (locahti type de Lourmhunosaurus [9], Port0 das Barcas (locahti type de Bmchzo-
saurus afalarenszs [5]), Port0 Dmhelro (locahte type de Dmhewosaurus [I]), et Valmitso (Ceruto-
saurus sp [lo])
four metatarsals (I-IV) and pedal phalanges Under the collection number ML 357 at
Museum of LourmM, Portugal figure 2)
Other referred matenal Femur (ML434, figure 7) from Prala do Canqal
Diagnosis:
CamptosauIldae with the followmg features
Max&try teeth with strong pnmary ridge with five tertiary ridges at the meslal
side The enamel on the labial surface of the tooth crown 1s lower distally than
mesially
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4 0 MATEUS, M T ANTUNES
Figure 2 Holotype of Draconyx Zourerrr~z gen et sp nov Ail these bones are. fkom a smgle mdwldual
(ML357)
Figure 2 Holotype de Druconyx Zourezmr gen et sp nov Tow les ossements concemes appartlennent &
un seul mdwldu (ML357)
Manual ungual phalanges higher than broad Tibia1 condyle of femur without lat-
eral projection into the flexor groove Flbular condyle of femur shortly proJected,
being shorter anteroposterlorly than wide medlolaterally Tibia1 condyle of femur
slightly proJected posteromedlally and concave on its medlal side Short cnemlal
crest on tibia Astragalus fused to calcaneum Distal tarsal II present Lateral distal
tarsal overlaps the medial distal tarsal Metatarsal V absent
DESCRIPTION
Teeth
Two max&try teeth were found The teeth are both higher than wide and rela-
tively compressed medlolaterally
Both teeth bear at their labial face one strong pnmary ridge,, five and two tertiary
ndges are present on their meslal and distal side respectively
The tooth enamel 1s present m labial and lingual crown, but m this last 1s
resticted to the distal half
Table I. Teeth Measurements
Tableau I. Mesures des dents
He&t Meslodistal length Lablolmgual width
164 95 59
11 2 85 88
All measurements are m mdhmetres
Toutes les mesures sont donnees en mdhmetres
- 64 -
DRACOMXLOUREIROI,A NEWCAMFTOSAURIDAE
figure 3 Draconyx Zouwzmr gen et sp nov Caudal vertebra m lateral and postenor views Scale bar 10 cm
figure 3 Draconyx lourerm gen et sp nov Vertkbre caudale, vue latemle et posteneure Echelle 10 cm
The root 1s sub-cylmdncal or slightly elhpsold
Small denticles can be seen on the margins of the crown
Axial skeleton (figure 3)
The three caudal vertebrae present have been part of the neural arch and centrum,
but the spines (neural and transverse processes) were not found
The three vertebrae are amphlplathyan or slightly oplsthocoehc In antenor view,
the centrum 1s rounded and m postenor view it 1s trapezoid-shaped
The neural channel 1s narrow and sub-round It 1s narrower than pedlcels.
In these vertebrae the dlapophyses are placed at the neural channel level and
apparently would be projected upwardly
The centrum 1s not pleurocoelous The facets for chevrons are very developed,
mainly on the ventro-antenor side
One single incomplete chevron 1s present It 1s not very expanded distally and 1s
as wide proximally as distally, with a small constnctlon m the middle
Table IL Caudal centra Measurements
Tableau II. Centres caudaux Mesures
Length He&t Width
52 58 60
47 52 50
46 46 46
All measurements are m mdhmetres
Toutes les mesures sont donnees en mdhmetres
-65-
6 0 MATEUS. M T ANTUNES
dc5
I
ul 1 2 cm I
Rgure 4 Druconyx louretroz gen et sp nov Carpal and metacarpal bones dc4- distal carpal 4, dc5- distal
carpal 5, mc- metacarpal, ul- ulnare
Flgure 4 Draconyx lourermr gen et sp nov Carplen and mkwxrp~en dc4- carplen distal 4 , dc5- carplen
distal 5 , mc- metacarplen , ul- ulnawe
Figure 5 Druconyx lourermr gen et sp nov Manual ungual phalanges Scale bar 1 cm
Figure 5 Draconyx lourezmz gen et sp nov Phalanges ungukales de la mam Echelle 1 cm
Forelimb (figures 4, 5)
A very badly preserved distal end of the humerus 1s present with two well defined
condyles
The manus 1s moderately preserved Three unksed carpal bones (distal carpal IV,
ulnare, and distal carpal V) and a proximal end of a metacarpal (W) were found m
anatomical connection One phalanx from the digit II or III and three ungual
phalanges were found without anatomical connection The three ungual phalanges
are higher than broad
Hindlimb (figures 6, 7, 6)
Only the holotype’s distal half of nght femur is preserved The distal half of the
femur 1s robust Its dlaphysls 1s slightly compressed anteroposterlorly
-66-
DRACONKX LOUREIROI, A NEW CAMPTOSAURIDAE 7
Figure 6 Draconyx louremt gen et sp nov Femur of holotype ML357
Figure 6 Draconyx Zoureml gen et sp nov F6mur de I’holotype ML357
An isolated left femur (ML434) with complete dlaphysls and part of the epl-
physes, was collected at Praia do Camqal, Just 1 km north of Druconyx holotype’s
site The lesser trochanter 1s well separated from the greater trochanter The femur 1s
very bowed postenorly The fourth trochanter 1s postenorly projected and mclmed
laterally Its edge is concave m posteromedlal view Adjacent and anterior to the
fourth trochanter there 1s a caudolhofemorahs muscle attachment
In the holotype (ML357) the two femoral condyles are well separated with well
developed antenor and postenor mtercondylar grooves The tibia1 condyle 1s slightly
concave m medial vrew and has no lateral proJection mto the flexor groove
fibular condyle is shortly projected postenorly, 1 e , shorter anteropostenorly
wide medlolaterally The fibular condyle 1s as wide anteriorly as postenorly
-67-
The
than
8 0 M~ATEUS, M T ANTLJNES
Figure 7 Femur of Draconyx Iourerm gen et sp nov ML434 Scale bar 10 cm
Figure 7 F&mm du Draconyx lourerm gen et sp nov ML434 kchelle 10 cm
The nght tibia is almost completely preserved (about 20 % of the mlddle of the
dlaphysls 1s lackmg) The proximal eplphysls 1s very strong 143 mm anteroposte-
norly and 111 mm medlolaterally The fibular condyle of the tibia 1s narrow and
lateroposterlorly proJected, it forms a notch with the posterror condyle The cnemlal
crest 1s very wide and short
In proximal view the eplphysls 1s flat and forms a very pronounced, convex half-
circle on its medial side The fibula was preserved entirely m cormectlon with the
lateral side of the fibular condyle
The dlaphyseal section 1s pointed oval shaped (water-drop shape) Distally the
tibia 1s fused to the astragalus and calcaneum and anatomically attached to the fibula,
tarsals and metatarsals It 1s much expanded lateromedlally and compressed antero-
postenorly
The fibula 1s very slender m comparison with the tibia It is anteroposterlorly
expanded and convex laterally On its medlal view, It bears the msertlon to the fibular
condyle of the tibia Distally the fibula 1s attached to the tlbla and the calcaneum
covers it dorsally
-68-
DRACONYX LOUREIROI, A NEW CAMPTOSAURIDAE
c 10cm t
Figure 8 Draconyx louwrmr gen et sp nov Tibia m proximal view (ML357)
figure 8 Draconyx loummz gen et sp nov Tlbla, we proxlmale (ML357)
Foot (figure 9)
The astragalus and calcaneum are almost entirely covered by the distal part of the
tibia and fibula The calcaneum is concave laterally
Three distal tarsals are present The lateral distal tarsal (IV) is 65 mm wide and
18 mm long It contacts dorsally with the calcaneum and astragalus and laterally with
the fibula It ventrally overlaps the medial distal tarsal (III) and the metatarsal IV
The preservation of tarsal II 1s very rare m dinosaurs In this case it 1s vestlglal
with ca 24 mm wide medlolaterally and 8 mm thick anteropostenorly It IS placed
between the tarsal III and the astragalus
Four metatarsals are present (I to IV) The (inner) metatarsal I is vestigial The
metatarsal I 1s not completely preserved but its lmpresslon on the metatarsal II 1s
perfectly preserved It 1s slightly bowed, fitting entirely mto the medial side of the
metatarsal II m the diagonal posltlon D&ally it 1s curved upwards
The metatarsal II has the largest proximal end of these metatarsals Its proximal
surface 1s longer anteropostenorly than wide medlolaterally
The metatarsal III 1s the longer one and with metatarsals II, IV, and distal tarsal
III The metatarsal IV has uniquely the both ends preserved
The metatarsal V 1s not preserved and there are no Impressions or anatomical
place for it We may therefore conclude that the metatarsal V was already absent m
Draconyx (derived character)
Draconyx holotype has three articulated phalanges of the pedal digit II Two other
phalanges (one ungual) are present The digit III Just has one articulated phalanx pre-
served Two ungual phalanges found might correspond to dlglt II and IV
-69-
mt3
I
4
0
I
mt3
.mtl
I
i 1
I ); f
1 Ocm 5
I I
Flpm 9 R&t foot ofLhconyx hre~~~ gen et sp nov (ML357) as astragalus, cal calcaueum, fi fibula, mtl-4 metatarsus I to IV, respectively, ta2-4 tarsals II, III
and IV respectwely, tl tlbla
Figure 9 Pled drod de Druconyx louremr gen et sp nov (ML357) as astragalus , cal calcaneum , ii p&on6 , m tl-4 metatarses I, II, III et IV, respectlvement ,
ta 2-4 tarslens II, III and IV, tl tlbla
DRACO~YX L~UREIROI, A NEW CAMPTOSAURIDAE 11
Table III. Metatarsals Measurements
Tableau III. Metatarslens Mesures
Length
Proximal width (anteropostenor/medlolateral)
I II III IV
53 152 175 7
10/6* 89/26* 7/35* ?I67
Distal width (anteropostenor/medlolateral) 16*/7 52145 57164 43143
All measurements are m mllhmetres * Estimated
Toutes les mesures sont don&es en mdhmetres * Estlmee
COMPARISONS AND DISCUSSION
Draconyx lourezroz IS clearly an Iguanodontla (sensu Carla & Salgado, 1996
= Tenontosaurus + Emguanodontla) because it shows the followmg characters
- Enamel present on the medial side of maxillary teeth [2]
- Leaf-shaped dentlcles [ 141
- Enamel restncted to the distal half of the crown on the medial side of the maxll-
lary teeth [ 141
- Presence of antenor mtercondylar groove on femur [2, 141
- Femur with fibular condyle medially placed
It 1s an Emguanodontra (Gaspannuaura + Dryomorpha) due to
Presence of lateral primary ndge m maxillary teeth [2]
It can be included among the Dryomorpha (Dryosaurus + Ankylopollexla, sensu
Sereno, 1986) due to
- Diamond-shaped maxillary tooth crown with rounded anterior and postenor
comers [ 141
- Enamel absent from the medial side of the maxillary teeth [ 141
- Channel like anterior mtercondylar groove on femur [ 1 l]
And among the Ankylopollexla Sereno 1986 (Camptosaundae + Styracostema)
(=Camptosauna [ 111) due to
- The prominent primary ndge on the labial surface of the maxillary teeth crowns
Vl, 141
- Partial fusion of ulnare and distal carpals 4 and 5 [ 141
- Pes digit 1 relatively shorter and less robust [ 141
- Metatarsal 1 markedly less robust relative to the other metatarsals [ 141
-71-
12 0 MATEUS, M T ANTUNES
It can be ascribed to Camptosaundae because
- The maxillary teeth have a strong vertical pnmary ridge on the distal side of the
labial crown
- The femur 1s curved and has a prominent lesser trochanter
Followmg Norman [l l] Camptosaundae IS composed by the genera Cumpto-
saunas, Cumnorza, Callovosaurus and an “unnamed species from Portugal (Galton,
1980)” Cumnona prestwzchzz was recogmsed mto the genus Camptosaurus (as
C prestwzchzz) by [3,4, 121 and commonly accepted Callovosaurus leedsz IS seen as
nomen dubzum because It was based m two mcomplete femoral eplphyses without
any strong dlagnostlc feature
Therefore, we restnct the family Camptosaurldae to Cumptosaurus (w&h three
species C dzspal; C amplus and C prestwzchzz) and Draconyx lourezroz that
includes the femur from Portugal described as Camptosaurus sp by Galton [3] and
cited by Norman [ 1 l]
Draconyx IS dlstmgmshable from Camptosaurus by (1) tibia1 condyle of femur
wlthout lateral proJection mto the flexor groove, (2) fibular condyle of femur wider
medlolaterally than long anteropostenorly, (3) vestigial first pedal digit, and (4)
absent pedal digit V
CHRONOLOGICAL AND GEOGRAPHICAL REMARKS
The descrlptlon of Druconyx as Camptosamdae 1s coherent with the chronology
of this family, placed m Knnmendglan and Tlthoman of North Amenca and England
(contemporaneous to Druconyx’s Vale Frades site) Galton [3] already cited the
family to Portugal and discussed its paleogeographlcal lmphcatlons The dinosaur
fauna1 list from the Late Jurassic of Portugal 1s related to the Momson formation
(with the same families presented m both areas) but often different at the genus level
showing some taxonomlcal divergence due to its paleogeographlcal separation
Acknowledgements - We are mdebted to Carlos Anuncla@o who found the specimen, to
GEAL - Museu da Lourmha team (Horilao Mateus and Vasco Rlbelro) and Mus&nn National
d’Hlstolre Naturelle team (Phlhppe Taquet) for the dlscusslon and preparation support We are
also grateful to LeaderOeste and Programa Cl&la Viva that financed part of the laboratonal
matenal This study was supported by PhD scholarship grant 21616/99 (to 0 M ) of PRAXIS
XXI of the Mmlstkno da C&ncla e Tecnologla (Centro de Estudos Geol6glcos) All draws by
Octivlo Mateus
VI
VI
REFERENCES
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DRACONYX LOCIREIROI, A NEW CAMPTOSAUR~DAE 13
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