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ABSTRACT
A new basal non-pterodactyloid pterosaur, Raeticodactylus filisurensis gen.
et sp. nov., is reported. It has been discovered in shallow marine sediments
from the Upper Triassic of the lowest Kössen beds (late Norian/early Rhae-
tian boundary) in the central Austroalpine of Canton Grisons (Switzerland).
The disarticulated specimen is comprised of an almost complete skull and a
partial postcranial skeleton. A high and thin bony, sagittal cranial crest char-
acterizes the anterodorsal region of the skull. The large mandible, with an ad-
ditional keel-like expansion at the front, partly matches the enlarged sagittal
cranial crest. A direct and close relationship to Austriadactylus cristatus, the
only known Triassic pterosaur with a bony cranial crest so far, cannot be estab-
lished. The teeth of the premaxilla are monocuspid and exhibit very strongly
bowed enamel wrinkles on the lingual side whereas the enamel is smooth on
the labial side. These monocuspid teeth are large and fang-like. The numer-
ous smaller teeth of the maxilla show three, four and five cusps. These are
very similar to the teeth of the Triassic pterosaur Eudimorphodon ranzii. The
humerus shows a thinner construction than that seen in other Triassic ptero-
saurs. The femur is quite unusual with a caput femoris perpendicular to the
shaft. The bones of the extremities are almost twice as long as the ones from
the largest Triassic specimen E. ranzii (MCSNB 2888). The newly described
pterosaur is an adult, with a wingspan of approximately 135 cm. A morpho-
functional analysis suggests that R. filisurensis was a highly specialized pisci-
vore and possibly a skim-feeder.
ZUSAMMENFASSUNG
Beschrieben wird ein früher langschwänziger Pterosaurier Raeticodactylus
filisurensis gen. et sp. nov. Entdeckt wurde dieser in den Flachwasserkarbo-
natablagerungen aus der oberen Trias aus den untersten Kössener Schich-
ten (Grenzbereich Norian/Rhaetian) des Zentralostalpins von Graubünden
(Schweiz). Der disartikulierte Fund enthält den beinahe kompletten Schädel
und Teile des postcranialen Skelettes. Der Schädel trägt auf der Schnauzen-
partie einen hohen und dünnen Knochenkamm. Im Zusammenhang mit dem
sagittalen Schädelkamm steht der hohe Unterkiefer mit einer im vorderen Un-
terkieferbereich zusätzlich auftretenden kielartigen Erhöhung. Eine direkte
und enge verwandtschaftliche Beziehung zu Austriadactylus cristatus, welcher
bisher der einzige bekannte Flugsaurier mit knöchernem Schädelkamm aus
der Trias war, konnte nicht festgestellt werden. Die Zähne der Prämaxilla sind
einspitzig und zeigen auf der lingualen Seite eine starke komplizierte, bogen-
artige Riefung, während sie lateral einen glatten Zahnschmelz aufweisen. Die
einspitzigen Zähne haben eine fangähnliche Funktion. Die vielen, kleinen
Maxillarzähne zeigen drei, vier und fünf Spitzen. Diese sind den Zähnen des
triassischen Flugsauriers Eudimorphodon ranzii sehr ähnlich. Der Humerus
zeigt im Vergleich zu andere triassischen Pterosaurier eine deutlich schlan-
kere Bauweise. Das Femur ist ungewöhnlich, weil der Gelenkkopf rechtwink-
lig zum Knochenschaft steht. Die Extremitätenknochen sind fast doppelt so
lang wie diejenigen des grössten Exemplares E. ranzii (MCSNB 2888). Der
neubeschriebene Pterosaurier stammt von einem adulten Exemplar, welches
eine Flügelspannweite von ungefähr 135 cm hatte. Die funktionsmorpholo-
gische Analyse zeigt, dass R. filisurensis gen. et sp. nov. ein hochspezialisierter
Fischfresser und möglicherweise ein Skimmer war.
Institutional abbreviations
BNM, Bündner Naturmuseum, Chur (Switzerland); BSP,
Bayerische Staatssammlung für Paläontologie und historische
Geologie, Munich (Germany); MCSNB, Museo Civico di Sci-
enze Naturali, Bergamo (Italy); MFSN, Museo Friulano di Sto-
ria Naturale, Udine (Italy); MGUH, Geological Museum, Uni-
versity of Copenhagen, (Denmark); MNHNL, National Mu-
seum of Natural History, Luxemburg (Luxemburg); MPUM,
Dipartimento de Scienze della Terra, Università di Milano
(Italy); PIMUZ, Paläontologisches Institut und Museum der
Universität Zürich (Switzerland); SMNS, Staatliches Museum
für Naturkunde, Stuttgart (Germany).
Introduction
Zambelli described the first known Triassic pterosaur, Eudimor-
phodon ranzii, in 1973. In the following 30 years new remains
of Triassic pterosaurs were discovered, especially in northern
Italy (Wild 1978, 1984, Dalla Vecchia 1995). Other specimens
A new Triassic pterosaur from Switzerland (Central Austroalpine,
Grisons), Raeticodactylus filisurensis gen. et sp. nov.
RICO STECHER
Key words: pterosaur, non-pterodactyloid, Upper Triassic, Ela nappe, Kössen Formation, Switzerland
DOI 10.1007/s00015-008-1252-6
Birkhäuser Verlag, Basel, 2008
Swiss J. Geosci.
Schellenbergstrasse 13, 7000 Chur, Switzerland. E-mail: ricostecher@yahoo.com
A new Triassic pterosaur from Switzerland 1
2 R. Stecher
from the Triassic were reported from Austria (Dalla Vecchia
et al. 2002, Wellnhofer 2003), France (Godefroit & Cuny 1997),
Britain (Fraser & Unwin 1990), Luxembourg (Cuny et al. 1995),
Greenland (Jenkins et al. 2001) and the USA (Murry 1986).
Remains of Triassic pterosaurs are also known from Switzer-
land. Clemens (1980) noted that some mammal-like teeth from
the Rhaetian of Hallau described by Peyer (1956) are poten-
tially teeth of Eudimorphodon. More recently, Fröbisch &
Fröbisch (2006) described an incomplete lower jaw from the
Schesaplana (Northern Calcareous Alps) in Switzerland and
established a new genus and species, Caviramus schesaplanen-
sis. Other Swiss pterosaur finds are from the Upper Jurassic: a
second wing phalanx of a pterodactyloid from the Solothurn
Turtle Limestone (Canton Solothurn, Meyer & Hunt 1999), a
wing phalanx fragment of a non-pterodactyloid from Porrent-
ruy (Canton Jura, Billon-Bruyat 2005) and unpublished bones
from the areas of Biel (Canton Bern) and Olten (Canton So-
lothurn) (Billon-Bruyat 2005). Swiss pterosaur finds can there-
fore be considered rare and generally poorly preserved. This
makes the pterosaur described in this paper the more complete
skeleton from Switzerland known so far.
The aim of this study is to describe this new pterosaur and
compare it to other Triassic pterosaurs. Currently, five genera
and seven species of Triassic pterosaurs are known: Austriadac-
tylus cristatus DALLA VECCHIA, WILD, HOPF & REITNER 2002
(Holotype: SMNS 56342, a poorly preserved specimen with a
bony crest on the skull; Seefeld Schichten, Austria); Caviramus
schesaplanensis F RÖBISCH & FRÖBISCH 2006 (Holotype: PIMUZ
A/III 1225, a right mandible that is overall poorly preserved;
Kössen Formation, Alplihorn Member, Schesaplana, Northern
Calcareous Alps, Switzerland); Eudimorphodon cromptonellus
Fig. 1. Map of Switzerland and the Canton Grisons indicating the locality of
Tinzenhorn, where Raeticodactylus filisurensis gen. et sp. nov. (BNM 14524)
was found.
Fig. 2. Photograph of Raeticodactylus filisuren-
sis gen. et sp. nov. (BNM 14524, Upper Triassic,
Tinzenhorn, Filisur, Switzerland), after prepara-
tion. The scale bar is in centimetres.
A new Triassic pterosaur from Switzerland 3
JENKINS, SHUBIN, GATE S Y & PADIAN 2001 (Holotype: MGUH
VP 3393, a disarticulated skeleton; Fleming Fjord Formation,
Orsted Dal Member, Greenland); Eudimorphodon ranzii
ZAMBELLI 1973 (Holotype: MCSNB 2888, a nearly complete
skeleton; Calcare di Zorzino, Cene, Bergamo, Italy); Eudimor-
phodon rosenfeldi D
ALLA VECCHIA 1995 (Holotype: MFSN
1797, a nearly complete skeleton with partially preserved skull;
Dolomia di Forno Formation, Enemonzo, Udine, Italy); Petei-
nosaurus zambellii W ILD 1978 (Holotype: MCSNB 2886, a com-
plete skeleton; Calcare di Zorzino, Cene, Bergamo, Italy); and
Preondactylus buffarinii WILD 1984 (Holotype: MFSN 1770, a
nearly complete postcranial skeleton without skull; Dolomia di
Forno Formation, Friuli, Udine, Italy).
Geology and depositional environment
The new pterosaur was found at the Tinzenhorn (Corn da Ti-
nizong, Canton Grisons), located in the Bergüner Stöcken (Piz
Ela, Tinzenhorn, Piz Michel), Mittelbünden, eastern Switzer-
land (Fig. 1). Tectonically, the Tinzenhorn belongs to the Ela
Nappe (Furrer 1993), and consists of Triassic and Jurassic ma-
rine sediments. The top of the Tinzenhorn is a lying anticline
and at its northern part the layers are vertical and show impres-
sively the transition from the Norian, Rhaetian to the Jurassic
sediments. The first detailed stratigraphical and tectonical de-
scriptions of this region were from Frei (1925) and Frei & Ott
(1926). Frei (1925) mentioned that the best locality for fossils
is the small valley next to the Fil da Stidier. It is noteworthy
that he only mentioned fossils of invertebrates, although those
of vertebrates are mostly black in colour and then very notice-
able. Almost 50 years later, Furrer (1974) described vertebrates
from the Kössen Formation found at the Piz Mitgel. Rohrbach
(1977) also found vertebrate remains at the Tinzenhorn includ-
ing teeth of hybodont sharks, teeth and scales of actinopter-
ygians. Studies by Duffin & Furrer (1981), Bürgin & Furrer
(1992, 1993) and Furrer (1993) uncovered yet more vertebrates
(mainly tooth plates and fin spines of myriacanthoid holo-
cephalans). They also mentioned remains of hybodont sharks,
actinopterygians, placodonts, ichthyosaurs and phytosaurs. The
vertebrates were usually found in bonebeds, on layer surfaces
and on hardgrounds.
In August 2005, the author discovered some long, thin,
blackish, fossil bones on a massive (12 cm thick) limestone slab
(40 cm by 40 cm), in the basal part of the Kössen Formation
(Fig. 2). Only the posterior part of the skull was visible. Some
bones were corroded by the weather and others had been de-
stroyed. The limestone slab was lying directly under vertical
beds from the Kössen Formation. It can be confidently assigned
to the Alplihorn Member of the Kössen Formation (Central
Austroalpine) (Figs. 3a, b). The Alplihorn Member, the most
basal Member of the Kössen Formation, is very rich in inverte-
brate and vertebrate fossils, however the vertebrates are mostly
highly disarticulated and fragmented. At the Fil da Stidier, the
Alplihorn Member can reach about 58 m in thickness. It origi-
nates from a short transition zone from the basal Uglix-Plat-
Fig. 3. a) Stratigraphy of the Alplihorn Member (Kössen Formation, Upper
Triassic) at the Fil da Stidier (Canton Grisons). b) View of Fil da Stidier as seen
from the Val Gravaratschas. The arrow shows the location where BNM 14524
was found. Abbreviations: AM, Alplihorn Member; KF, Kössen Formation;
SM, Schesaplana Member of KF; UP, Uglix-Plattenkalk.
4 R. Stecher
tenkalk (Hauptdolomit-Gruppe). It consists of interbedded
strata of black, brown, yellow weathered clayey shales, dark-
grey micritic limestones and olive calcareous dolomite, often
laminated micrites and marly intercalations. This sequence was
interpreted as a wide, flat lagoon or shallow basin on the ex-
tensive carbonate platform of the western margin of the Tethys
Ocean in the Late Triassic (Furrer 1993).
Under the pterosaur-bearing layer, a blue-grey weathered,
massive limestone occurs. Immediately below the specimen, the
sediment shows conspicuous bioturbation. The sediment encas-
ing the pterosaur is a dark grey limestone. The sedimentary unit
(12 cm thick) including the fossil can be divided into three main
parts (Fig. 4): 1) Basal most part (3 cm thick): a micritic lime-
stone with less fossil shells than seen elsewhere; the base is ero-
sional with scour-and-fill-structures; 2) Middle part (5 cm): the
matrix is micritic and contains many ooids, and shells (mostly
convex side up) and shell fragments; lamellibranchs with both
shells preserved are rare; the ooids often have fragments of
bivalves and gastropods as nuclei; towards the upper part the
ooids show a decrease in grain size, indicating a small scale fin-
ing upward sequence; 3) Upper part (4 cm): this micritic part is
the matrix of the pterosaur, it shows marly laminated interca-
lations; the ooid content decreases gradually, they are missing
in the uppermost part; the last part of the slab is built up by
laminated, yellow, calcareous marl; the latter yields thin shell
fragments and rare ganoid scales.
The layer above the specimen is of a 1 cm thick grey clay
followed by a 28 cm thick finely laminated grey limestone. The
presence of ooids indicates shallow turbulent water in a sub-
tidal environment (Furrer 1993). The ooids from the middle
part come from such an environment but were transported and
redeposited by a storm. The normal graded bedding, the con-
vex up/down position of the shells and the frequent telescoping
position support this. The layer including the pterosaur is inter-
preted as a storm deposit, a tempestite that was subsequently
bioturbated (presence of Thalassinoides). The increased marl
content, the complete absence of ooids in the uppermost part
and the clay layer indicate laminar, non-turbulent lower energy
conditions. The thin layer of grey clay can be interpreted as a
fine deposition during a quiet sedimentation period. The finely
laminated, yellow, limestone is interpreted as cyanobacterial
mats developed in the intertidal zone.
It is suggested that the pterosaur body sank shortly after the
storm to the bottom of the shallow marine environment and
was rapidly buried (excluding a long post-mortem transport),
leading to the excellent preservation of the specimen.
Material and methods
The reported specimen (BNM 14524) consists of a disarticu-
lated skeleton, two isolated teeth, an isolated rib fragment
and an isolated hyoid fragment. One of the isolated teeth lay
6 cm from the skull, completely embedded in the limestone; it
was fully separated from the slab for further examination. The
second isolated tooth was still in the mouth but not attached to
the jaws, it was also removed. The rib fragment was a rib stem
lying on the lower jaw, covering several teeth. To allow further
examination of these teeth, it was necessary to separate a part
of this bone from the main slab.
The preparation was challenging because of the very frag-
ile skull. The bones were embedded in a very hard and massive
limestone. The preparation was first carried out using a weak
formic acid; this allowed the extraction of the black bones from
the light-grey weathered limestone. Bones were protected
from acid by using a high quality acrylic resin (“OSTEO-
FIX”). The upper part of the limestone slab is a finely lami-
nated calcareous marl, making impossible a preparation with
acid alone; an abrasive fibreglass eraser (“ECOBRA”) was
used. The final conservation of the fossil was completed with
“OSTEO-FIX”.
Systematic palaeontology
Class Reptilia LAURENTI 1768
Subclass Archosauria COPE 1869
Order Pterosauria KAUP 1834
Although the tail is not preserved, BNM 14524 can be assigned
with confidence to a non-pterodactyloid (“short-tailed”) ptero-
saur according to the following features (Wellnhofer 1993, Un-
Fig. 4. Sedimentological section through the layer where the reported ptero-
saur was found. 1) Micrit limestone with only few shells. 2) Micrit, limy ma-
trix with many ooids (the largest are 0.8 mm in diameter) and shells (mostly
convexly adjusted). 3) Micrit, limy matrix with only few ooids. Bioturbation.
4) Micrit, limy matrix with little ooids (the largest are 0.2 mm in diameter)
and broken shells which are convexly adjusted. 5) Micrit, limy matrix with
rare ooids but many shells. 6) Laminated calcareous marl with few shells but
without any ooids. The scale bar is in centimetres.
A new Triassic pterosaur from Switzerland 5
win 2003a): The narial and antorbital fenestrae are separated
by a bony strip; the occipital condyle is arranged backwards;
the wing phalanges have a furrow on the posterior side; a long
5th toe is present.
Genus Raeticodactylus gen. nov.
Etymology. – Raetia (Latin): old name of the Swiss Canton Gri-
sons, where the pterosaur was found; dactylus (Greek): for the
long fourth wing finger.
Diagnosis. – As for the type species.
Species Raeticodactylus filisurensis sp. nov.
(Figs. 2, 5–11)
Etymology. – Filisur (Romansch): name of the village where
the pterosaur was found; ensis (Latin): for “from”.
Holotype. – The holotype is housed in the Bündner Naturmu-
seum (BNM) of Chur (Grisons, Switzerland), with the collec-
tion number BNM 14524 (Fig. 2).
Material. – A disarticulated skeleton including a nearly com-
plete skull, a mandible, two isolated teeth and a postcranial
skeleton (right humerus, right ulna, right metacarpals I–III,
right and left wing phalanges 1–4, right femur, right tibia, part
of the right fibula, part of the right pes, several cervical, dorsal
and several ribs).
Locus typicus. – 1.25 km northeastern of the Tinzenhorn at the
Fil da Stidier. The Tinzenhorn (Corn da Tinizong) is a moun-
tain of the Bergüner Stöcke in Mittelbünden, commune Filisur,
Canton Grisons, Switzerland (Fig. 1). The exact geographical
coordinates are deposited in the BNM.
Stratum typicum. – The layer in which BNM 14524 was found is
located in the lower part of the Alplihorn Member of the Kös-
sen Formation. Characteristics for the Alplihorn Member are
the interbedded strata of black, brown, yellow weathered shale
Fig. 5. Drawing of Raeticodactylus filisurensis gen. et sp. nov. (BNM 14524). Abbreviations: cv, cervical vertebra(e); dr, dorsal rib(s); dv, dorsal vertebra(e); fe,
femur; fi, fibula; gsc, ganoid fish scale; h, humerus; it, isolated tooth; l, left; m, mandible; mc I–III, metacarpals I–III; mt I–V, metatarsals I–V; rad, radius; r, right;
s, skull; ti, tibia; tv, thoracic vertebra(e); u, ulna; wph 1–4, wing phalanges 1–4. The scale bar is in centimetres.
6 R. Stecher
clay, dark-grey micritic limestones and the olive calcareous-do-
lomite, often laminated micrites and marl layers (Figs. 3a, b).
Age. – Late Triassic: late Norian (Sevatian) or Rhaetian. Furrer
(1993) states that the age from the Alplihorn Member is not ex-
actly known, most probably it is Sevatian (late Norian) in age.
Diagnosis. – The holotype specimen of the monospecific ge-
nus Raeticodactylus presents the following diagnostic features:
Bony, anterior cranial crest formed from the premaxilla; keel
like extension of the anterior part of the mandible; heterodont
dentition with large monocuspid teeth in the anterior part and
multicuspid (tricuspid, quadricuspid, quinticuspid) teeth in the
posterior part of the dentition; diastema in the upper jaw be-
tween the premaxilla and maxilla; lingual side of the anterior
teeth with heavy and conspicuous bowed enamel wrinkles;
multicuspid teeth of the upper jaw more bulbous than the teeth
of the lower jaw; multicuspid teeth of the upper jaw aligned in
one row and show distinct gaps between the teeth; multicuspid
teeth of the lower jaw packed close together causing the orien-
tation of the teeth to slope and laterally overlap; very long, slen-
der (length to diameter shaft ratio: 18.2) and straight humerus,
with a subrectangular deltopectoral crest.
Description. – The holotype (BNM 14524) is mainly a disarticu-
lated skeleton. Nevertheless, some parts are in natural associa-
tion, especially the skull and the mandible, the wing phalanges
and elements of the right pes (Fig. 5). The pectoral girdle, the
pelvis, the majority of the ribs and vertebrae, particularly all
caudal vertebrae and some bones of extremities are missing.
The ossification of the epiphysis, the complete and fine porous
surface of the bones (possibly) and the relatively small orbit
(see below the comparisons of the skull-orbit length index)
suggest that BNM 14524 was fully grown.
Skull and mandible (Figs. 6a–c). The skull is well preserved.
Only some teeth are broken or lost and the posterior part of
the skull was distorted by diagenesis. The skull lies on its right,
lateral side. The mandible is still articulated with the skull. The
skull has a low and long form; it measures 95 mm from the tip
of the snout to the occipital condyle. At its highest point (above
the orbit) the skull is 19 mm high. The largest opening of the
skull is the orbit with a diameter of 20 mm. The antorbital fe-
nestra is oval and longer than high (19 mm and 11 mm respec-
tively). The naris lies in a posterior position on the snout. It
has an unusual “tear-drop” shape with the rounded part located
rostrally. The naris is 14 mm long and 4.5 mm high.
The outstanding anatomical feature of the skull is the anterior,
large, sagittal bony crest, extending from the premaxilla and
lying over the naris. It ascends from the tip of the snout di-
rectly to a height of about 21 mm and descends with a slight
concave curve smoothly to end above the middle part of the
antorbital fenestra. The anterior part of the bony crest is about
1 mm thick; the posterior part is only a fraction of a millimetre.
The bony crest surface shows strong radial ridges, as does the
anterior part of the skull. It is supposed that the bony crest but
also the tip of the snout were covered by a keratinized sheath
(a rhamphothecae). A conspicuous suture like structure, visible
between the bony crest and the premaxilla, may have separated
the soft tissue part from the rest of the skull.
The very thin mandible is 84 mm long and 9.5 mm height. The
right and left lower jaws meet in an extremely thin, common
keel. Its anterior end descends to form a keel-like expansion
measuring 14 mm at its deepest point. The tip of the mandible
is toothless, pointed and directed dorsally. The surface of the
tip of the mandible reveals similar ridges than on the premax-
illa and on the bony crest; the anterior part of the mandible
was probably covered by a keratinous rhamphothecae. Numer-
Fig. 6. Skull of Raeticodactylus filisurensis gen. et sp. nov. (BNM 14524) in
left lateral view. Photograph (a), drawing (b) and reconstruction (c). Abbre-
viations: a, angular; ar, articular; co, coronoid; d, dentary; f, frontal; j, jugal; m,
maxilla; n, nasal; pf, postfrontal; pm, premaxilla; pmsc, premaxillar sagittal
crest; po, postorbital; prf, prefrontal; q, quadrate; sa, surangular; sp, splenial;
sq, squamosal. The scale bar is in centimetres.
A new Triassic pterosaur from Switzerland 7
ous foramina are found on both the maxilla and mandible. The
maxilla exhibits 13 irregularly spaced, backwards facing foram-
ina. Seven pocket-shaped foramina are present in the anterior
part of the mandible.
Dentition (Figs. 7–9). Some teeth were broken before the
death of the pterosaur, or were lost post mortem, but before
fossilisation. The teeth have a single root in both the upper and
lower jaws. Based on the teeth and alveoli, it is possible to state
the original number of teeth of the specimen: the left upper jaw
and mandible have 17 and 22 teeth respectively, that is a total
of 78 teeth. In the left upper jaw, the 11th–15th and 17th teeth
are broken. The 7th, 8th and 16th teeth show strong wear facets.
In the left mandible, the 5th and 6th teeth were lost before the
pterosaur was preserved. The 8th, 20th and 22th teeth are broken.
The 7th tooth shows also important wear facets, so much so that
is impossible to reconstruct its original form. In the premaxilla
are four monocuspid, fang-like teeth. Distally the 4th tooth has
a conspicuous bulge. The first three teeth are slightly recurved.
Between the teeth of the premaxilla and the maxilla a 5 mm
long diastema is apparent. The thirteen teeth of the maxilla are
tricuspid to quinticuspid. The first quinticuspid tooth in the up-
per jaw is the 9th.
In the anterior part of the mandible, three large monocuspid,
fang-like teeth are visible. Another monocuspid tooth and some
tri-, four-, and quinticuspid teeth succeed these three teeth. The
first quinticuspid tooth of the mandible is the 10th.
The monocuspid teeth in the anterior dentition are large and
have a crown up to 4 mm in length. On the labial side there
is – to a greater or lesser extent – smooth enamel with hardly
any enamel wrinkles. In contrast, on the lingual side strong and
conspicuous enamel wrinkles are apparent. The narrow stand-
ing enamel wrinkles are partially bifurcated and have a bowed
form along the crown (Fig. 8).
The multicuspid teeth in the posterior part have generally a
conical, flat form and show no or one to two small accessory
cusps mesially and/or distally to the main cusp. The cutting dis-
tal and mesial edges have a convex form. Some of the most
anterior multicuspid teeth exhibit thickened bulges along the
cutting edges. The multicuspid teeth of the maxilla are slightly
more bulbous, as the conical teeth of the mandible, and have
larger gaps between them. The multicuspid teeth of the man-
dible are very close together, in a sloping position and mutu-
ally overlap. The five supplementary teeth of the mandible are
within the same distance, therefore the upper/lower teeth ratio
is 0.78.
Some teeth of the upper jaw and mandible mainly show two
types of strong wear facets, at the top of the crown (horizon-
tal wear) and on the labial side of mandible teeth. This can be
traced back to contact of the teeth of the upper jaw. No wear
facets are present on the labial side of teeth of the upper jaw.
The horizontal heavy wear facet of the main cusp suggests a
durophagous diet.
Examining the mutual position of the skull to the mandible, this
latter seems slightly displaced forwards. Moving the mandible
backwards would enable its 3rd tooth to fit exactly in the dia-
stema of the upper jaw. However, this would result in a shorter
Fig. 7. Detailed drawing of the upper and lower jaws of Raeticodactylus filisu-
rensis gen. et sp. nov. (BNM 14524). The scale bar is in centimetres.
Fig. 8. Detailed drawings and photograph of
fang-like teeth of Raeticodactylus filisurensis gen.
et sp. nov. (BNM 14524). a) Labial view of the 2nd
tooth of the left upper jaw. b) Lingual view of the
2nd tooth of the right upper jaw. c) Photograph of
the 2nd tooth (~ lingual view) of the right upper
jaw. The scale bar is in millimetres.
mandible compared with the upper jaw. A keratinized beak,
at the tip of the mandible, could have completed this missing
length.
The two isolated teeth are very thin and show two small ac-
cessory cusps mesially and/or distally of the main cusp (Fig. 9).
The size and shape of the two teeth suggest that both are from
the mandible.
Vertebrae and ribs. Eight vertebrae are preserved, including
two cervicals and six dorsals. The cervical vertebrae look very
large and massive. They are up to 20 mm long and 14 mm wide.
They are very weak, pneumatized and show very thin bony
walls. The six dorsal vertebrae lie on their lateral faces, nearly
in articulation. These vertebrae are smaller (7 mm in length)
and more pneumatized. No caudal vertebrae are preserved.
Six ribs are preserved. The length of three dorsal ribs ranges
between 30 and 35 mm. The ribs are two headed and then taper
distally to a point. Some vertebrae cover two further ribs. The
6th rib is preserved in abutted position at the humerus; it has
only one head, is very thin with a strongly evolved arch in the
proximal part and is straight in the distal part.
Forelimb (Fig. 10). The right humerus lies parallel next to
the right ulna with adjacent articular ends. The humerus lies
on its lateral face. The shaft of the humerus is partly broken
and shows that it is hollow with a large pneumatic cavity. The
humerus has a length of 82 mm, the diameter of the shaft is
4.5 mm and at its widest it is 22 mm across. It has a straight and
slender form with a subrectangular, wide and well-developed
deltopectoral crest.
The right ulna is long, slender with well-developed articular
ends. It is 106 mm long and pneumatized. Only a few centime-
tres from the distal end of the humerus and of the proximal
end of the ulna, a fragment of another bone with parts of an
articular end is present; it was not possible to identify it.
The metacarpals I–III of either the right or the left side are pre-
served in association. These slender bones have approximately
similar lengths (metacarpal I: 40 mm; II: 41 mm; III: 42 mm).
The distal ends of the metacarpals are slightly wider than the
proximal ends. A well preserved manual claw lies close to these
metacarpals. The wing metacarpal (IV) is not preserved.
The wing phalanges of BNM 14524 are nearly complete from
both left and right wings and are mostly articulated. All wing
phalanges of the right wing are preserved, at least partially. Of
the 1st wing phalanx, the dorsal side is visible. It is 113 mm long
and the most robust of the wing phalanges. The 2nd wing phalanx
is 109 mm long. Only the proximal articular ends of the 3rd and
4th wing phalanx are preserved, the distal ends were destroyed
during weathering. The 1st and 2nd wing phalanges of the left
wing are most likely complete. However, they lie underneath
the skull and therefore make impossible the determination of
the length of these two bones. The 1st wing phalanx shows its
ventral side. The proximal end of the 3rd wing phalanx is pre-
served, however the distal end is missing. By contrary, only
the distal end of the 4th wing phalanx is preserved, it ends in a
very fragile, flattened point. Two vague impressions, from these
missing articular ends are present on the slab; the length of the
3rd and 4th wing phalanx are roughly estimated to 117 mm and
83 mm, respectively. The pteroid was not found.
Hindlimb and pes (Fig. 11). Only the right femur is pre-
served, it is 56 mm long and shorter than the tibia. The shaft of
the femur is slender and slightly bowed, and circular in cross-
Fig. 10. Left humeri of various Triassic pterosaurs and the Liassic Campylo-
gnathoides liasicus (QUENDSTEDT), in anconal view. a) Raeticodactylus filisu-
rensis gen. et sp. nov. (BNM 14524), right humerus is mirrored. b) Eudimorph-
odon ranzii, Holotype (MCSNB 2888), after Wild (1978). c) Eudimorphodon
cf. ranzii (BSP 1994 I 51), after Wellnhofer (2003). d) Eudimorphodon sp., Mi-
lano specimen (MPUM 6009), after Wild (1978). e) Eudimorphodon cf. ran-
zii (MCSNB 2887), after Wild (1978). f) Eudimorphodon sp. (MCSNB 8950),
after Wild (1994). g) Eudimorphodon cromptonellus, Holotype (MGUH VP
3393), right humerus is mirrored, after Jenkins et al. (2001). h) Eudimorph-
odon rosenfeldi, Holotype (MFSN 1797), after Dalla Vecchia (2004). i) Eu-
dimorphodon sp. (MFSN 1922), after Dalla Vecchia (2004). j) Peteinosaurus
zambellii (MCSNB 3359), after Wild (1978). k) Preondactylus buffarinii, Ho-
lotype (MFSN 1770), after Wild (1984). l) Campylognathoides liasicus (CM
11424) after Wellnhofer (1974). The scale bar is in centimetres.
Fig. 9. Drawing of two isolated multicuspid teeth of Raeticodactylus filisuren-
sis gen. et sp. nov. (BNM 14524). a, b) Labial (a) and lingual (b) views of an
originally isolated tooth. c, d) A second isolated tooth (that initially lay in the
mouth of the specimen) in labial (c) and lingual (d) views. Stippling represents
where the teeth are broken. The scale bar is in millimetres.
8 R. Stecher
A new Triassic pterosaur from Switzerland 9
section. The caput femoris has a stout neck. The head is per-
pendicular to the shaft. The shaft of the femur was weathered
along the shaft; it shows that the femur is pneumatized and
hollow. The pneumatic space has been filled with an orange-
red stone matrix. The tibia is very thin, straight, hollow, and
83 mm in length. It is adjacent to the relic of the 1.5 mm wide
fibula. The right tarsus is partially articulated and connected
to the tibia. In addition, the calcaneus, the tarsalia and the four
metatarsals are articulated. The metatarsals are about 32 mm
long. The phalanges are disarticulated. Next to the pes is a toe,
which seems less robust and less strongly bowed. Bones from
the elongate 5th toe, typical of non-pterodactyloid pterosaurs,
are also preserved.
Comparisons. – Comparison with Caviramus (Fig. 12). Fröbisch
& Fröbisch (2006) recently described a new Triassic pterosaur
from Switzerland, Caviramus schesaplanensis, based on a
poorly preserved mandible (PIMUZ A/III 1225). It consists of
three bone fragments with two teeth. According to Fröbisch &
Fröbisch (2006), the teeth of Caviramus show many similarities
with those of Eudimorphodon. It is noteworthy that this mandi-
ble was found in the Alplihorn Member (Kössen Formation) of
the Schesaplana (Northern Calcareous Alps, Canton Grisons).
In other words, Caviramus was found in the same stratigraphi-
cal unit than Raeticodactylus, but in another tectonic nappe.
Unfortunately, the exact stratigraphical and geographical data
where PIMUZ A/III 1225 was found are not mentioned.
The following differences have been established thanks to a
personal examination of Caviramus: PIMUZ A/III 1225 does
not present any quinticuspid tooth; BNM 14524 presents only
seven cup-shaped structures (which might be part of the at-
tachment for the rhamphothecae) on the anterior part of the
mandible, which are hardly visible and smaller; the lower edge
of the mandible of PIMUZ A/III 1125 is convex instead of con-
cave in BNM 14524; parallel to the tooth row, PIMUZ A/III
1225 has large, oval foramens, located every 2nd tooth, BNM
14524 has also such foramen but every 3rd tooth. The following
features are common in Raeticodactylus and Caviramus: a rela-
tively tall mandible; multicuspid teeth; retroarticular process
angled at about 30–35 degrees. Elsewhere, Fröbisch & Fröbisch
(2006) suppose that the dentition of Caviramus is isodont and
that the anterior end of the mandible reveals its original form
but this seems weakly supported. Although Raeticodactylus
could be interpreted as relatively similar to Caviramus at first
glance, the above differences are well supported. In conclusion,
Raeticodactylus and Caviramus are defined as two different
genera. Their precise relationship cannot be established, Cavi-
ramus being too incomplete.
Comparison with other Triassic pterosaurs (Tables 1–3).
Kellner (1996, 2003) and Unwin (2003a) separately investi-
gated the phylogeny and the evolutionary history of ptero-
saurs. Using cladistic analyses they both concluded that Tri-
assic pterosaurs are not related as previously thought. Un-
win (2003a) defined that all Eudimorphodon species belong
to a single family, the Campylognathoididae (including also
Campylognathoides and Austriadactylus). Kellner (1996,
2003) showed that E. ranzii and Campylognathoides liasicus
(QUENSTEDT) are sister taxa belonging to the Campylogna-
thoididae; however, he also suggested that ‘E’. rosenfeldi and
Peteinosaurus zambellii are in a different clade, in other words
that Eudimorphodon species do not necessarily belong to a
single genus or family. It is not the purpose of this paper to
revise Eudimorphodon but the variations observed within
this genus are considered. The following anatomical sections
will detail the cranial and postcranial differences and similari-
ties between Raeticodactylus and other Triassic pterosaurs. A
Fig. 12. Lower jaws of the two Triassic pterosaurs from Switzerland. a) Draw-
ing of the lower jaw of Raeticodactylus filisurensis gen. et sp. nov. (BNM
14524). b) Drawing of the mirrored lower jaw of Caviramus schesaplanensis
(PIMUZ A/III 1225), modified after Fröbisch & Fröbisch (2006). The scale
bar is in centimetres.
Fig. 11. Femora of various Triassic pterosaurs, in lateral view. a) Raeticodacty-
lus filisurensis gen. et sp. nov. (BNM 14524), right femur. b) Eudimorphodon
ranzii, Holotype (MCSNB 2888), left femur, after Wild (1978). c) Eudimor-
phodon cf. ranzii (MCSNB 2887), right femur, after Wild (1978). d) Eudimor-
phodon sp. (MCSNB 8950), left and right femur, after Dalla Vecchia (2002).
e) Eudimorphodon cromptonellus, Holotype (MGUH VP 3393), right fe-
mur, after Jenkins et al. (2001). f) Peteinosaurus zambellii (MCSNB 3496),
left femur, after Dalla Vecchia (2003b). g) Preondactylus buffarinii, Holotype
(MFSN 1770), after Wild (1984). The scale bar is in centimetres.
comparison with the holotypes of Triassic pterosaurs having
multicuspid teeth is summarized in Table 1.
Skull and mandible. The overall skull morphology of Raeti-
codactylus is different from that of Eudimorphodon (MCSNB
2888) and Austriadactylus (SMNS 56342) (Fig. 13). The size
– i.e. the wingspan – in comparison to the skull length makes it
clear that Raeticodactylus has a relatively small skull compare
to other Triassic pterosaurs (BNM 14524: 14.2; E. sp., MPUM
6009: 11.3; E. ranzii, MCSNB 2888: 8.3–11.1; A. cristatus, SMNS
56342: 10.9). The length/height ratio of the skull of Raeticodac-
tylus (without the sagittal crest) is relatively high with a value
of 3.84 (E. ranzii, MCSNB 2888: 2.84; E. sp., MPUM 6009: 2.41;
A. cristatus, SMNS 56342: 3.0).
The comparison of the skull-orbit length index shows that the
orbit of Raeticodactylus is relatively small, only Austriadacty-
lus has maybe a smaller orbit (BNM 14524: 21.4%; E. ranzii,
MCSNB 2888: 24.5%; E. sp., MPUM 6009: 28.5%; A. cristatus,
SMNS 56342: 20.8%). The relatively small orbit also suggests
that Raeticodactylus was fully grown (Wild 1978). While the
narial fenestra of Eudimorphodon (MCSNB 2888) and Raeti-
codactylus show certain similarities, the antorbital fenestra of
BNM 14524 is larger and with a different shape (Fig. 13). Only
Raeticodactylus and Austriadactylus have a bony, sagittal crest
on the skull. Comparing the mandibles of Triassic pterosaurs,
Raeticodactylus mandible is taller than that of E. ranzii; this
is obvious using the length to height ratio (BNM 14524: 8.8;
10 R. Stecher
Austriadactylus
cristatus
(Holotype, SMNS
56342)
Caviramus schesap-
lanensis
(Holotype, PIMUZ
A/III 1225)
Eudimorphodon
cromptonellus
(Holotype, MGUH
VP 3393)
Eudimorphodon
ranzii
(Holotype, MCSNB
2888)
Eudimorphodon
rosenfeldi
(Holotype, MFSN
1797)
Raeticodactylus filisu-
rensis gen. et sp. nov.
(Holotype, BNM
14524)
Formation Seefeld Schichten Kössen Formation /
Alplihorn-Member
of the Lechtal nappe
Fleming Fjord For-
mation / Orsted Dal
Member
Zorzino-Calcari basic part of the
Dolomia di Forno
Formation
Kössen Formation /
Alplihorn-Member of
the Ela nappe
Age late Norian (late
Alaunian–early
Sevatian)
late Norian–early
Rhaetian
Norian–Rhaetian middle Norian, late
Alaunian
middle Norian
(Alaunian 2–3)
late Norian
(Sevatian) or early
Rhaetian
Bony crest 20 mm high; long
crest
– no no – 21 mm high; crest on
the preamaxilla and
naris
Largest skull opening narial fenestra – – orbit orbit orbit
Skull-orbit length index 20.8% – – 24.5% – 21.4%
Mandible length / height
ratio
– 7.4–8.7 – 14.6 more than 13.8 8.8
Shape of the anterior end
of the mandible…
– … forms a rounded
edge
– … is curved
ventrally
– … is keel-shaped
and ends in a dorsally
deflected tip
Dentition and teeth heterodont: mono-
cuspid and triangular
formed multicuspid
(up to 12 denticles
on each cutting
edge) teeth
isodont: at least
tri- and four-cuspid
teeth in the lower
jaw
heterodont: mono-,
tri-, four- and five-
cuspid teeth
heterodont: mono-,
tri-, five-cuspid teeth
heterodont: at least
mono- and five-
cuspid teeth
heterodont: mono-,
tri-, four- and five-
cuspid teeth
Comparison of teeth size
between upper and
lower jaws
teeth of the upper
jaw are larger
– teeth of the lower
jaw are partially
smaller
same size – teeth of the upper jaw
are larger
Teeth of the upper jaw… … have small gaps
between them
– … have wide gaps
between them
… have no gaps
between them
… have small gaps
between them
… have small gaps
between them
Teeth of the lower jaw… … are aligned in
one row and have
small gaps between
them
– … are aligned in
one row and have
wide gaps between
them
… are standing very
close together in
one row, but do not
overlap
… are aligned in
one row and have
small gaps between
them
… are standing very
close together and
overlap partially
Number of teeth in the
upper jaw
17–25 – 12 or more 29 – 17
Number of teeth in the
lower jaw
– at least 12, maxi-
mum 17
12 (?) 28 – 22
Table 1. General data of Raeticodactylus filisurensis gen. et sp. nov. and various holotypes of Triassic pterosaurs having multicuspid teeth.
A new Triassic pterosaur from Switzerland 11
E. ranzii, MCSNB 2888: 14.6; E. sp., MPUM 6009: 13.1). A fur-
ther difference is the anterior tip of the mandible, the anterior
end of Eudimorphodon (MCSNB 2888) is curved ventrally,
whereas that of Raeticodactylus is keel-shape and ends in a
dorsally deflected tip.
Dentition. Like all other Eudimorphodon specimens and
Austriadactylus, Raeticodactylus has a heterodont dentition.
The anterior teeth are monocuspids whereas the posterior
ones are multicuspids. Raeticodactylus has the largest differ-
ence in the number of teeth between the upper and lower jaws
compared to Eudimorphodon (MCSNB 2888, MPUM 6009).
The mandible of Raeticodactylus has five supplementary teeth
than the upper jaw; in contrast, the upper jaw of Eudimorph-
odon has a supplementary tooth. In the posterior part of the
upper and lower jaws, E. ranzii has equally sized multicuspid
teeth (Wild 1978). In comparison, Raeticodactylus has slightly
bulbous teeth in the upper jaw and slightly shorter teeth in the
mandible. The maxillary teeth of Raeticodactylus are aligned
in one row and have small gaps between them; the mandible
exhibits very narrowly standing teeth that partially overlap.
These characteristics are only present on BSP 1994 I 51 (E. cf.
ranzii) and to a certain degree on MPUM 6009 (E. sp.). Raeti-
codactylus has no enlarged pseudo-unicuspid teeth under the
processus ascendens, like in Eudimorphodon (MCSNB 2888).
At this position, Raeticodactylus has multicuspid teeth, which
are slightly larger and of similar form than the surrounding
Austriadactylus
cristatus
(Holotype, SMNS
56342)
Caviramus schesap-
lanensis
(Holotype, PIMUZ
A/III 1225)
Eudimorphodon
cromptonellus
(Holotype, MGUH
VP 3393)
Eudimorphodon
ranzii
(Holotype, MCSNB
2888)
Eudimorphodon
rosenfeldi
(Holotype, MFSN
1797)
Raeticodactylus filisu-
rensis gen. et sp. nov.
(Holotype, BNM
14524)
Monocuspid teeth with
fang-like function in the
upper jaw
5 – 1 or 2 4 (3rd teeth is tri-
cuspid)
–4
Monocuspid teeth with
fang-like function in the
lower jaw
–012–3
Number of pseudo-uni-
cuspid teeth under the
processus ascendens
1 or 2–02–0
First five cusped tooth in
the upper jaw
––5
th place 17th place (left), 18th
place (right)
–9
th place
First five cusped tooth in
the lower jaw
––10
th place (left), 8th
place (right)
6th place (left), 8th
place (right)
–10
th place
Diastema – – between the 3rd and
4th tooth in the upper
jaw and between the
9th and 10th tooth in
the lower jaw
no – large between the 4th
and 5th tooth in the
upper jaw
Shape of the humerus slightly bowed with
a wide deltopectoral
crest
– slightly bowed with
subtriangulary delto-
pectoral crest
bowed and robust
with rectangular del-
topectoral crest
slightly bowed and
slender with a small,
perpendicular delto-
pectoral crest
straight, slender,
relatively long, with a
well-developed sub-
rectangular and wide
deltopectoral crest
Humerus length / shaft
diameter ratio
– – 13 7.4 10.4 18.2
Humerus length / width
ratio
– – 3.5 2.2 3.0 3.7
(humerus + ulna) / (femur
+ tibia) ratio
– – 0.95 1.23 1.05 1.34
(∑ humerus + ulna +
metacarpal IV / ∑ femur
+ tibia + average of
metatarsals) × 100
– – 90.0% – 103.8% at least 134.3%
Data from Dalla Vecchia et al.
(2002)
Fröbisch & Fröbisch
(2006)
Jenkins et al.
(2001)
Zambelli (1973),
Wild (1978)
Dalla Vecchia
(1995)
This study
Table 1.
12 R. Stecher
teeth. The mandible of Raeticodactylus has teeth along 55.4%
of its length. In comparison with other Triassic pterosaurs
this is relatively low, e.g. in Eudimorphodon (MCSNB 2888:
66.2%; MPUM 6009: 75.8%).
The rough-textured and rippled surface at the ends of the jaws
of Raeticodactylus indicates a keratinized, beak-like edge on
the jaws (see above), as Wild (1978) also suggested for Eu-
dimorphodon (MCSNB 2888, MPUM 6009).
Humerus (Fig. 10). In contrast to Eudimorphodon (MCSNB
2888) and other humeri of Triassic pterosaurs, Raeticodactylus
has a very straight and remarkably slender humerus. Its length
to the shaft diameter ratio is 18.2. This ratio is clearly higher than
from other Triassic pterosaurs (E. ranzii, MCSNB 2888: 7.8; E.
rosenfeldi, MFSN 1797: 10.4; E. cromptonellus, MGUH VP 3393:
13). The length to width ratio of the BNM 14524 humerus is 3.7
and higher than other Triassic pterosaurs (E. ranzii, MCSNB:
2.2; E. rosenfeldi, MFSN 1797: 3.0). Only E. cromptonellus shows
a similar value of 3.5. The humerus has a wide subrectangular
deltopectoral crest, similar to that of E. rosenfeldi. The proxi-
mal articular end of the humerus exhibits two concave saddles.
This is an apomorphic feature (Unwin 2003a) known in Juras-
sic non-pterodactyloid pterosaurs (Campylognathoides, Rham-
phorhynchus). By contrary, E. ranzii (MCSNB 2888) shows the
plesiomorphic state with a single concave saddle.
Femur (Fig. 11). BNM 14524 has a caput femoris that lies
perpendicular to the shaft. Wellnhofer (1993) stated that all Tri-
assic pterosaurs have a caput femoris that faces diagonally up-
wards, which is also a feature of Jurassic and Cretaceous ptero-
saurs. However, it is noteworthy that the specimens MCSNB
8950 (identified as Eudimorphodon sp., Wild 1994), MGUH
VP 3393 (E. cromptonellus, holotype) and MCSNB 3496 (Pe-
teinosaurus zambellii) also show a distinct perpendicular caput
femoris. Interestingly, this is a very common feature in dino-
saurs (O. Rauhut, pers. comm. 2007), it is now identified in some
Triassic pterosaurs.
Comparison of skeletal lengths (Tables 1–3; Fig. 14). The
skull length of Raeticodactylus is only slightly longer than in the
adult specimen of Eudimorphodon ranzii (MCSNB 2888), their
postcranial skeletons, however, clearly differ in size (Table 2).
Raeticodactylus was considerably larger than MCSNB 2888;
with longer and relatively thinner bones, Raeticodactylus was
more gracile.
s m h u mcIV fe ti wph1 wph2 wph3 wph4 Wing-
span
Preondactylus buffarinii
(Holotype, MFSN 1770) 2, 4, 7
56* 54 32 42 19 34 49.5 30.8 39 39 28* 45–50
?Peteinosaurus zambellii
(MCSNB 3359) 5
– – 38.5 48 17 37 49 43 43 46.5 34.8 60
Austriadactylus cristatus
(Holotype, SMNS 56342) 8
110 – 83 – – – – – 101 103.5 85.5* 120
Eudimorphodon ranzii
(Holotype, MCSNB 2888) 1, 3
90* 74.5 47 65 29 41 50* 80* – – – 75–100
Eudimorphodon cf. ranzii
(MCSNB 2887) 1, 9
– – 28 38 14* 21.2 28.5 39.5 36.5* – – –
Eudimorphodon sp.
(MPUM 6009) 1, 3, 9
40 34 26 36 10.5 18.5 25* 38.5 33* 36.2 34* 45
Eudimorphodon cf. ranzii
(BSP 1994 I 51) 5
– – 40 – – – 57.7 52.9 – – – 70–80
Eudimorphodon sp.
(MCSNB 8950) 3, 5, 9
– – 26 33.5 9.3 19.6 25.5 33 35.2 36.2 32 45
Eudimorphodon rosenfeldi
(Holotype, MFSN 1797) 6
– – 40.5 55 21 37 54.2 64 58.2 63.2 51.5 65–70
Eudimorphodon cromptonellus
(Holotype, MGUH VP 3393) 10
– – 18.15 20.15 8.4 19.7 20.5* 18* 20.5 20.5* – 24
Raeticodactylus filisurensis gen.
et sp. nov.
(Holotype, BNM 14524)
95 84 82 106 – 56 84 113 109 117* 83* 135
Table 2. Overview of skeletal lengths (mm) and wingspan (cm) in Raeticodactylus filisurensis gen. et sp. nov. and various Triassic pterosaurs.
* estimated
Abbreviations: fe, femur; h, humerus; m, mandible; mcIV, wing metacarpal (IV); s, skull; ti, tibia; u, ulna; wph1–4, wing phalanges 1–4.
Data sources: 1 Wild (1978); 2 Wild (1984); 3 Wild (1994); 4 Wellnhofer (1993); 5 Wellnhofer (2003); 6 Dalla Vecchia (1995); 7 Dalla Vecchia (1998); 8 Dalla Vecchia
et al. (2002); 9 Dalla Vecchia (2003a); 10 Jenkins et al. (2001).
A new Triassic pterosaur from Switzerland 13
There are many differences between the bone length ra-
tios (Table 3) of Raeticodactylus, E. ranzii (MCSNB 2888),
E. rosenfeldi (MFSN 1797) and E. cromptonellus (MGUH VP
3393). Raeticodactylus shows close similarities with MCSNB
2887, (E. cf. ranzii), a highly disarticulated specimen, only
partially preserved and lacking skull. Raeticodactylus is ap-
proximately three times larger in size than MCSNB 2887. Two
main conclusions from the comparison of Raeticodactylus with
Eudimorphodon, Peteinosaurus and Preondactylus are drawn:
Raeticodactylus exhibits the shortest femur compared to the
tibia, and also the longest humerus compared to the femur.
The forelimb (humerus + ulna) to hindlimb (femur + tibia) ra-
tio (Table 1) is very high in Raeticodactylus with 1.34 (E. ranzii,
MCSNB 2888: 1.23; E. rosenfeldi, MFSN 1797: 1.05; E. cromp-
tonellus, MGUH VP 3393: 0.95). It is similar to that of E. cf.
ranzii (MCSNB 2887) and E. sp. (MCSNB 8950) with a value
of 1.32; only the ratio of E. sp. (MPUM 6009) is higher with a
value of 1.43. Jenkins et al. (2001) also compared the dimen-
sions of the forelimbs and hindlimbs in various pterosaurs,
using the length of the metacarpal IV and the average length
of metatarsals. The ratio was as follows: (∑ humerus + ulna
+ metacarpal IV / ∑ femur + tibia + average of metatarsals)
× 100. Jenkins et al. (2001) determined it in the following Tri-
assic pterosaurs: E. cromptonellus (90%), Preondactylus buf-
farinii (95%), Peteinosaurus zambellii (100%) and E. sp. (MC-
SNB 8950) (130%). These values could be completed with
E. rosenfeldi (103.8%). The ratio for the specimens MCSNB
2888 (E. ranzii) and SMNS 56342 (A. cristatus) cannot be de-
termined. The metacarpal IV of BNM 14524 is not preserved,
however, as this metacarpal is never shorter but rather longer
than the metacarpal III in oth er Triassic pterosaurs, the length
of the metacarpal III can be used as a proxy. The resulting ratio
is of at least 134.3%, Raeticodactylus reveals the highest value
of this compar ison.
With a wingspan of 135 cm (Table 2), Raeticodactylus is one of
the largest Triassic pterosaur known so far. Only two other Tri-
assic specimens suggest the existence of even larger pterosaurs
during the early history of this group: three wing phalanges
from an Italian specimen, MCSNB 4562 (Pterosaur indet.), in-
dicate a wingspan of 150–160 cm (Padian 1980); an isolated 4th
wing phalanx from an Italian specimen, MFSN 19836 (Ptero-
saur indet.), indicates a wingspan of approximately 175 cm
(Unwin 2003b).
An overall graphic comparison of skeletal lengths is pre-
sented in Figure 14. It well summarizes that even though Rae-
ticodactylus has some similarities with some Triassic specimens
(e.g. MCSNB 2887, Eudimorphodon cf. ranzii), this new genus
is clearly distinct from other Triassic pterosaurs.
Functional morphology and palaeoecology
Bony, sagittal cranial crest
Bony cranial crests are known from many pterosaurs. Ac-
cording to Wellnhofer (1993), there are three groups of
crests: short or long crests on the posterior part of the skull
(e.g. Pteranodon, Dsungaripterus); long and low crests on the
middle part of the skull (e.g. Germanodactylus, Gnathosaurus,
Ctenochasma and Dsungaripterus); high crests on the anterior
part of the snout, which look like an inverse keel (e.g. Orni-
thocheirus).
Bony sagittal crests are only known from Triassic pterosaurs in
Austriadactylus cristatus and now in Raeticodactylus filisuren-
sis. Both specimens have the crest on the premaxilla, but that
of A. cristatus elongates all along the skull. One could hypoth-
esize that the bony crests moved to a more posterior position
during the Jurassic and Cretaceous, although preorbital sagittal
crests are very common in the Cretaceous (e.g. Ornithocheirus).
From this point of view the crests do not reflect an evolutionary
trend in pterosaur evolution, the premaxillary bony crests could
rather indicate a functional adaptation to a particular lifestyle.
Until now, the function of these crests is not fully understood,
different explanations are found in the literature: a structure
linked to the sexual dimorphism (this has been argued for Pter-
anodon) (Bennett 1992); a display structure (Kellner 2002);
imprints of bloodstreams on the bony crests show that they
were strongly supplied with blood, they could have regulated
the body temperature (e.g. in Thalassodromeus) (Campos &
Kellner 1996, Kellner 2002); an aerodynamic function (Kellner
2002); an hydrodynamic role to feed by skimming, notably by
analogy with the jaw anatomy of the modern skimming bird
Rhynchops (Wellnhofer 1993).
It could be argued that Raeticodactylus caught fishes by
skimming, as suggested by Wild (1978) for Eudimorphodon
ranzii. Beyond the presence of premaxillary and mandibular
crests, the following features (Kellner 2002) also suggest that
R. filisurensis could be a skimming pterosaur: laterally com-
pressed upper and lower jaws with a bladelike horny covering
(rhamphothecae); large cervical vertebrae for an important
neck musculature; large adductor mandible complex muscu-
lature; several foramina on the tip of the premaxilla and of
the mandible indicate that this region was well irrigated by
blood vessels and likely well supplied by nerves; due to the
possibility that the snout was dived into the water at least for
a short time, the posterior position of the narial fenestra of
BNM 14524 in relation to the skull could have been advanta-
geous. A recent study by Humphries et al. (2007), however,
strongly challenged the hypothesis of skimming pterosaurs,
even in smaller forms.
Specialized dentition
Dalla Vecchia (2003a) mentioned that most Triassic pterosaurs
have multicuspid or serrated teeth. Wellnhofer (1993) stated
that multicuspid teeth do not reflect a primitive feature of rep-
tiles, but rather a specialization to its environment and there-
fore to prey (e.g. Eudimorphodon as a piscivore). Multicuspid
teeth are surely useful to penetrate and cut hard food (e.g.
ganoid fishes, crustaceans, large insects or other animals with
a robust exoskeleton) into small pieces (Dalla Vecchia 2003a).
14 R. Stecher
u/h h/mcIV u/mcIV h/fe h/ti u/fe u/ti ti/fe fe/mcIV ti/mcIV wph1/h wph1/u wph1/
mcIV
wph1/
fe
wph1/
ti
wph2/
wph1
wph3/
wph2
wph3/
wph4
wph3/
wph1
Preondactylus buf-
farinii (Holotype;
MFSN 1770)
1.31 1.68 2.21 0.94 0.65 1.24 0.85 1.46 1.79 2.61 0.96 0.73 1.62 0.91 0.62 1.27 1 1.39* 1.27
?Peteinosaurus zam-
bellii (MCSNB 3359)
1.25 2.26 2.82 1.04 0.79 1.3 0.98 1.32 2.18 2.88 1.12 0.9 2.53 1.16 0.88 1 1.08 1.34 1.08
Austriadactylus cris-
tatus (Holotype;
SMNS 56342)
––––––––––––––––1.02 – –
Eudimorphodon
ranzii (Holotype,
MCSNB 2888)
1.38 1.62 2.24 1.14 0.94* 1.58 1.3 1.22* 1.41 1.72* 1.7* 1.23* 2.75* 1.95* 1.6* ––––
Eudimorphodon cf.
ranzii (MCSNB
2887)
1.36 2 2.27 1.32 0.98 1.79 1.33 1.34 1.57 2.03 1.43* 1.05* 2.82 1.89* 1.39* 0.91* – – –
Eudimorphodon sp.
(MPUM 6009)
1.37 2.5 3.43 1.42 1.05* 1.95 1.44* 1.35* 1.76 2.38* 1.48 1.04 3.57 2.03 1.5* 0.88 1.1* 1.06 0.96*
Eudimorphodon cf.
ranzii
(BSP 1994 I 51)
––––0.69 –––––1.32 – – – 0.92 ––––
Eudimorphodon sp.
(MCSNB 8950)
1.29 2.89 3.72 1.33 1.02 1.71 1.34 1.27 2.18 2.78 1.27 1.01 3.78 1.73 1.29 1.04 1.02 1.12 1.06
Eudimorphodon
rosenfeldi
(Holotype, MFSN
1797)
1.31 1.93 2.62 1.13 0.75 1.49 1.02 1.46 1.76 2.58 1.58 1.16 3.05 1.73 1.18 0.91 1.09 1.23 0.99
Eudimorphodon
cromptonellus
(Holotype, MGUH
VP 3393)
1.11 2.16 2.39 0.92 0.89* 1.02 0.98* 1.04* 2.34 2.44* 0.99* 0.9* 2.14* 0.91* 0.88* 1.14* 1* – 1.14*
Raeticodactylus filisu-
rensis gen. et sp. nov.
(Holotype, BNM
14524)
1.29 – – 1.46 0.98 1.89 1.26 1.5 – – 1.38 1.07 – 2.02 1.35 0.96 1.07* 1.41* 1.04*
* estimated
Abbreviations: fe, femur; h, humerus; mcIV, wing metacarpal (IV); ti, tibia; u, ulna; wph1–4, wing phalanges 1–4.
Table 3. Bone length ratios of Raeticodactylus filisurensis gen. et sp. nov. and various Triassic pterosaurs. The ratios were calculated with the measurements of Table 2.
A new Triassic pterosaur from Switzerland 15
Fastnacht (2005) noted that Eudimorphodon, with its small
multicuspid teeth, could better hold very small food items, in
contrast to Preondactylus.
Raeticodactylus filisurensis also has a heterodont dentition.
In the anterior part are the long, monocuspid and fang-like
teeth. On the labial side the enamel is more or less smooth, but
has very strong wrinkles on the lingual side. The wrinkles are
aligned very close to each other and show bowed ridges, which
are partially bifurcated. The function of these enamel wrinkles
can be interpreted as follows: better catching and holding (fixa-
tion through the enamel wrinkles) of slick nutrition (fish); in-
creased stability of the tooth by construction ridges (less wide
ridges or many fine ridges) like in R. filisurensis with its fine
teeth.
One isolated fang tooth of E. cf. ranzii (BSP 1994 I 51) also
reveals bowed enamel wrinkles on its lingual side. It is simi-
lar to BNM 14524, however, not so strongly developed as for
R. filisurensis. The multicuspid teeth are equal in size in the up-
per and lower jaws. The wear facets on the labial side of the
mandible teeth indicate a displaced contact between the upper
and lower jaws. When the snout was closed, the upper multi-
cuspid teeth were located on the outside whereas those of the
mandible were inside. The cutting edges of some teeth of the
upper jaw, distally and mesially in convex form, show a thick-
ened bulge. These features indicate a cutter-like function of the
dentition. Wellnhofer (1993) pointed out that in addition to the
teeth, some pterosaurs (e.g. Rhamphorhynchus) had keratin-
ized, beak-like edges on the tips of the jaws that were useful for
fishing. Based on the structure of the bones, it is very likely that
Raeticodactylus had such a soft tissue structure on the anterior
part of its snout. These keratinized edges would have helped in
cutting prey. The diastema between the monocuspid and mul-
ticuspid teeth in the upper jaw shows an analogous feature to
other animals. Some theropod dinosaurs (e.g. Baryonyx, Irrita-
tor) and recent crocodiles reveal such diastema at similar posi-
tion. This could indicate that Raeticodactylus also had a grasp-
ing and hunting style.
Conclusions
Raeticodactylus filisurensis gen. et sp. nov. is so far the best pre-
served pterosaur from Switzerland. This basal non-pterodac-
tyloid pterosaur is very likely late Norian in age. Comparisons
with other Triassic pterosaurs show clearly that it is a distinct
genus.
Raeticodactylus filisurensis has a dentition that character-
izes a grasping and holding preying style. In the anterior part
of the dentition are monocuspid teeth (with strong enamel
wrinkles on the lingual side) with a fang like function. In the
posterior part of the dentition are tricuspid, quadricuspid and
quinticuspid teeth with a crack or cutting-like function. The
bony cranial crest of R. filisurensis is different from that in
Austriadactylus cristatus. It is linked to the keel-like increase
of the mandible and interpreted as a skim-feeder adaptation.
With the discovery of R. filisurensis, two Triassic pterosaurs
are known with bony sagittal cranial crests, suggesting that
such crests were more common during the Late Triassic than
previously thought. The femur of R. filisurensis is quite unusual
with a caput femoris perpendicular to the shaft. Comparisons
of skeletal measurements show that R. filisurensis was a grac-
ile flyer with an important wingspan (at least 135 cm) for the
Late Triassic.
Finally, R. filisurensis confirms that pterosaurs were diverse
and highly specialized flying reptiles since the beginning of
their long history. It gives additional evidence to the hypothesis
of Unwin (2003a) suggesting that there was a significant radia-
tion of basal pterosaurs during the Late Triassic.
Acknowledgements
I especially thank Ulrich Schneppat (BNM Chur) for his faithful and con-
tinuous help to solve different problems. Many thanks to Dr. Jürg Paul Mül-
ler (BNM, Chur) who provides me a stereomicroscope for observation and
preparation. Thanks to Dr. Fabio Dalla Vecchia (Italy) for discussions about
Eudimorphodon. Many thanks to Dr. Oliver Rauhut and Dr. David Hone
(BSPG, Munich) for fruitful discussions, and the chance to examine specimen
BSP 1994 I 51. Many thanks to Dr. Heinz Furrer (PIMUZ, Zurich) for access
Fig. 13. Comparison of Triassic pterosaur skulls. a) Eudimorphodon ranzii
(MCSNB 2888), after Wild (1978). b) Raeticodactylus filisurensis gen. et sp.
nov. (BNM 14524). c) Austriadactylus cristatus (SMNS 56342), after Dalla Vec-
chia et al. (2002). The scale bar is in centimetres.
16 R. Stecher
to the holotype of Caviramus. For the correct proofs of a previous version of
the manuscript in German, I especially thank my colleague Katrin Bebié. For
the correct proofs of the first English version of the manuscript, I thank my
colleague Dr. Carlo Casty. Special thanks to Dr. David Hone (BSPG, Munich)
for many suggestions and improvements of the English. Many thanks also to
Prof. Kevin Padian (Univ. of California, Berkeley), Dr. Christian A. Meyer
(NM Basel) and an anonymous reviewer for their constructive comments.
Dr. Christian A. Meyer also helped me for the sedimentological interpreta-
tion. Many thanks to the Editor Dr. Jean-Paul Billon-Bruyat for his great help
with the final version of the manuscript. Most of all I am thankful to my wife
Irmgard Stecher and our children Gianna and Claudio for their understanding
of my hobby. Finally, I warmly thank the KSPA (Kommission des Schweizeri-
schen Paläontologischen Abhandlungen) for its sponsorship of the additional
pages.
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Manuscript received March 14, 2007
Revision accepted January 22, 2008
Editorial handling: J.-P. Billon-Bruyat
