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A new rhamphorhynchid pterosaur (Pterosauria) from Jurassic deposits of Liaoning Province, China


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Compared to pterosaurs from the Early Cretaceous from China, Late Jurassic pterosaurs are relatively rare. A new rhamphorhynchid pterosaur, Orientognathus chaoyngensis gen. et sp. nov., is erected based on an incomplete skeleton from the Upper Jurassic Tuchengzi Formation of Chaoyang, Liaoning Province, China. It is identified by the following characters: the toothless tip of the lower jaw is slightly pointed; the length ratio of wing metacarpal to humerus is 0.38, the ulna is shorter than each wing phalanx and the tibia is nearly equal to femur in length. A phylogenetic analysis recovers Orientognathus chaoyngensis as a rhamphorhynchid pterosaur. Orientognathus chaoyngensis is perhaps the youngest Jurassic pterosaur from western Liaoning Province of China.
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Accepted by S. Brusatte: 24 Nov. 2014; published: 16 Jan. 2015
ISSN 1175-5326 (print edition)
(online edition)
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zoota xa
A new rhamphorhynchid pterosaur (Pterosauria) from Jurassic deposits
of Liaoning Province, China
Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037 China; Key Laboratory of Stratigraphy and Paleontol-
ogy, Ministry of Land and Resources. E-mail:; Tel. (fax). 00861068999707
Henan Geological Museum, Zhengzhou 450016, Henan Province, China
Cores and Samples Center of Land and Resources, China Geological Survey, Sanhe 065201, Hebei Province, China
Subdepartment of Evolution and Development, Department of Organismal Biology, Evolutionary Biology Centre, Uppsala University,
Norbyvägen, 18A
, SE-75236 Uppsala, Sweden
Compared to pterosaurs from the Early Cretaceous from China, Late Jurassic pterosaurs are relatively rare. A new rham-
phorhynchid pterosaur, Orientognathus chaoyngensis gen. et sp. nov., is erected based on an incomplete skeleton from
the Upper Jurassic Tuchengzi Formation of Chaoyang, Liaoning Province, China. It is identified by the following charac-
ters: the toothless tip of the lower jaw is slightly pointed; the length ratio of wing metacarpal to humerus is 0.38, the ulna
is shorter than each wing phalanx and the tibia is nearly equal to femur in length. A phylogenetic analysis recovers Ori-
entognathus chaoyngensis as a rhamphorhynchid pterosaur. Orientognathus chaoyngensis is perhaps the youngest Juras-
sic pterosaur from western Liaoning Province of China.
Key words: Upper Jurassic, Tuchengzi Formation, Pterosauria, Rhamphorhynchidae
Pterosaurs from the Early Cretaceous Jehol Biota of western Liaoning and its surrounding areas are numerous, but
those from the pre-Early Cretaceous deposits of China are rarer, although recently they have been increasing both
in number and variety. The most important discovery among these Jurassic forms is Darwinopterus, a transitional
form with a cranium and cervical series that is of typical pterodactyloid morphology, whereas the remainder of the
postcranial skeleton is almost identical to that of typical long-tailed non-pterodactyloid pterosaurs (Lü et al. 2010a;
2011a). This taxon is pivotal for understanding the evolution of pterodactyloid pterosaurs, and their characteristic
anatomy, from more basal forms. Another spectacular discovery is a female Darwinopterus preserved with an egg,
which has implications for understanding sexual dimorphism and reproductive biology in pterosaurs (Lü et al.
2011b). Other Middle Jurassic pterosaurs from China include the non-pterodactyloid Changchengopterus (Lü
2009); the rhamphorhynchids Pterorhynchus (Czerkas and Ji 2002), which is regarded as close to wukongopterid
pterosaurs (Andres et al. 2014), and Qinglongopterus (Lü et al. 2012); the scaphognathines Fenghuangopterus (Lü
et al. 2010b), Jianchangnathus (Cheng et al. 2012; = Scaphognathus, Bennett 2014), Jianchangopterus (Lü and
Bo 2011c); and the anurognathid Dendrorhynchoides mutodungensis (Lü and Hone 2012).
Here we report a new pre-Cretaceous pterosaur, a rhamphorhynchid designated as the new genus and species
Orientognathus chaoyangensis, from the possible Late Jurassic deposits of Chaoyang, Liaoning Province, of north-
eastern China (Fig. 1). A phylogenetic analysis recovers Orientognathus chaoyangensis as a rhamphorhynchid
pterosaur. The discovery of Orientognathus chaoyangensis provides important new information on the
evolutionary history of the Rhamphorhynchinae and on pterosaur evolution in the Jurassic more generally.
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FIGURE 1. The map of the fossil locality. Solid pentagon represents the fossil site.
Geological setting
The specimen was collected by a local farmer. One of the authors (JL) later visited the fossil locality, guided by the
farmer. Because the specimen was not excavated by a technically accomplished collector, it was heavily damaged
during excavation. During the visit to the site, JL and the collector found a fragment of black bone from a
pterosaur, which is identical to the present specimen in color. Thus, we are confident that the locality the farmer
pinpointed is the locality where the specimen described here was collected. The specimen comes from gray-green
tuffaceous shales (Fig. 2), which based on lithology are most likely belonging to the second member of the Upper
Jurassic Tuchengzi Formation (Bureau of Geology and Mineral Resources of Liaoning Province, 1989).
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FIGURE 2. The strata of the fossil site.
Systematic paleontology
Pterosauria Kaup, 1834
Breviquartossa Unwin, 2003
Rhamphorhynchidae Seeley, 1870 (sensu Unwin, 2003)
Rhamphorhynchinae Nopcsa, 1928 (sensu Unwin, 2003)
Orientognathus chaoyangensis gen. et sp. nov.
Holotype. An incomplete skeleton (41HIII-0418), housed at the Henan Geological Museum, Zhengzhou, Henan
Type locality and horizon. Jucaigou, Dongshan Village (41º13'9"N, 120º5'51"E), Shengli Town, Chaoyang
County, Liaoning Province; most likely the Tuchengzi Formation (Bureau of Geology and Mineral Resources of
Liaoning Province, 1989) (Figs. 1–2).
Etymology. “Orient” derives from the Latin word oriens meaning "east"; “gnathus” is derived from Greek,
meaning “jaw”. Orientognathus means “pterosaur from the east”. The specific name refers to where the specimen
was found, in Chaoyang.
Diagnosis. A large rhamphorhynchine pterosaur with a wing span about 1.1 meters, bearing the following
unique combination of characters: ratio of wing metacarpal to humerus 0.38 (it is smaller than those in other basal
pterosaurs); the ulna shorter than each wing phalanx (similar to basal pterosaurs such as Preondactylus and
Dimorphodontidae, but it is reversed condition in Scaphognathus); the ulna shorter than tibia (ratio of ulna to tibia
0.92; it is greater than 1 in other rhamphorhynchine pterosaurs); tibia nearly equal to femur in length; length ratio
of pteroid to humerus 0.21.
Description and comparisons. The specimen is not well preserved (Fig. 3; Table 1). The wing span is
estimated to be 1.1 meters. The individual is not ontogenetically mature, based on its unfused extensor tendon
process of the first wing phalanx, and lack of fusion in the scapulocoracoid and pelvis. The skull was heavily
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damaged before burial. All the elements near the braincase are scattered, and it is difficult to identify them. Only
the lower jaws and some teeth can be clearly observed. The third and fourth wing phalanxes were destroyed or lost
as a result of excavation.
TABLE 1. Measurements (mm) of Orientognathus chaoyangensis gen. et sp. nov. (41HIII-0418).
There are about 18 teeth distributed near the rostral portion of the skull. The nearly triangular shaped bone
bearing two teeth is identified as a maxilla in which the teeth are orientated anteroventrally. There is a plate-like
bone with one thickened margin and one articular end, which may be the quadrate. Some teeth are still in the tooth
sockets of the maxillae and dentary. The anterior teeth are relatively small (9 mm in length) and slightly curved,
whereas the posterior teeth are large (20 mm) and nearly straight. The cross-sections of the teeth are oval. The
lower jaw is deep with parallel dorsal and ventral margins. The left ramus of the lower jaw is preserved in situ, but
the right ramus is upside down, which may indicate that the mandibular symphysis was short and the two dentaries
were weakly joined together. The ventral margin of the lower jaw is straight. The distal end of the lower jaw is
short and pointed in lateral view (Fig. 4a).
There are four cervical vertebrae preserved. Although they are poorly preserved, the exposed lateral surface
exhibits clear pneumatic foramina. The dorsal vertebrae are short, and as with the cervical vertebrae, their detailed
structures are not clear. The dorsal ribs are slender. There are four sacral vertebrae (inferred by their position
between the preacetabular processes of the ilia), although they are slightly displaced. The anterior two are fused
and the posterior two are not fused, corroborating the juvenile status of the specimen. The tail is missing its distal
portion. The first two preserved caudal vertebrae are square in shape dorsolateral view. Beginning with the third
caudal, the caudal vertebrae are completely encased by thread-like zygapophyses and elongated haemal arches
(Fig. 4e).
The scapula and coracoid are not fused, which further corroborates the immature ontogenetic stage of the
individual. These bones are not well-preserved. The distal end of the coracoid is missing, and thus its length ratio is
not clear. The left humerus is missing its distal portion and a small piece of proximal portion, but there is a full
impression of this bone in the matrix. Thus, the length and the shape of the humerus can be restored. The distal
portion of the humerus is curved, but this may be an artefact of preservation. The remaining part of the shaft is
straight. The deltopectoral crest is located proximally and is semicircular in shape. Its ventral margin is rounded.
The ulna and radius are displaced at their proximal portions, but the impressions of their distal ends allow their
lengths can be measured. They are straight bones that extend parallel to each other. The length ratio of the ulna to
the tibia is 94.2%.
Three carpals can be identified. The proximal carpals are fused into a rectangular unit, whose ventral surface is
concave. There are two distal carpals; the lateral one that articulates with the pteroid is smaller than the medial one,
which articulates with the proximal ends of metacarpals II-IV. Most of the ventral surface of the medial distal
length width
Isolated teeth 19.3; 17.6; 10.9 2.4; 2.5; 2.1
Lower jaw 105.6 9.0 (depth)
Caudal vertebrae (1–13) 150 -
Humerus 77.2 7.5
Ulna/radius 65.1 7.6/4.7
Pteroid 15.9 2.2
Metacarpals I–IV 24.2/24.2/24.2/29.3 1.7/1.7/2.1
Wing phalanx 1/Wing phalanx 2 85.7/118.3 6.5/4.8
Femur 68.3 6.1
Tibia 69.1 5.2
Metatarsals I–V 35.2/35.3/35/27.5/? 1.7/1.0/1.0/1.6
Digits 1 and 2 of the fifth toe 25.6/24.6 1.6/0.6
Angle of digit 2 of the fifth toe 140° -
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carpal is concave. The lateral distal carpal is circular in shape. The pteroid is very short and stout (Fig. 4b). The
distal end of the pteroid is blunt, not as pointed as in pterodactyloids such as dsungaripterids (Lü et al. 2009). The
length ratio of the pteroid to the humerus is 0.21. Metacarpals I-III are essentially equal in size and length, and they
are slightly shorter than metacarpal IV. The ratio of the length of metacarpal IV to that of the humerus is 0.38. This
falls slightly out of the range of ratios seen in Rhamphorhynchinae (Unwin 2003; Hone et al. 2012: ratio range:
0.39–0.68) (Table 2).
FIGURE 3. The holotype of Orientognathus chaoyangensis gen. et sp. nov. (41HIII-0418). a, photograph; b,line drawings.
Abbreviations: cr, coracoid; crs, crushed skull; cau, caudals; dor+dorr, dorsals + dorsal ribs; f, femur; h, humerus; ja, jaw; jt,
jaw point; isc, ischium; mcl, manual claw; mtc, metacarpals; mtt, metatarsals; pt, pteroid; sft, second phalange of the fifth toe;
t, tooth; ti, tibia; u/r, ulna and radius; wph1-3, wing phalanxes 1-3.
The right hand is not well preserved, but the left hand is in better condition. The third finger is the longest
among the first three fingers, whereas the first finger is the shortest (Fig. 3). The manual unguals are large and
likely larger than those of the pes (although the pedal claws are not preserved, their small size is inferred from the
slender pedal digits), which is normal for pterosaurs. The unguals bear sharp tips and their proximal ends and are
expanded, with strong flexor tubercles. The articular surfaces with the distal digits occupy half of the width of the
proximal end of each ungual. The distal part of wing phalanx 3 and wing phalanx 4 are missing. Wing phalanx 1 is
shorter than wing phalanx 2. The proximal articular end of the first wing phalanx bears an unfused extensor tendon
process, which indicates that the specimen is not osteologically mature.
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FIGURE 4. The holotype of Orientognathus chaoyangensis gen. et sp. nov. (41HIII-0418); a, Close up of the rostral end of
the jaw; b, Close up of near the wrist (showing the pteroid); c, Whole skeleton; d, Close up of the pes (showing the fifth toe), e,
Close up of the anterior portion of the tail.
The right ilium is only partially preserved. It bears an elongated preacetabular process and there is a clear
concavity on the lateral surface that articulates with the femoral head. The postacetabular process is relatively
short. The pubis is not preserved. The ischium is well preserved; it is fan-like, with a rounded posteroventral
margin, and it bears a distinct neck. The left femur is nearly complete. It has a straight shaft, and a ball-shaped head
set off on a distinct neck. The proximal portion of the tibia is missing, but based on its position relative to the femur
and impression near the distal end of the femur, its length can be measured. It is slightly longer than the femur, with
a tibia: femur length ratio of 1.01. The distal end of the fibula is not fused to the tibia. Two small round bones near
the distal end of the tibia may be the tarsals. The pes is narrow and slender. The metatarsals are long and thin, with
metatarsal II being the longest. Metatarsal III is shorter than metatarsal I but longer than metatarsal IV. The ratio of
the length of the third metatarsal to that of the tibia is 0.51. Metatarsal V is small and perhaps missing a small
portion. The first digit of the fifth toe is straight and nearly similar to metatarsal I-IV in size and shape. The second
digit of the fifth toe is slender, long and curved, and the angle of the second digit of the fifth toe is 140° (Fig. 4d).
Phylogenetic analysis. To evaluate the systematic position of Orientognathus among non-pterodactyloid
pterosaurs, the new taxon was added to the data matrix published byet al. (2012) (Appendix 1). The dataset
consists of three taxa as outgroups (Preondactylus, Dimorphodon and Peteinosaurus) and 21 taxa as ingroups.
Parsimony analysis of the dataset was carried out using PAUP* software (version 4.0b1) (Swofford 2002) under the
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following settings: characters were treated as unordered and of equal weight (except for characters 5 and 55, which
were treated as ordered); multiple states for a single taxon were interpreted as a polymorphism; addition sequence
was set as furthest; default character state optimization was set as DELTRAN. Support for individual nodes was
estimated through a bootstrap analysis (1000 replicates) using a heuristic search and the settings listed above. The
analysis recovered 2931 most parsimonious trees (MPTs), each with a tree length of 192, consistency index (CI) of
0.6510, retention index (RI) of 0.7774, and rescaled consistency index of 0.5061. The strict consensus of these
MPTs (Fig. 5) shows that Orientognathus falls within the clade Rhamphorhynchinae and it is more derived than
Pterorhynchus. Orientognathus shares the following two unambiguous characters with Rhamphorhynchinae:
tapering, spike-like rostral process; and a pteroid that is less than 30% of the length of the humerus. The
autapomorphies of Orientognathus are (1) teeth are peg-like and widely spaced; (2) absence of a lateral pneumatic
foramen on the cervical centra; (3) deltopectoral crest of humerus that is large and subtriangular with a proximally
directed apex; (4) an ulna whose length is less than 133% of that of the humerus; (5) ulna:tibia ratio: 0.9–1.2; and
(6) pubis and ischium are unfused.
FIGURE 5. Strict consensus of the 2931 most parsimonious trees (Tree length = 192, CI = 0.6510, RI = 0.7774, RCI = 0.5061),
showing the relationship of Orientognathus chaoyangensis gen. et sp. nov. (41HIII-0418) to other basal pterosaurs.
, Anurognathidae;
, Scaphognathinae;
, Rhamphorhynchidae;
, Monofenestrata.
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Comparison and discussion
Orientognathus is assigned to the Rhamphorhynchidae based on the following characters: mandible tips fused into
a short symphysis bearing a forward-projecting prow; a number of large, fang-like, procumbent teeth forming a
fish grab (Unwin 2003); and jaws with edentulous tips (shared with Rhamphorhynchus muensteri [Wellnhofer
1975a, 1978; Bennett 1995] and Bellubrunnus rothgaengeri [Hone et al. 2012]); anterior teeth long and angled
forward and laterally; femur shorter than the humerus (shared with Rhamphorhynchus muensteri [Wellnhofer
1975a, 1978; Bennett 1995] and Qinglongopterus guoi [Lü et al. 2012]).
FIGURE 6. Jaw structure comparisons of Rhamphorhynchus muensteri, Scaphognathus (modified from Wellnhofer, 1975a)
and Orientognathus (reversed). Scale bars = 1 cm.
The ulna is shorter than the tibia in Orientognathus, similar to basal pterosaurs such as Preondactylus and
Dimorphodontidae, which exhibit a primitive condition (Unwin et al. 2000). The edentulous tip of the jaw is not as
long as in Rhamphorhynchus muensteri (Bennett 1995; Wellnhofer 1975a) and it is also not as short and deep as in
Scaphognathus (Wellnhofer 1975a) (Fig. 6). The tip morphology of Orientognathus is somewhat intermediate
between the conditions of Rhamphorhynchus and Scaphognathus, more similar to that of Dorygnathus (Padian
2008). The tooth morphology is similar to Rhamphorhynchus.
The distal end of the pteroid is expanded, similar to the condition in most basal pterosaurs, such as
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Qinglongopterus (Lü et al. 2012), Rhamphorhynchus, Dorygnathus (Padian 2008), and Campylognathoides
liasicus (Wellnhofer 1974), but different from the condition in pterodactyloid pterosaurs, where the distal end of
the pteroid is slender and pointed (e.g., Dsungaripterus: Lü et al. 2009; Darwinopterus: Lü et al. 2010a). The
length of the pteroid is 21% of the humerus length, which is similar to the ratio in Qinglongopterus (22.5%: Lü et
al. 2012). In more distantly related taxa this ratio is much greater, as it is about 40% in Campylognathoides liasicus
(Wellnhofer 1974) and 70% in Darwinopterus robustodens (Lü et al. 2011a). Like basal non-pterodactyloids such
as Preondactylus, Dimorphodontidae and Anurognathidae (Unwin et al. 2000), the metatarsals of Orientognathus
exhibit the primitive condition in being highly elongate, slender and subequal to each other in length (Table 1).
TABLE 2. Proportions of the lengths of major elements among various rhamphorhynchines (modified from Hone et al.
2012). Fenghuangopterus from Lü et al. (2010b), Scaphognathus (SMNS 59395) from Bennett (2014); Orientognathus
(41HIII-0418) from this study. Note: in the table 3 of Hone et al. (2012), the last column should refer to the metacarpal
IV/humerus ratio.
Orientognathus shares one character with Rhamphorhynchus, Eudimophodon and Campylognathoides: the
second wing phalanx is longer than the ulna, whereas in other most non-pterodactyloid pterosaurs, the ulna is
longer than wing phalanx two (Unwin et al. 2000).
Orientognathus differs from Scaphognathus in that the ulna is shorter than each wing phalanx in
Orientognathus, whereas the ulna is longer than each wing phalanx in Scaphognathus (Bennett 2014). The second
phalanx of pedal digit V is at an angle of 140° in Orientognathus, but bears an angular flexure of 40-45º at mid-
length in Scaphognathus (Bennett 2014).
Orientognathus differs from Rhamphorhynchus in the shape of the deltopectoral crest of the humerus: there is
no clear neck on the crest in Orientognathus, but a distinct neck on the tongue-shaped deltopectoral crest in
Rhamphorhynchus (Wellnhofer 1991). The length ratio of the metacarpal IV to humerus is approximately 0.38 in
Orientognathus, which is much smaller than in Rhamphorhynchus, which has a ratio of approximately 0.60
(Wellnhofer 1975b) (Table 2).
Orientognathus differs from Qinglongopterus, Pterorhynchus, Darwinopterus, Fenghuangopterus,
Jianchangopterus and Jianchangnathus, which come from the Middle to Late Jurassic deposits of western
Liaoning and surrounding areas of China.
Taxon Humeral length
Humerus/Femur Ulna/Tibia Metacarpal IV /
Humerus ratio
Rhamphorhynchinae - 1.03–1.48 1.25–1.81 0.39–0.68
Rhamphorhynchus (1) TM 6924 16.5 1.32 1.72 0.60
Rhamphorhynchus (1) BSP 1889 XI.1 15.5 1.32 1.72 0.61
Rhamphorhynchus (1) MB 14.5 1.32 1.60 0.54
Rhamphorhynchus (1) MMK V45/1 13.5 1.23 1.62 0.63
Rhamphorhynchus (1) UB E 554 21.8 1.28 1.70 0.63
Rhamphorhynchus (1) 19 1.19 1.86 0.53
Rhamphorhynchus (2) MLU 33 1.18 1.56 0.58
Rhamphorhynchus (2) SMD 34 1.21 1.45 0.53
Dorygnathus BSP 1938 149 51 1.21 1.38 0.49
Dorygnathus UUPM R 157 61 1.22 1.48 0.48
Qinglongopterus D3080 17.8 1.46 1.85 0.93
Nesodactylus AMNH 2000 46.5 - - 0.57
Bellubrunnus BSP–1993–XVIII–2 14 1.40 1.83 0.64
Fenghuangopterus 63 1.11 0.85 0.56
Scaphognathus (SMNS 59395) 29 1.06 1.54 0.46
Orientognathus 77.2 1.13 0.92 0.38
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Orientognathus differs from Qinglongopterus in having a smaller ratio of the length of the metacarpal IV to
the humerus (0.38 vs. 0.51; Lü et al. 2012). The shape of the deltopectoral crest in Orientognathus is also different
from that of Qinglongopterus, where it is more developed compared with the humeral shaft.
Orientognathus differs from Pterorhynchus (Czerkas and Ji 2002) in that the anterior teeth are stouter and
longer in Orientognathus than in Pterorhynchus. The length ratio of metacarpal IV to humerus is also smaller in
Orientognathus (0.38 vs. 0.55).
Orientognathus differs from the scaphognathine pterosaur Fenghuangopterus (Lü et al. 2010b) in that the teeth
are straight in Orientognathus whilst they are curved in Fenghuangopterus. The length ratio of metacarpal IV to
humerus is smaller in Orientognathus, which also has a shorter length ratio of the metacarpals to humerus (0.38 vs.
0.56). The rostral end of the lower jaw is toothless and pointed in Orientognathus, but expanded in
It is difficult to compare Orientognathus with Jianchangnathus (Cheng et al. 2012), because of the lack of
overlapping complete elements. The teeth of Jianchangnathus are relatively smaller and stouter than these of
Orientognathus differs from Jianchangopterus (Lü and Bo 2011c) in that the tibia is much longer than the
femur in Jianchangopterus, whilst the length of tibia is nearly equal to the femur in Orientognathus. The length
ratio of the pteroid to the humerus is smaller in Orientognathus (approximately 0.21) than in Jianchangopterus
(approximately 0.42), and the length ratio of metacarpal IV to the humerus is approximately 0.38 in
Orientognathus but much larger in Jianchangopterus (approximately 0.82).
Orientognathus differs from the transitional form Darwinopterus in that the length ratio of the pteroid to the
humerus is smaller in Orientognathus (0.21 vs. 0.59). Furthermore, the pteroid is short with an expanded distal end
in Orientognathus, but it is slender with a pointed distal end in Darwinopterus (Lü et al. 2009; Wang et al. 2010).
The length ratio of the metacarpal IV to the humerus is approximately 0.66 in Darwinopterus, which is much larger
than in Orientognathus (approximately 0.38).
Orientognathus most likely comes from the Late Jurassic Tuchengzi Formation, which is younger than the
Tiaojishan Formation (Bureau of Geology and Mineral Resources of Liaoning Province 1989). It therefore most
likely represents the youngest Jurassic pterosaur found in China thus far.
Orientognathus is the largest rhamphorhynchinae pterosaur discovered in the Upper Jurassic deposits of
northeastern China thus far. The discovery of Orientognathus may help fill the temporal gap between the better-
sampled Middle Jurassic and Early Cretaceous pterosaur faunas of China, and indicates that more Upper Jurassic
pterosaurs may remain to be found.
The authors are indebted to Dr. Dave Hone and an anonymous reviewer, who reviewed the first draft of the
manuscript and made constructive suggestions. This research was supported by grants from the National Natural
Science Foundation of China (grant no. 41272022) and the China Geological Survey (grant no. 12120114026801)
to Lü JC. The specimen was prepared by Y. Q. Zhang (CAGS).
Andres, B., Clark, J. & Xu, X. (2014) The earliest pterodactyloid and the origin of the Group. Current Biology, 24 (9),
Bennett, S.C. (1995) A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year classes of a
single large species. Journal of Paleontology, 69, 569–580.
Bennett, S.C. (2014) A new specimen of the pterosaur Scaphognathus crassirostris, with comments on constraints of cervical
vertebrae number in pterosaurs. Neues Jahrbuch für Geologie und Paläontologie,Abhandlungen, 271, 327–348.
Zootaxa 3911 (1) © 2015 Magnolia Press
Bureau of Geology and Mineral Resources of Liaoning Province (1989) Regional geology of Liaoning province. Geological
Publishing House, Beijing, 856 pp.
Cheng, X., Wang, X.L., Jiang, S.X. & Kellner, A.W.A. (2012) A new scaphognathid pterosaur from western Liaoning, China.
Historical Biology, 24, 101–111.
Czerkas, S.A. & Ji, Q. (2002) A new rhamphorhynchoid with a headcrest and complex integumentary structures. In: Czerkas,
S.J. (Ed.), Feathered Dinosaurs and the origin of flight. The Dinosaur Museum Journal 1. The Dinosaur Museum of
Blanding, Utah, pp. 15–41
Hone, D.W.E., Tischlinger, H., Frey, E. & Roper, M. (2012) A new non-pterodactyloid pterosaur from the Late Jurassic of
Southern Germany. PLoS ONE, 7, e39312.
Lü, J.C. & Bo, X. (2011) A New rhamphorhynchid pterosaur (Pterosauria) from the Middle Jurassic Tiaojishan Formation of
Western Liaoning, China. Acta Geologica Sinica, 85, 977–983.
Lü, J.C. & Hone, D.W.E. (2012) A new Chinese anurognathid pterosaur and the evolution of pterosaurian tail lengths. Acta
Geologica Sinica, 86, 1317–1325.
Lü, J.C., Unwin, D.M., Zhao, B., Gao, C.L. & Shen, C.Z. (2012) A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae)
from the Middle/ Upper Jurassic of Qinglong, Hebei Province, China. Zootaxa, 3158, 1–19.
Lü, J.C., Xu, L., Chang, H.L. & Zhang, X.L. (2011a) A new darwinopterid pterosaur from the Middle Jurassic of western
Liaoning, Northeastern China and its ecological implications. Acta Geologica Sinica, 85, 507–514.
Lü, J.C., Unwin, D.M., Deeming, D.C., Jin, X.S., Liu, Y.Q. & Ji, Q. (2011b) An egg-adult association, gender, and reproduction
in pterosaurs. Science, 331, 321–324.
Lü, J.C., Unwin, D.M., Jin, X.S., Liu, Y.Q. & Ji, Q. (2010a) Evidence for modular evolution in a long-tailed pterosaur with a
pterodactyloid skull. Proceedings of the Royal Society B, 227, 383–389.
Lü, J.C., Fucha, X.H. & Chen, J.M. (2010b) A new scaphognathine pterosaur from the Middle Jurassic of Western Liaoning,
China. Acta Geoscientica Sinica, 31, 263–266.
Lü, J.C., Azuma, Y., Dong, Z.M., Barsbold, R., Kobayashi, Y. & Lee, Y.N. (2009) New material of dsungaripterid pterosaurs
(Reptilia: Pterosauria) from western Mongolia and its paleoecological implications. Geological Magazine, 146, 690–700.
Padian, K. (2008) The early Jurassic pterosaur Dorygnathus banthensis (Theodori, 1830). Special Papers in Palaeontology, 80, 1–64.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Sinauer Associates,
Sunderland, Massachusetts.
Unwin, D.M. (2003) On the phylogeny and evolutionary history of pterosaurs: In: Buffetaut, E. & Mazin, J.M. (Eds.),
Evolution and Palaeobiology of Pterosaurs, 217, pp. 139–190. [Geological Society, London]
Unwin, D.M., Lü, J.C. & Bakhurina, N.N. (2000) On the systematic and stratigraphic significance of pterosaurs from the Lower
Cretaceous Yixian Formation (Jehol Group) of Liaoning, China. Mitteilungen aus dem Museum für Naturkunde Berlin,
Geowissenschaftliche Reihe, 3, 181–206.
Wang, X.L., Kellner, A.W.A., Jiang, S.X. & Meng, X. (2009) An unusual long-tailed pterosaur with elongated neck from
western Liaoning of China. Anais da Academia Brasileira de Ciências, 81, 793–812.
Wang, X.L., Kellner, A.W.A., Jiang, S.X., Cheng, X., Meng, X. & Rodrigues, T. (2010) New long-tailed pterosaurs
(Wukongopteridae) from western Liaoning, China. Anais da Academia Brasileira de Ciências, 82, 1045–1062.
Wellnhofer, P. (1974) Campylognathoides liassicus (Quenstedt), an Upper Liassic pterosaur from Holzmaden - The Pittsburgh
specimen. Annals Carnegie Museum of Natural History, 45, 5–34.
Wellnhofer, P. (1975a) Die Rhamphorhynchoidea (Pterosauria) der oberjura-plattenkalke süddeutschlands. Teil I. allgemeine
skelletmorphologie. Paläontographica A, 148, 1–33.
Wellnhofer, P. (1975b) Die Rhamphorhynchoidea (Pterosauria) der oberjura-plattenkalke süddeutschlands. Teil II.
systematische beschreibung. Paläontographica A, 148, 132–186.
Wellnhofer, P. (1978) Handbuch der paläoherpetologie. Teil 19. Pterosauria. Verlag Gustav Fischer, Stuttgart, 82 pp.
Wellnhofer, P. (1991) The illustrated encyclopedia of pterosaurs. Salamander Books, London, 192 pp.
APPENDIX 1. Character codings for Orientognathus chaoyangensis gen. et sp. nov. for the phylogenetic analysis of Lü
et al. (2012). The complete matrix includes 82 characters and 24 taxa (with Orientognathus chaoyangensis gen. et sp.
nov. added).
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Pterosaurs were the first vertebrates to evolve powered flight. The timing of their origin is still debated, and hypotheses range from the end of the Permian Period, to the lower Mesozoic Era, and through to the Middle–Late Triassic epochs. Regardless of when they originated, the oldest records are restricted to the Upper Triassic Norian Stage in the northern hemisphere (Europe, USA and Greenland). We report two new raeticodactylid pterosaurs, Yelaphomte praderioi gen. et sp. nov. and Pachagnathus benitoi gen. et sp. nov. from the upper Norian to Rhaetian Quebrada del Barro Formation in north‐western Argentina. The new specimens (an isolated dentary symphysis, partial rostrum, and distal half of ulna) are the first unequivocal Triassic records of pterosaurs in the southern hemisphere, confirming that the absence of pterosaurs outside north‐western Pangaea during the Late Triassic was the result of poor sampling rather than true absence. These new discoveries provide evidence of a greater diversity of pterosaurs living in terrestrial habitats and a wider global distribution of pterosaurs from the beginning of their evolution on Earth.
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Morphology forms the most fundamental level of data in vertebrate palaeontology because it is through interpretations of morphology that taxa are identified, creating the basis for broad evolutionary and palaeobiological hypotheses. Assessing maturity is one of the most basic aspects of morphological interpretation and provides the means to study the evolution of ontogenetic changes, population structure and palaeoecology, life‐history strategies, and heterochrony along evolutionary lineages that would otherwise be lost to time. Saurian reptiles (the least‐inclusive clade containing Lepidosauria and Archosauria) have remained an incredibly diverse, numerous, and disparate clade through their ~260‐million‐year history. Because of the great disparity in this group, assessing maturity of saurian reptiles is difficult, fraught with methodological and terminological ambiguity. We compiled a novel database of literature, assembling >900 individual instances of saurian maturity assessment, to examine critically how saurian maturity has been diagnosed. We review the often inexact and inconsistent terminology used in saurian maturity assessment (e.g. ‘juvenile’, ‘mature’) and provide routes for better clarity and cross‐study coherence. We describe the various methods that have been used to assess maturity in every major saurian group, integrating data from both extant and extinct taxa to give a full account of the current state of the field and providing method‐specific pitfalls, best practices, and fruitful directions for future research. We recommend that a new standard subsection, ‘Ontogenetic Assessment’, be added to the Systematic Palaeontology portions of descriptive studies to provide explicit ontogenetic diagnoses with clear criteria. Because the utility of different ontogenetic criteria is highly subclade dependent among saurians, even for widely used methods (e.g. neurocentral suture fusion), we recommend that phylogenetic context, preferably in the form of a phylogenetic bracket, be used to justify the use of a maturity assessment method. Different methods should be used in conjunction as independent lines of evidence when assessing maturity, instead of an ontogenetic diagnosis resting entirely on a single criterion, which is common in the literature. Critically, there is a need for data from extant taxa with well‐represented growth series to be integrated with the fossil record to ground maturity assessments of extinct taxa in well‐constrained, empirically tested methods.
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A new non-monofenestratan pterosaur with multicusped dentition, Seazzadactylus venieri, is described from the Upper Triassic (middle-upper Norian) of the Carnian Prealps (northeastern Italy). The holotype of S. venieri preserves a complete mandibular and maxillary dentition, along with a nearly complete premaxillary one, showing unique features. Furthermore, the arrangement of the premaxillary teeth and the shape of jugal, pterygoid, ectopterygoid, scapula and pteroid are unique within non-monofenestratan pterosaurs. S. venieri is similar and closely related to Carniadactylus rosenfeldi and Austriadraco dallavecchiai, which are also from the Alpine middle-upper Norian of Italy and Austria, respectively. In a parsimony-based phylogenetic analysis, S. venieri is found to nest within a clade of Triassic pterosaurs composed of Arcticodactylus cromptonellus, Austriadraco dallavecchiai, Carniadactylus rosenfeldi and a trichotomy of Raeticodactylus filisurensis, Caviramus schesaplanensis and MCSNB 8950. This unnamed clade is basal within the Pterosauria, but is not the basalmost clade. Eudimorphodon ranzii lies outside this clade and is more derived, making the Eudimorphodontidae paraphyletic. S. venieri increases the diversity of Triassic pterosaurs and brings the number of pterosaur genera and species in the Dolomia di Forni Formation to four.
Although both the Yanliao and Jehol vertebrate assemblages are known for exceptional preservation of feathered dinosaurs, mammals, pterosaurs, lizards, salamanders, and fish, the Early Cretaceous Jehol Biota also contains birds, choristoderes, frogs, and turtles that are currently lacking in the Jurassic Yanliao Biota. The vertebrate assemblages of the Yanliao and Jehol biotas are very distinct from each other, with the salamander Liaoxitriton being the sole unquestionably shared taxon on generic level. Both assemblages contain mainly stem relatives of major clades of extant vertebrates, with all genera and species extinct. Crown group taxa are restricted to some fishes, salamanders, frogs, and turtles.
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A reassessment of the systematic relationships of pterosaurs from the Lower Cretaceous Yixian Formation of Liaoning Province, China, shows that Dendrorhynchoides should be reassigned to the Anurognathidae (“Rhamphorhynchoidea”) and that Eosipterus possibly belongs within Ctenochasmatidae (Pterodactyloidea). These pterosaurs formed an integral part of a diverse community that inhabited lowland terrestrial environments in the region of northeast China in the Early Cretaceous. A new compilation of data for the Lower Cretaceous hints at a broad differentiation between pterosaurs that lived in continental habitats (anurognathids, ctenochasmatoids, dsungaripteroids) and those that frequented marine environments (ornithocheiroids). Moreover, there is evidence of further differentiation within continental habitats, between pterosaurs living in lowland and coastal regions (anurognathids. ctenochasmatoids) and those living in more inland environments (dsungaripteroids). The temporal and geographical range extensions for high rank taxa that are implied by the Yixian pterosaurs further emphasise the incompleteness and unevenness of the pterosaur fossil record and its unreliability for biostratigraphic zonation. Eine Neubewertung der systematischen Stellung der Flugsaurier von der unterkretazischen Yixian-Formation der Provinz Liaoning, China, zeigt, dass Dendrorhynchoides den Anurognathiden (“Rhamphorhynchoidea”) zugeordnet werden kann und dass Eosipterus vermutlich zu den Ctenochasmatiden (Pterodactyloidea) gehört. Diese beiden Flugsaurier bilden einen integralen Bestandteil einer diversen Fauna, die in der Unteren Kreide ein terrestrisches Flachland-Ökosystem im Nordosten Chinas bewohnte. Fasst man die für die Untere Kreide verfügbaren Daten zusammen, so zeigt sich eine weitgehende Differenzierung zwischen Flugsauriern, die überwiegend in kontinentalen Ökosystemen lebten (Anurognathidae, Ctenochasmatoidea, Dsungaripteroidea) und jenen, die auch oft in marinen Bereichen auftreten (Ornithocheiroidea). Darüber hinaus gibt es auch Hinweise auf eine Differenzierung innerhalb der kontinentalen Habitate, zwischen Pterosauriern, die sich in den Ebenen und küstennahen Bereichen aufhielten (Anurognathidae, Ctenochasmatoidea) und den Bewohnern von mehr küstenfernen Ökosystemen (Dsungaripteroidea). Die von den Taxa der Yixian-Formation angezeigte Erweiterung der stratigraphischen und geographischen Reichweite für Taxa höheren Ranges unterstreichen die Unvollständigkeit und Unausgewogenheit des Fossilberichtes der Flugsaurier und seine Unzulänglichkeit für biostratigraphische Zonierungen. doi:10.1002/mmng.20000030109
A statistical and morphological study of the pterosaur Rhamphorhynchus from the Solnhofen Limestone shows that specimens fall into discrete size-classes that are year-classes resulting from seasonal mortality or preservation of specimens. Taxonomic characters used in the past to separate species of Rhamphorhynchus are all related to size and ontogeny, and all specimens belong to a single species, R. muensteri. Thus, the collections of Rhamphorhynchus provide an unequaled record of the growth and development of a Jurassic rhamphorhynchoid pterosaur. Rhamphorhynchus did not have rapid determinate growth; the growth rate was comparable to that of extant crocodilians, and growth continued over the course of at least three years after individuals began to fly. -Author
Over 30 skeletons and dozens of isolated bones of the Liassic pterosaur Dorygnathus have been recovered from the Early Jurassic (Toarcian) of Baden-Württemberg and Lower Saxony in Germany, and from Nancy, France. All but one specimen have been assigned to the species D. banthensis; the exception was assigned to a larger species, D. 'mistelgauensis', which new discoveries suggest is simply a large individual of D. banthensis. The form of the lower jaw and premaxillary teeth are diagnostic for the genus, as are several other features. Here I review the history of the understanding of Dorygnathus, describe the known specimens in public repositories, and characterize the general morphology and systematic position of the genus. Dorygnathus is distinguished by its extremely large anterior teeth (four premaxillary and three or four anterior dentary teeth), which are proportionally larger than in any other pterosaur, Its deep maxilla gives the skull a high, straight, gradual slope, and its long, deepened, upwardly curved mandibular symphysis is diagnostic for the taxon. Other features such as the proportions of the wing elements, the form of the pelvis, and the shape and proportions of the toes are equally characteristic. Dorygnathus is most closely related to Rhamphorhynchus and the Pterodactyloidea, and represents this lineage in the Early Jurassic of Europe.
A heavily compressed, but nearly complete fossil skeleton recovered from the Middle/Upper Jurassic Tiaojishan Formation of Mutoudeng, Qinglong County, Hebei Province, China, represents a new genus and species of long-tailed pterosaur, Qinglongopterus guoi gen. et sp. nov. The holotype and only known specimen has an estimated forelimb length of 0.18 m. The new taxon is distinguished by a relatively short skull, a remarkably short pteroid with a distinctive knob-like distal expansion, and a prepubis with a relatively slender distal process. Phylogenetic analysis demonstrates that Qinglongopterus is a member of Rhamphorhynchidae, exhibiting many of the unique character states found in members of this clade. Qinglongopterus is strikingly similar to Rhamphorhynchus and more closely related to this taxon than to any other rhamphorhynchine, this pairing is supported by morphometric data and several synapomorphies (short, broad nasal process of the maxilla; forelimb length more than four times that of the hind limb; wing-phalanx one more than twice the length of the tibia). Qinglongopterus demonstrates that the highly derived skeletal morphology of Rhamphorhynchus, known only from the latest Jurassic (Tithonian) of Europe, had already appeared by the start of the Late Jurassic. This hints at evolutionary stasis in Rhamphorhynchinae, a phenomenon seemingly also present in two other clades of basal pterosaurs, Anurognathidae and Scaphognathinae, and contrasting sharply with basal monofenestratans which appear to have undergone extensive evolutionary change during the same interval.
A new complete and fully articulated juvenile specimen of the rhamphorhynchoid pterosaur Scaphognathus crassirostris from the Upper Jurassic Solnhofen Limestone of southern Germany is only the third known specimen of the species. The specimen is described and compared to the other two specimens. Based on the comparisons, the skull of Scaphognathus is reinterpreted as having two premaxillary, six maxillary, and five dentary teeth per jaw side, and a broad boat-shaped snout. Scaphognathus is compared to Jianchangnathus robustus, and revised diagnoses of the genus and family are presented. In addition, the position of the cervico-dorsal transition in the vertebral column of pterosaurs is reviewed, and an apparent constraint to nine cervical vertebrae is noted. © 2014 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.
— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
A new anurognathid pterosaur specimen from the Middle Jurassic Tiaojishan Formation of Qinglong, northern Hebie Province is described. The new specimen is referred to Dendrorhynchoides, based on the general morphology of the skeleton, but it represents a new species, named here as Dendrorhynchoides mutoudengensis sp. nov.. It is characterized by the presence of short, robust and straight teeth, and bearing wing metacarpal approximately 40% of the length of humerus. The new specimen provides further osteological information for anurognathid pterosaurs, especially the presence of a relatively elongate tail.