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Short note on a new anurognathid pterosaur with evidence of perching behaviour from Jianchang of Liaoning Province, China


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A new anurognathid pterosaur, Vesperopterylus lamadongensis gen. et sp. nov., is erected based on a complete skeleton with a skull preserved. It is characterized by two short distinct ridges present on the ventral surface of the cervical vertebrae; coracoids slightly longer than scapula; humerus, wing phalanx 3 and tibia nearly the same in length; grooves clearly present on the posterior surface of the wing phalanges 1-3; and the first toe reversed. It is the first anurognathid pterosaur from China with a definitively short tail, and the first pterosaur with a reversed first toe. The reversed first toe of Vesperopterylus indicates that it had arboreal habitats. The discovery of Vesperopterylus lamadongensis from the Jiufotang Formation strongly expands the geological age range for anurognathid pterosaurs.
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Short note on a new anurognathid pterosaur with evidence
of perching behaviour from Jianchang of
Liaoning Province, China
Institute of Geology, Chinese Academy of Geological Sciences,
Beijing 100037, China
Key Laboratory of Stratigraphy and Palaeontology, Ministry of Land and Resources
of China, Beijing 100037, China
Beijing Museum of Natural History, Beijing 100050, China
Abstract: A new anurognathid pterosaur, Versperopterylus lamadongensis gen. et sp. nov., is
erected based on a complete skeleton with a skull preserved. It is characterized by twoshort distinct
ridges present on the ventral surface of the cervical vertebrae; coracoids slightly longer than scap-
ula; humerus, wing phalanx 3 and tibia nearly the same in length; grooves clearly present on the
posterior surface of the wing phalanges 13; and the first toe reversed. It is the first anurognathid
pterosaur from China with a definitively short tail, and the first pterosaur with a reversed first toe.
The reversed first toe of Versperopterylus indicates that it had arboreal habitats. The discovery of
Versperopterylus lamadongensis from the Jiufotang Formation strongly expands the geological
age range for anurognathid pterosaurs.
Supplementary material: The character list and data matrix for phylogenetic analysis are avail-
able at
The anurognathid pterosaurs were a group of small
pterosaurs known mainly in Europe and Asia dur-
ing the Jurassic period. At present, four genera are
reported: Anurognathus, from the Late Jurassic of
Germany (Do
¨derlein 1923; Bennett 2007); Batra-
chognathus, from the Late Jurassic of Kazakhstan
(Ryabinin 1948; Unwin & Bakhurina 2000; Costa
et al. 2013) and also a specimen from Mongolia
(Bakhurina & Unwin 1995); Dendrorhynchoides,
from the Middle Jurassic of China (Ji & Ji 1998;
Ji et al. 1999; Lu
¨& Hone 2012; Jiang et al.
2015); and Jeholopterus (Ji & Yuan 2002; Wang
et al. 2002; Sullivan et al. 2014) from the Middle
to Late Jurassic of China. Here, we report a new
anurognathid pterosaur: Versperopterylus lama-
dongensis gen. et sp. nov. from the Early Creta-
ceous Jiufotang Formation of Lamamdong, in
Jianchang of Liaoning Province (Fig. 1). It repre-
sents the youngest anurognathid pterosaur in geo-
logical age and this substantially expands the
temporal range of anurognathid pterosaurs. The
reversed first toe of Versperopterylus lamadongen-
sis provides direct evidence of its perching
Systematic palaeontology
Pterosauria Kaup, 1834
Anurognathidae Kuhn 1937
Versperopterylus lamadongensis gen. et sp. nov.
Etymology.Versper-, Latin word for ‘dusk’
implying that the new pterosaur may seek food at
dusk; -pteryl, Latin word for ‘wing’. The specific
name is referred to the fossil locality, lamadong of
Jianchang County, Liaoning Province.
Type specimen. An almost complete skeleton
with skull and jaws (BMNHC-PH-001311). The
specimen is now stored in the Beijing Museum of
Natural History.
Locality and horizon. Lamadong goumen, Jian-
chang County, Liaoning Province; Jiufotang
Diagnosis. A anurognathid pterosaur with the
following combination of characters: the skull is rel-
atively smaller than other anurognathids; the ratio of
length to width is approximately 79.1%; the poste-
rior region of the skull is rounded (shared with Anu-
roganthus and Bactrachognathus); the anterior end
From:Hone, D. W. E., Witton,M.P.&Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology.
Geological Society, London, Special Publications, 455,
#2017 The Author(s). Published by The Geological Society of London. All rights reserved.
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of the lower jaw is horizontally expanded; the cau-
dal end of the mandible is without a distinct dorsal
‘coronoid’ eminence; postexapophyses are present
on the cervical vertebrae; the number of caudal ver-
tebrae is 15 or fewer (shared with Anuroganthus);
metacarpals I III are the same length (shared with
Batrachognahus and Dendrorhynchoids); a promi-
nent anteriorly directed tubercle on the dorsal apex
of the external trochanter of the femur; the length
of metatarsal IV is shorter than metatarsals IIIII
(shared with Jeholopterus); coracoids are slightly
longer than the scapula; the humerus, wing phalange
3 and tibia are nearly the same length; the wing
metacarpal is approximately 36% of the length of
the humerus; grooves are clearly present on the pos-
terior surface of the wing phalange 1 to the wing
phalange 3; and the first toe is reversed.
The specimen is almost complete, except for miss-
ing one humerus and some cervical vertebrae (Figs
24; Table 1). The wing span is about 1 m. The
fused scapulocoracoid, the smooth long bone sur-
faces and the fusion of extensor process to the
wing phalanx 1 indicate that it is at least a sub-adult
animal. The skull is remarkably small on this ani-
mal. The ratio of skull width to humeral length is
0.57, which is smaller than that Dendrorhynchoides
mutoudengensis (this ratio is 0.74: Lu
¨& Hone
2012). It is unlike other anurognathids and its ven-
tral surface is exposed. The skull is slightly wider
than long. The ratio of length to width of the skull
is approximately 79.1%, which is similar to that of
Dendrorhynchoides mutoudengensis (this ratio is
80%: Lu
¨& Hone 2012). It is semi-circular in ventral
view (Fig. 3a). The premaxilla and maxilla are slen-
der, rod-like. The upper jaw teeth are relatively stout
with blunt tips (Fig. 3b), unlike some longer teeth
with curved tips found in Dendrorhynchoides
mutoudengensis (Lu
¨& Hone 2012). There are
eight teeth preserved. Although the exact number
of teeth is not sure due to the poor preservation,
there are, perhaps, 12 teeth in the upper jaw accord-
ing to the distance between two teeth and the
dentition length.
There are four cervical vertebrae preserved. The
anterior two are rectangular in ventral view. There
are two short, distinct longitudinal ridges on the cen-
tral portion of the centrum, which form a groove
between them. The cervical vertebrae bear a distinct
postexapophysis. The neural spine is high and
blade-like. There are no cervical ribs associated
with the cervical vertebrae.
The anterior dorsal vertebrae bear a distinct neu-
ral spine. There are 14 dorsal vertebrae preserved
with their ventral surfaces exposed. Both the ante-
rior and posterior articular ends are flat. The ventral
surfaces are smooth. The shaft of the third dorsal rib
is the widest of the dorsal ribs. The seventh dorsal
rib is the longest and the first one is the shortest.
At least 13 caudal vertebrae can be observed.
The anterior part of the tail is covered by the
ischium, but it is inferred that no more than two cau-
dal vertebrae were covered. This means that there
are no more than 15 caudal vertebrae (Fig. 4d).
Fig. 1. A map of Chinese anurognathid pterosaur sites: Dendrorhynchoides;Jeholopterus;
Dendrorhynchoides mutoudengensis;ÐVersperopterylus (BMNHC-PH-001311).
The anterior caudal vertebrae are rectangular in lat-
eral view. The last one bears a pointed distal end.
The left scapula and coracoid are fused into
the scapulocoracoid. The shaft of the scapula is
slightly concave and it is slightly shorter than the
coracoid. The coracoid is stouter than the scapula
and it bears a distinct brachial flange. The shaft of
the coracoid is straight. The shaft of the humerus
Fig. 2. Photograph (a) and line drawings (b)ofVersperopterylus (BMNHC-PH-001311). Abbreviations: cav, caudal
vertebrae; csk, crushed skull; co, coracoids; dv, dorsal vertebrae; dr, dorsal ribs; f, femur; fot, fourth toe; fi, fibula;
ft, first toe; h,, humerus; lj, lower jaw; mc, maniual claw; mcI-IV, metacarpals I –IV; mttI-IV, metatarsals I IV;
mttv, metatarsal V; pt, pteroid; rd, radius; sc, scapula; ti, tibia; ul, ulna.
Fig. 3. (a). Photograph of the skull of Versperopterylus; and (b) Close-up of the dentition, showing the teeth
(arrows point). Scale bar is 0.5 cm in (b).
Fig. 4. Photographs of the holotype Versperopterylus (BMNHC-PH-001311): (a) whole specimen; (b) close-up of
the pes; (c) close-up of wing phalanges 1 and 2; and (d) close-up of the tail. Abbreviations: cav, caudal vertebrae;
fot, the fourth toe; gr, groove; mtt I– IV, metatarsal I–IV; rft, the reversed first toe; wph1, 2, wing phalanges 1 and
2; st, second toe; tt, third toe. Scale bars are 1 cm in (b) and (d).
is straight. There is no pneumatic opening near its
proximal end. The deltopectoral crest of the
humerus is relatively small, and is longer than
wide. It is located proximally. There is a distinct
ridge near the distal end of the humerus. The lateral
margin of the deltopectoral crest is straight.
The ulna and radius are parallel to each other.
The ulna is longer than the radius. The width ratio
of ulna to radius is 1.17. The ulna bears a distinct
olecranon process at its proximal end. The shaft of
the radius is straight with slightly expanded distal
and proximal ends.
The proximal carpal is irregular in shape with a
concave surface in ventral view. It is larger than
any other carpals. It articulates with the distal end
of the ulna. There are two unfused distal carpals.
The proximal portion of the pteroid is slightly
curved. The distal end of the pteroid is expanded,
and knob-like. The length ratio of pteroid to
humerus is 0.19. Metacarpals I IV are the same
length. Metacarpal IV is stout. The first phalanx of
the first manual digit is long and slender, slightly
thinner than metacarpals I III. The ungual is deep
at its proximal end and curved with a sharp tip.
The rest of the digits of the left hand are missing.
The wing phalanges decrease in length from the
first to the third. The ventral surfaces of all the right
wing phalanges are exposed. Although the phalan-
ges were heavily depressed during preservation, a
distinct groove appears posterior to the middle
shaft of wing phalanx 1 (Fig. 4c). This groove is
on the posterior surface of the wing phalanx 1.
The extensor tendon process is fused with the first
wing phalanx. The proximal end of the first wing
phalanx is expanded. The first wing phalange is
the longest. The proximal end of the second wing
phalanx is expanded, larger than its distal end.
The distal end of the wing phalanx 2 is slightly
expanded. The proximal end of the third wing pha-
lange is much more expanded than its distal end,
which becomes thinner but is slightly expanded.
There is also a distinct groove on the posterior sur-
faces of wing phalanges 2 and 3. The fourth wing
phalanx is rod-like, quite small and straight with
pointed distal end. This is different to the curvature
seen in the distal phalanges of other anurognathids
(see Hone et al. 2015), but this is also very small
and coupled with the apparent tapering of phalanx
3; thus, it might explain the apparent absence of
the fourth phanalx in the juvenile Anurogahtus.
Table 1. Measurements (in mm) of holotype of Versperopterylus lamadongensis gen. et sp. nov
Length Width
Skull 32.1 40.6 (between quadrate)
Single cervical 9.4 6.8
Dorsals +sacrals 112.9
Tail 31.0
Scapula 37.2
Coracoid 38.3 3.4
Humerus 71.0 6.4
Deltopectoral crest 12.5 5.2
Ulna 96.4 5.6
Radius 92.9 4.8
Metacarpals 25.8
Pteroid 13.7 1.7
Manus digit I: 1; 2 15.8; 13.6 (claw) 1.1; 5.4 (proximal end)
Width of phalange 1 121.3 (with extensor process);
117.8 (without extensor process)
Width of phalange 2 95.4 2.9
Width of phalange 3 71.0 1.9
Width of phalange 4 14.3 1.0
Acetabulum 7.8 4.7 (height)
Pre-acetabulum process 32.0
Femur 52.6 5.7
Tibia 71.9 6.6
Fibula 47.9
Metatarsals I– IV (whole) 34.5 8.0
Metatarsals I– V 30.9; 34.0; 34.0; 33.0; 13.9 3.5; ; ; ;
Pedal digit I: 1; 2 12.2; 8.1 2.1; 4.3
Pedal digit II: 1; 2; 3 6.3; 11.7; 11.2 2.5; 1.6; 4.4
Pedal digit III: 1; 2: 3; 4 5.4; 4.9; 9.8; 10.6 2.7; 2.9; 1.9; 5.1
Pedal digit IV: 1; 2; 3; 4; 5 4.6; 3.9; 2.1; 7.5; 9.2 2.6; 2.6; 2.5; 1.9; 4.1
Pedal digit V: 1; 2 19.8; 1.8
The pelvic girdle is not well preserved. It seems
that the pubis, ischium and ilium are fused together.
The preacetabular process of the ilium is slender and
long, covering at least four posterior dorsal verte-
brae. The acetabulum bears a distinct dorsal margin
and anterior margin. The shapes of ischium and
pubis are unclear because of being covered by the
proximal portion of the femur. The femoral head
has a distinct neck, which forms an angle of 1208
with the shaft of the femur. There is a distinct
ridge on the anterolateral surface near the proximal
end. The femoral head bears a sharp margin. The
shaft of the femur is straight. The tibia is straight
with an oblique proximal end. The proximal end
of the fibula is fused with the proximal end of the
tibia. The proximal portion of the fibula is rod-like
but below it, by about 0.7 cm, it becomes sharply
expanded, which may be a pathology. The distal
end of the tibia bears two distinct condyles. Metatar-
sals I IV are in close contact with each other. Their
proximal ends are fused. Metatarsal II and metatar-
sal III are the same length and size. Metatarsal I is
shorter than metatarsal IV, which is, in turn, shorter
than metatarsals II and III. Metatarsal V is sepa-
rated from other metatarsals. It is stout and shorter
than other metatarsals. The first phalange of digit 5
is straight but its length cannot be determined due
to a missing distal portion. The first toe of the
right pes is clearly reversed (Fig. 4b), although
the left toe is slightly destroyed. The tips of the
pedal unguals of digits II, III and IV are pointed
in the same direction, whilst the tip of the first toe
is pointed in the opposite direction in both pedes;
thus, it is inferred that the first toe is reversed.
The naturally articulated dorsal to caudal series
and all other elements that are not missing indicate
that they are not transported far and are quickly
buried after death. Therefore, they are regarded as
genuine, not caused by preservation. The digital
formula is 2–3–4–5–2. The pedal unguals are
curved with sharp tips. They bear a large extension
process. The pedal unguals are slightly smaller than
the manual unguals.
Comparison and discussion
Versperopterylus differs from Dendrorhynchoides
(Ji & Ji 1998; Lu
¨& Hone 2012; Jiang et al. 2015)
in that the length ratio of wing phalange 2 to
Fig. 5. The living scene of Versperopterylus (drawn by Zhao Chuang).
Fig. 6. Strict consensus of 630 most parsimonious trees (MPTs) obtained by TNT, based on analysis of 68 taxa and
124 characters, showing the phylogenetic position of Versperopterylus. gen. et sp. nov. (tree length, 503). The
numbers adjacent to each node are Bremer support values.
phalange 1 (i.e. 0.79) is smaller than that of Dendro-
rhynchoides (i.e. 0.88). The length ratio of wing
metacarpal to humerus is also smaller than that of
Dendrorhynchoides (it is 36% in Versperopterylus:
however, it is 40.6% in Dendrorhynchoides mutou-
¨& Hone 2012). Versperopterylus has
a tail that is shorter than Dendrorhynchoides.Den-
drorhynchoides was first reported to bear a long
tail because it was not clearly preserved, and was
once thought to have had the tail added onto the
specimen (Unwin et al. 2000). However, the discov-
ery of more material further confirms that there is a
greater likelihood that it had a long tail, as more spec-
imens indicate (Lu
¨& Hone 2012; Jiang et al. 2015).
Versperopterylus differs from Jeholopterus
(Wang et al. 2002; Sullivan et al. 2014) in that Jeho-
lopterus does not bear a tail (according to the holo-
type (IVPP V12705) and the referred specimen
(CAGS-IG-02-81), which came from the same
place, both do not have a visible tails; therefore,
we inferred that Jeholopterus does not bear a tail,
although we cannot exclude the possibility that the
tail is not preserved even if they do have a tail,
the tail should be very short); the length ratio of
wing phalange 2 to phalange 1, and the ratio of
ulna to humerus and femur to tibia, are smaller
than those of Jeholopterus (IVPP V12705) (they
are 0.79, 1.36 and 0.73 in Versperopterylus, whilst
they are 0.88, 1.44 and 0.80 in Jeholopterus,
Versperopterylus differs from Anurognathus
¨derlein 1923; Dalla Vecchia 2002; Bennett
2007) in that the length ratio of the wing metacarpal
to the metatarsals is greater than that of Anurogna-
thus (this ratio is 0.61 in Anurognathus (BSP
1922.I.42), whilst it is 0.76 in Vesperopterylus).
Versperopterylus differs from Batrachognathus
(Ryabinin 1948; Unwin & Bakhurina 2000) in that
its length ratio of ulna to humerus is greater than
that of Batrachognathus (the ratio is 1.0 in Batra-
chognathus, whilst it is 1.36 in Vesperopterylus).
At present, there are three kinds of anurognathid
pterosaurs from western Liaoning and its surround-
ing areas, in terms of their body bauplans: a tail-less
one, such as Jeholopterus; one with a short tail, such
as Versperopterylus; and one with a longer tail, such
as Dendrorhynchoides. The reversed first toe of Ver-
speropterylus indicates that it had arboreal habitats,
which seems like more of a gripping adaptation: ver-
tical clinging, branch climbing and/or branch cling-
ing (above or below the branch) (Fig. 5).
Phylogenetic analysis
A phylogenetic analysis was carried out which in-
cluded 88 ingroup taxa and 124 characters (based
on the modified data matrix of Lu
¨et al. 2016). We
subjected the dataset to a maximum parsimony anal-
ysis in TNT v1.1 (Goloboff et al. 2008). We first
conducted a ‘new technology’ search (with default
parameters for sectorial search, ratchet, tree drift
and tree fusion) that recovered a minimum length
tree in 10 replicates. This procedure aims to broadly
sample tree space and identify individual tree
islands. We then subjected the recovered most par-
simonious trees (MPTs) to a traditional search with
TBR branch swapping, which more fully explores
the tree islands found in the ‘new technology’
search. This process returned a total of 3 043 358
MPTs in 503 steps. Bremer values were used to
assess clade support. The strict consensus of the
630 MPTs (Fig. 6) recovers a monophyletic group
of anurognathid pterosaurs, with Versperopterylus
basal to other anurognathid pterosaurs. The clade
Anurognathidae is supported by the following syn-
apomorphies: skull broad with very short preorbital
region; palatal elements reduced to thin bars of
bone; teeth are small, peg-like and widely spaced;
cervical ribs are highly reduced or absent; the com-
bined length of the caudal vertebrae is shorter than
the dorsal series; the ulna is 133150% of the
humerus length; the manus digit IV (wing-finger)
phalanx 1 is 1.5 2.0 times longer compared to the
length of the tibiotarsus; the contribution of the
wing-phalanx 1 to the wing-finger length is 30
40%; and preacetabular process of the ilium is lon-
ger than the postacetabular process.
Versperopterylus is the first anurognathid pterosaur
from China preserved with a clearly reduced tail,
and it is also the youngest anurognathid pterosaur
in geological age. The reversed first toe of Versper-
opterylus may indicate that it, perhaps, has a grip-
ping adaptation.
We thank Dr Dale Winkler (Southern Methodist Univer-
sity, Dallas, TX, USA) for providing thorough comments
on the earlier version of this paper. This study was sup-
ported by the National Natural Science Foundation of
China (grant Nos 41688103 and 41672019), the Funda-
mental Research Funds for the Chinese Academy of
Geological Sciences (grant No. JB1504) and the project
from the China Geological Survey (DD 20160126) to
Junchang Lu
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... They shared a distinctive suite of morphological characters consistent with being specialized aerial insectivores active in low light conditions. These include (1) a wide mouth with a large gape and a sharp isodont dentition adapted for catching and holding insects (Bennett, 2007a;Ősi, 2011;Hone, 2020), (2) small size, a deep wing with curved wingtip and a short pliable tail, which together facilitated slow and highly maneuverable flight in dense forests (Bennett, 2007a;Ősi, 2011;Hone et al., 2015;Hone, 2020) and (3) large, anterolaterally directed eyes that were likely adapted for nocturnal and/or crepuscular predation (Bennett, 2007a;Lü et al., 2018). The gross morphology of anurognathids changed little during their 40-Ma existence, which suggested they had a rather conservative bauplan Bennett, 2007a;Hone, 2020). ...
... Nine specimens (including specimens of Anurognathus, Dendrorhynchoides, Luopterus, Sinomacrops, and Batrachognathus, and specimens NJU-57003 and CAGS-Z070 [Dalla Vechia, 2002;Yang et al., 2019;Hone, 2020;Wei et al., 2021]) show characteristics of immaturity, e.g., unfused articular bones, scapula-coracoid, pelvis and cranial elements. In contrast, the holotypes of Jeholopterus and Vesperopterylus show osteological maturity and are thus (sub)adults (Wang et al., 2002;Lü et al., 2018). Two specimens lack detailed descriptions, and their ontogenetic state is unknown (Gao et al., 2009;Jiang et al., 2015). ...
... Inadequate recognition of ontogenetic status in previous studies of anurognathids has raised concerns about confounding ontogenetic variation with systematics (Hone, 2020). In particular, many length ratios used as diagnostic (e.g., Wang et al., 2002;Lü and Hone, 2012;Wei et al., 2021) or in analyzing characters (e.g., Kellner, 2003;Lü et al., 2018) may change greatly during growth (Delfino and Sánchez-Villagra, 2010). Previous statistical studies on ontogenetic changes in other pterosaurs have suggested or prompted taxonomic revisions for some taxa (e.g., Bennett, 1995Bennett, , 1996Bennett, , 2006Bennett, , 2007bBennett, , 2013Jouve, 2004;Vidovic and Martill, 2014). ...
... The two specimens referred to Batrachognathus volans come from the Upper Jurassic (Oxfordian-Kimmeridgian) Karabastau Formation (Karabastau Svita; Unwin & Bakhurina 2000: 423) of the Karatau Ridge, Kazakhstan. Specimen PIN 52-2 includes the better preserved anurognathid skull reported to date, although it is slightly disar- Wang, Liu, Shen & Zhang, 2018;and Sinomacrops bondei Wei, Pêgas, Shen, Guo, Ma, Sun & Zhou, 2021 were added to the list. Unnamed and only preliminarily described anurognathids are reported from the Sinuiju Beds (Lower Cretaceous) of North Korea (Gao et al. 2009) and the Middle Jurassic of China (Jiang et al. 2015). ...
... Under Unwin's definition, 'Dimorphodon' weintraubi would not be considered an anurognathid. Similarly, sensu Unwin (2003), Vesperopterylus lamadongensis would not be considered an anurognathid in the strict consensus tree by Lü et al. (2018), and Dendrorhynchoides curvidentatus would not be considered an anurognathid in the strict consensus tree obtained from the matrix of Dalla Vecchia (2019) modified according to the results of this paper (see below). To avoid problems, the definition of Hone (2020) was used in this paper. ...
... Unlike other pterosaurs, the skull is comparatively small with respect to body size, especially in largest and probably more mature individuals. This is evident in the holotypes of Jeholopterus ningchengensis (see Wang et al. 2002) and Vesperopterylus lamadongensis (see Lü et al. 2018); the small size of the skull is independent from the extreme reduction of the rostrum. ...
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The anurognathids are peculiar pterosaurs characterized by broad skulls with very short rostra and broadly arched jaws. The presence of distinct or confluent external naris and antorbital fenestra in these pterosaurs has been debated in the last years. The relatively well-preserved specimens of Batrachognathus volans show that the antorbital fenestra was confluent with the orbit forming an enormous orbitoantorbital fenestra. This feature is evident also in Jeholopterus ningchengensis. The consequent modification of the matrices of two recently published phylogenetic analyses about the in-group pterosaur relationships shows that the Anurognathidae are a derived clade of non-monofenestratan pterosaurs. Anurognathidae (including also 'Dimorphodon' weintraubi according to the definition by Hone 2020) are still a scarcely known clade because only a few specimens have been adequately described in the literature.
... Recently, a new genus has been erected to accommodate this species: Luopterus, named after the late Prof. Junchang Lü (Hone, 2020). Moreover, a second Cretaceous anurognathid was also named recently, Vesperopterylus lamadongensis, known from an almost complete holotype from the late Aptian Jiufotang Formation (Lü et al., 2018). ...
... Despite being crushed to the point of obliterating many details, the specimen is rather complete and provides new information for the group, including the first record of an anurognathid skull exposed in mostly lateral view. In other specimens, the skull is either exposed in mostly internal view, as in the holotype of Anurognathus ammoni (Döderlein, 1923;Wellnhofer, 1975;Bennett, 2007), or dorsoventrally crushed, as in all other specimens that preserve a skull (Riabinin, 1948;Ji & Ji, 1998;Bennett, 2007;Gao et al., 2009;Lü & Hone, 2012;Lü et al., 2018). ...
... We infer that this hemimandible is complete because its length equals that of the upper jaw. It is only slightly bowed, as in Batrachognathus volans, instead of strongly semicircular as in the jaws of Dendrorhynchoides (Ji & Ji, 1998), Luopterus (Lü & Hone, 2012;Hone, 2020), Jeholopterus (Wang et al., 2002), Anurognathus (Bennett, 2007) or Vesperopterylus (Lü et al., 2018). ...
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Anurognathids are an elusive group of diminutive, potentially arboreal pterosaurs. Even though their monophyly has been well-supported, their intrarelationships have been obscure, and their phylogenetic placement even more. In the present work, we present a new genus and species from the Middle-Late Jurassic Tiaojishan Formation, the third nominal anurognathid species from the Jurassic of China. The new species provides new information concerning morphological diversity for the group. Furthermore, we provide a new phylogenetic analysis incorporating into a single data set characters from diverging phylogenetic proposals. Our results place them as the sister-group of Darwinoptera + Pterodactyloidea, as basal members of the Monofenestrata.
... They were likely the only clade of nonpterodactyloid pterosaurs to have crossed the Jurassic-Cretaceous boundary (Hone and Benton, 2007). Anurognathids are rare, with less than a dozen specimens reported, and most of these having been discovered only in the last fifteen years (Bennett, 2007a;Gao et al., 2009;Lü and Hone, 2012;Jiang et al., 2015;Lü et al., 2018;Yang et al., 2019). ...
... Most recently, Vesperopterylus (Lü et al., 2018) has been named and briefly described. This unusual anurognathid had a relatively small skull and apparently also a reversed first toe giving it some grasping ability. ...
... In both, the anurognathids are recovered in an early branching position either as the earliest-diverging clade within Pterosauria (Kellner, 2003) or branching off after only Preondactylus and the Dimorphodontidae (Unwin, 2003a). Most other analyses (both before and after those of 2003) consistently reflected these positions with anurognathids either as the first branching clade in the pterosaurian tree (Kellner, 1996;Lü and Ji, 2006;Bennett, 2007a;Wang et al., 2008;Lü et al., 2018) or in an early branching position (Unwin, 1995(Unwin, , 2003bViscardi et al., 1999;Lü et al., 2010) with only one or two taxa in earlier branching positions. Recently, Vidovic and Martill (2018) recovered the anurognathids as a clade within a Scaphognathinae (and therefore suggested they should be renamed the Anurognathinae) based on the curvature of the skull and teeth, and shape of the pubis, and this hypothesis should not be overlooked. ...
The anurognathids are an enigmatic and distinctive clade of small, non‐pterodactyloid pterosaurs with an unusual combination of anatomical traits in the head, neck, wings and tail. They are known from very limited remains and few have been described in detail, and as a result, much of their biology remains uncertain. This is despite their importance as potentially one of the earliest branches of pterosaur evolution or even lying close to the origins of pterodactyloids. This review covers the taxonomy and palaeoecology of the anurognathids, which remain an interesting branch of pterosaurian evolution.
... Within Pterosauria, the clade Anurognathidae, whose phylogenetic position is still contended as either early or late branching (Kellner, 2003;Wei et al., 2021), demonstrates atypical features when compared to other pterosaurs. Anurognathidae currently consists of eight monotypic genera: Anurognathus ammoni (Döderlein, 1924), Batrachognathus volans (Ryabinin, 1948), Dendrorhynchoides curvidentatus (Ji & Ji, 1998), Jeholopterus ningchengensis (Wang, 2002), Luopterus mutodengensis (Lü & Hone, 2012), Vesperopterylus lamadongensis (Lü et al., 2018), Sinomacrops bondei (Wei et al., 2021), and Cascocauda rong (Yang et al., 2022). All anurognathids exhibit mediolaterally wide skulls (≥120% skull width: length) making the depth and width of the skull subequal lengths, ultimately facilitating a mediolaterally wide gape and accompanied by peg-like teeth (Hone, 2020). ...
... Anurognathid feet are plantigrade with predominantly anteriorly directed toes. This would give limited hindlimb perching ability although Lü et al. (2018) suggested that digit I is reversed in Vesperopterylus, which if correct, would have given it a much greater capacity for grasping. More importantly, as with all non-pterodactyloid pterosaurs, anurognathids also possessed three clawed free fingers (that were not integrated with the wing membrane) that would have been employed in any form of terrestrial or arboreal locomotion (Bennett, 2007) and are better suited to gripping than the pes. ...
Across the evolution of powered flight, the ecological niche of aerial insectivore has been occupied by members of the three volant vertebrate clades—Aves and Chiroptera, and the first known volant vertebrates, pterosaurs. However, morphological and quantitative evidence to support pterosaurs exhibiting this ecology remains scant. Anurognathids are an unusual group of pterosaurs in which the skull exhibits the unique morphology of being mediolaterally expanded, so much so that their skulls may be wider than rostrocaudally long. Here, we conduct quantitative comparative cranial measurements and dental morphology in anurognathids against extant avian and chiropteran taxa, respectively, with ecologies and behaviors that are similar to predicted putative behaviors of anurognathids. Comparative analyses of both skull and dental morphology suggest anurognathid specimens in similar morphospaces as insectivorous crepuscular and nocturnal extant volant taxa. Our results support that this unique group of pterosaurs likely occupied a niche of mid‐flight insectivorous capture in low‐light conditions. Utilizing neontological approaches and comparative anatomy our results support that anurognathid pterosaurs likely occupied a niche of mid‐flight insectivory in in low‐light conditions.
... The earliest bone fossil remains of a pterodactyloid are recorded for the Callovian-Oxfordian boundary (Middle-Upper Jurassic boundary, [122]), while tracks are referred to the Aalenian (lower Middle Jurassic, [111]). On the other hand, the youngest osteological record of non-pterodactyloid pterosaur corresponds to the anurognathids from the lower Aptian Jiufotang Formation of China (upper Lower Cretaceous, [123]). Consequently, any pterosaur tracks younger than the latter age, including the entire Upper Cretaceous, and the Campanian age of the Anacleto Formation of the present study, cannot be supported by the non-pterodactyloid pterosaur record to date. ...
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The Campanian Anacleto Formation holds an abundant and diverse ichnofossil and body-fossil vertebrate record. Despite the striking diversity of this record, pterosaur fossils had never been described from the unit. Here, we report four pterosaur manus tracks from fluvial red beds cropping out in the Área Natural Protegida Municipal Paso Córdoba (Río Negro Province, northern Patagonia, Argentina). Tracks are longer than wide, tridactyl with digit impressions of different lengths (I < II < III), anteriorly directed and laterally asymmetrical. Being on loose slabs and lacking direct examination of pes morphology, the material is classified as undetermined pterosaur tracks. The new find represents the first occurrence of pterosaurs from the lower–middle Campanian of Argentina and one of the few evidences from South America for this time interval. In addition, it is one of the few ichnological pterosaur records from Gondwana, thus shedding light on the palaeobiogeography of this clade during the latest Cretaceous. Pterosaur tracks from the Anacleto Formation allow us to integrate the body-fossil record from the unit and to add a new component, along with birds, to the flying archosaur fauna coexisting with non-avian dinosaurs, notosuchians, chelonians, squamates and mammals in the Campanian of northern Patagonia.
... 5: Vorläufige Größenschätzung aufgrund der Halswirbelreste JME-ETT-04391 bei möglicher Bestimmung als Rhamphorhynchus (siehe Text): Der Neufund würde ein Individuum mit reichlich 180 % der Größe des bisher belegten Maximums dieser Gattung implizieren. Maßstab gesamt 2 m. wenig verlängert sind (Lu et al. 2017), kommen diese selbst bei den größten weltweit bekannten Exemplaren kaum auf 1 cm Länge (Wang et al. 2002;Kellner et al. 2010) und scheiden sie als Bestimmung für JME-ETT-04391 aus. ...
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Die Spur einer Spur-ein möglicher erster Flugsaurier aus Ettling Trace of a trace-a putative first pterosaur from the Ettling locality Archaeopteryx, 37: 75-83; Eichstätt 2021 Zusammenfassung Aus den Plattenkalken von Ettling sind im Vergleich zu anderen Fundstellen des oberjurassischen Solnhofen-Archipels bislang nur wenige Reptilreste und keine von Flugsauriern geborgen worden. Ein neu identifiziertes Fossil von mäßiger Erhaltung wird mit Vorbehalt als zwei assoziierte, aber dislozierte Halswirbel interpre-tiert. Ihre Form und ursprünglich filigrane Knochen-struktur legen einen Flugsaurier nahe. Sollte dies zu-treffen, liegt wahrscheinlich ein überdurchschnittlich großer Rhamphorhynchide vor, zudem einer der größ-ten Juraflugsaurier weltweit. Abstract Compared to other localities of the Upper Jurassic Sol-nhofen Archipelago, the Plattenkalk from Ettling has so far produced only a few reptilian remains, none of them from pterosaurs. A newly identified, badly preserved fossil is tentatively interpreted as two associated but dislocated cervical vertebrae. Their shape and originally delicate bone structure suggest a pterosaur. If true, an outstandingly large rhamphorhynchid would be present, moreover one of the biggest Jurassic ptero-saurs on a global scale. Abb. 1: Neu identifiziertes Fossil aus der Grabung Ettling, JME-ETT-04391, zwei wahrscheinlich in Beziehung stehende, stark ge-löste und daher mutmaßliche Halswirbel eines Pterosauriers. Fig. 1: Newly identified fossil from the Ettling excavation, JME-ETT-04391, two likely associated, but heavily dissolved and therefore only supposed cervical vertebrae of a pterosaur.
... Circles represent genera (Andres and Ji, 2008;Kellner, 1985, 1997;Dong and Lü, 2005;Frey and Martill, 1994;Howse et al., 2001;Ji and Ji, 1997;Jiang and Wang, 2011;Kellner, 1984;Kellner and Tomida, 2000;Lawson, 1975;Lü, 2003;Lü et al., 2012aLü et al., , 2012bLü et al., , 2017Lü and Ji, 2005;Lü and Qiang, 2005;Lü and Zhang, 2005;Marsh, 1876;Rodrigues et al., 2015;Steel et al., 2005;Sweetman and Martill, 2010;Unwin, 2001;Veldmeijer et al., 2005;Wang et al., 2005Wang et al., , 2007Wang et al., , 2012Wang et al., , 2014Wang and Lü, 2001;Wang and Zhou, 2006;Wellnhofer, 1987). ...
A new genus and species of edentulous pterodactyloid pterosaur with a distinctive partial rostrum from the mid-Cretaceous (?Albian/Cenomanian) Kem Kem beds of southeast Morocco is described. The taxon is assigned to Chaoyangopteridae based upon its edentulous jaws, elongate rostrum and slightly concave dorsal outline. The rostral cross-section is rounded dorsally and concave on the occlusal surface. The lateral margins are gently convex dorsally becoming slightly wider toward the occlusal border, and a row of small lateral foramina parallel to the dorsal margin determines it as a taxon distinct from other chaoyangopterids. Apatorhamphus gyrostega gen et sp. nov. is a pterosaur of medium to large size (wingspan likely somewhere between ~3 m and ~7 m). This new species brings the number of named Kem Kem azhdarchoids to three, and the number of named Kem Kem pterosaurs to five, indicating a high pterosaur diversity for the Kem Kem beds.
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The pterosaur is the earliest and largest powered flying vertebrate, even earlier and larger than the other extant archosaurian group, birds. However, evidence for this flying reptile, including the diversity of the small-sized pterosaur after the mid-Cretaceous, and their ecology, has remained elusive. Here we present numerous and dense pterosaur track assemblages from the Hwasun Seoyuri tracksite in the Upper Cretaceous Jangdong Formation of the Neungju Basin in Korea. The pterosaur track assemblage, assigned to Pteraichnus isp., consists of various sized, randomly oriented manus-dominated tracks with several pes claw marks. These features commonly indicate the semi-aquatic behavior and multi-age gregariousness of pterosaurs. The supposed trackmaker of pterosaur tracks would be the small-sized pterodactyloid that inhibited the Late Cretaceous Korean Peninsula, but that has not previously been reported. This ichnological evidence for the global distribution of small-sized pterosaurs could be interpreted to mean that the pterosaur fauna in the Late Cretaceous was more distributed and diverse than was previously known.
Pterosaurs, which lived during the Mesozoic, were the first known vertebrates to evolve powered flight.¹,² Arboreal locomotion has been proposed for some taxa,³,⁴ and even considered to have played a role in the origin of pterosaur flight.⁵,⁶ Even so, there is still need for comprehensive quantitative ecomorphological analyses.³,⁴ Furthermore, skeletal adaptations correlated to specialized lifestyles are often difficult to recognize and interpret in fossils. Here we report on a new darwinopteran pterosaur that inhabited a unique forest ecosystem from the Jurassic of China. The new species exhibits the oldest record of palmar (or true) opposition of the pollex, which is unprecedented for pterosaurs and represents a sophisticated adaptation related to arboreal locomotion. Principal-coordinate analyses suggest an arboreal lifestyle for the new species but not for other closely related species from the same locality, implying a possible case of ecological niche partitioning. The discovery adds to the known array of pterosaur adaptations and the history of arborealism in vertebrates. It also adds to the impressive early bloom of arboreal communities in the Jurassic of China, shedding light on the history of forest environments.
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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
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Although there are nine genera of ctenochasmatoids reported from the Jehol Biota, at present each is known from a specimen that has either a skull or a relatively complete postcranial skeleton. A nearly complete juvenile specimen of Gladocephaloideus from the Lower Cretaceous Jiufotang Formation of Sihedang, Lingyuan of Liaoning Province is the most complete ctenochasmatoid preserved to date with a skull and postcranial skeleton. Based on the holotype (IG-CAGS 08–07) and the nearly complete new specimen (JPM 2014–004), the diagnosis of Gladocephaloideus is amended: approximately 50 teeth in total with sharp tips; small nasoantorbital opening, occupying approximately 13% of total skull length; ratio of prenarial rostrum length to skull length approximately 0.63; deep groove along the mid-line of the mandibular symphysis; length to width ratio of the longest cervical vertebra = 4.1; ratio of femur length to tibia length = 0.61; tibia as long as the wing-phalange 1. Phylogenetic analysis recovers Gladocephaloideus within the clade Ctenochasmatidae. Gladocephaloideus has a closer relationship to the Chinese Pterofiltrus rather than to other ctenochasmatid pterosaurs. Microstructure of limb bones implies that JPM 2014–004 represents an early juvenile of Gladocephaloideus jingangshanensis, and that the type specimen is not a fully grown specimen either. We assume that the holotype may equate to the late juvenile or sub-adult developmental stage of Gladocephaloideus.
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Pterosaurs consist of an extinct group of flying reptiles that show short- and long-tailed species. Among those are the anurognathids whose phylogenetic position has been considered quite controversial. So far, there are about 10 described specimens from the Anurognathidae, from which only a few show the preservation of caudal elements. Here, we report a new anurognathid specimen (IVPP V16728) from Mutoudeng, Qinglong, Hebei, China that shows the most complete tail of this non-pterodactyloid clade. The preserved part of the tail has at least 20 caudal vertebrae, some showing extended chevrons and zygapophyses, which is a very primitive character within pterosaurs.
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The Early Cretaceous Jehol Biota of northeastern China has become famous over the last two decades as a source of feathered avialan and non-avialan theropods, preserved alongside an array of other fossil vertebrates, invertebrates, and plants. Still more recently, a rich assemblage referred to in this paper as the Daohugou Biota has begun to emerge from Jurassic strata in the same region. Like their counterparts from the Jehol Biota, Daohugou Biota vertebrate specimens are typically preserved in fine-grained lacustrine beds and often retain feathers and other soft-tissue features. At present, 30 vertebrate taxa (five salamanders, one anuran, two lizards, 13 pterosaurs, five dinosaurs, and four mammals) are known from the Daohugou Biota, which was first recognized at the Daohugou locality in Inner Mongolia. The presence of the salamander Chunerpeton tianyiensis, proposed in this paper as an index fossil for the Daohugou Biota, links the Daohugou locality to five other fossil-producing areas in the provinces of Hebei and Liaoning. The strata containing the Daohugou Biota are close to the Middle–Upper Jurassic boundary and belong at least partly to the regionally widespread Tiaojishan Formation. In general, the vertebrate fauna of the Daohugou Biota is strikingly different from that of the Jehol Biota, although paravian dinosaurs, anurognathid pterosaurs, and salamanders with cryptobranchid and hynobiid affinities occur in both. Nevertheless, the Daohugou Biota and the Jehol Biota are two successive Lagerstätte assemblages that collectively offer a taphonomically consistent window into the Mesozoic life of northeast Asia over a significant span of geologic time.
The leading edge and shape of the pterosaur wing is constrained by the skeleton. Although it has long been known that at least some pterosaurs had posteriorly curved distal wing phalanges, affecting the shape of the wingtip, this has been little studied despite that this may have profound effects on flight performance. Here we examine the evidence for curved wingtips in pterosaurs and evaluate the possible aerodynamic and aeronautical effects. Curved wingtips are shown to be common in both pterosaurs likely to have inhabited terrestrial environments, and those which were strongly pelagic. The recently described genus Bellubrunnus provides new anatomical novelty for pterosaurs having anteriorly directed wingtips and thus likely had a different flight profile to all previously known pterosaurs.
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.
Pterosaurs are represented in China by five genera and some isolated bones ranging in age from the Middle Jurassic to the Late Cretaceous period. Four of these genera belong to the derived monophyletic subgroup Pterodactyloidea; only the Middle Jurassic Angustinaripterus from Dashanpu, Sichuan, is a non-pterodactyloid (traditionally `rhamphorhynchoids', a paraphyletic taxon). Two further pterosaurs (Fig. 1) from the Chaomidianzi Formation of the Beipiao area, western Liaoning Province, occur in the Liaoning beds, several metres higher than the compsognathid coelurosaur Sinosauropteryx and the basal bird Confuciusornis. Our analysis of these two fossils and other components of the fauna suggest a Late Jurassic biostratigraphic age for the Liaoning beds, which are important in the study of avian origins.