Journal of Systematic Palaeontology

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A new tanystropheid (Diapsida:

Archosauromorpha) from the Middle Triassic
of SW China and the biogeographical origin of
Tanystropheidae

Yu-Ting Lu & Jun Liu

To cite this article: Yu-Ting Lu & Jun Liu (2023) A new tanystropheid (Diapsida:
Archosauromorpha) from the Middle Triassic of SW China and the biogeographical
origin of Tanystropheidae, Journal of Systematic Palaeontology, 21:1, 2250778, DOI:
10.1080/14772019.2023.2250778

To link to this article: https://doi.org/10.1080/14772019.2023.2250778

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Published online: 02 Oct 2023.

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 Journal of Systematic Palaeontology, 2023
Vol. 21, No. 1, 2250778
http://dx.doi.org/10.1080/14772019.2023.2250778

A new tanystropheid (Diapsida: Archosauromorpha) from the Middle Triassic

of SW China and the biogeographical origin of Tanystropheidae
Yu-Ting Lu and Jun Liu

Division of Geology, School of Resources and Environmental Engineering, Hefei University of Technology,
193 Tunxi Road, Hefei 230009, China
(Received 4 January 2023; accepted 17 August 2023)

Tanystropheidae is a clade of early archosauromorphs with high morphological disparity and a wide geographical
distribution. The origin and early radiation of Tanystropheidae are still incompletely understood. Here we report
Luxisaurus terrestris gen. et sp. nov., a new archosauromorph collected from the marine Upper Member of Guanling
Formation (Pelsonian substage, Anisian, Middle Triassic) in Luxi County, Yunnan Province, China, and
phylogenetically recovered as a tanystropheid. The morphology of Luxisaurus is consistent with a terrestrial lifestyle.
This is the first occurrence of a tanystropheid found from the Anisian Guanling Formation in SW China and also the
earliest tanystropheid from the eastern Tethys. Phylogenetic analysis shows that Dinocephalosauridae forms the sister
group of Tanystropheidae, Fuyuansaurus occupies the most basal position in Tanystropheidae, and Luxisaurus forms the
sister group to a clade consisting of all other tanystropheids. Considering that Fuyuansaurus and Luxisaurus are found
exclusively in South China, we hypothesize that Tanystropheidae originated in the Early Triassic of South China and
then dispersed globally. The discovery of Early Triassic tanystropheids in South China is needed to further support this
hypothesis. The discovery of Luxisaurus expands the stratigraphical and geographical distribution of Tanystropheidae,
and provides new insights into the origin and early radiation of the clade.
http://zoobank.org/urn:lsid:zoobank.org:pub:07DE12AC-8AE7-48F0-8BAD-0B73F04C0858
Keywords: marine reptile; Dinocephalosauridae; phylogeny; Guanling Formation

Introduction Protorosaurus speneri, Tanystropheidae, Rhynchosauria and

Allokotosauria (Ezcurra, 2016; Spiekman et al., 2021),
As the largest ecological disaster in Earth’s history, the among which Tanystropheidae reached peak diversity dur-
Permo–Triassic Mass Extinction (PTME) led to the ing the Middle Triassic.
extinction of almost all life on Earth, and biological The known Tanystropheidae fossil record extends from
recovery from this event began gradually in the Triassic the Early Triassic (Elessaurus gondwanoccidens from the
(Chen & Benton, 2012). During the Triassic, terrestrial Induan–Olenekian of Southern Brazil) (De-Oliveira et al.,
reptiles diversified, and marine reptiles began to domin- 2020) to the Late Triassic (Sclerostropheus fossai from the
ate the oceans (Q. Li & Liu, 2020; Qiao et al., 2022). late Norian of Italy) (Rigo et al., 2009; Tackett & Tintori,
Since Triassic reptiles are generally positioned in the 2019). While most basal archosauromorphs are terrestrial,
higher levels of the food chain, revealing their true tanystropheids are an exception. Tanystropheidae has a
diversity in the fossil Lagerst€atten is important in under- worldwide distribution, which led to the development of
standing the timing and pattern of recovery from the different body plans and adaptations to distinct environ-
PTME (Benton et al., 2013; Chen & Benton, 2012; ments, including terrestrial, semi-aquatic, entirely aquatic,
Motani et al., 2015; Scheyer et al., 2014). and possibly gliding forms (Liu et al., 2017; Miedema
Archosauromorpha is a diverse clade of diapsid reptiles, et al., 2020; Olsen, 1979; Rieppel et al., 2008; Spiekman
which originated in the Permian and then gradually radiated et al., 2020; Vecchia, 2000). Among them, Macrocnemus,
to become one of the dominant vertebrate groups in the Elessaurus gondwanoccidens and Langobardisaurus are
Triassic (Butler et al., 2015; De-Oliveira et al., 2020; considered to be terrestrial forms (De-Oliveira et al., 2020;
Ezcurra, 2016; Liu et al., 2017; Pritchard et al., 2015). The Rieppel, 1989; Saller et al., 2013), while Fuyuansaurus
earliest members of Archosauromorpha are the non-archo- acutirostris, Tanytrachelos ahynis and Tanystropheus
sauriform archosauromorphs, including Prolacerta broomi, hydroides are considered to be typically aquatic forms

Corresponding author. Email: [email protected]

# The Trustees of the Natural History Museum, London 2023. All rights reserved.

Published online 02 Oct 2023

2 Y.-T. Lu and J. Liu

(Casey et al., 2007; Fraser et al., 2013; Spiekman et al., Hefei, Anhui, China; PIMUZ, Palaeontological Institute
2020). Augustaburiania vatagini is suggested to be semi- and Museum, Zurich, Switzerland.
aquatic (Sennikov, 2011). Ozimek volans may even glide
between trees (Dzik & Sulej, 2016).
Many phylogenetic analyses of Tanystropheidae have Material and methods
been carried out, but the results are controversial (De-
Oliveira et al., 2020; Liu et al., 2017; Pritchard & The new material comprises the proximal caudal verte-
Nesbitt, 2017; Pritchard et al., 2015, 2016; Spiekman brae, gastralia, and some part of the pelvic girdle,
et al., 2021). These conflicting phylogenetic hypotheses together with most of the limbs (Fig. 2). The specimen
may be the result of inadequate sampling of tanystrop- was prepared using mounted needles in the palaeonto-
heid taxa. Up to now, the most comprehensive phylo- logical laboratory of HFUT. To test the phylogenetic
genetic analysis was conducted by Spiekman et al. affinities of Luxisaurus terrestris, we used the matrix
(2021), but one of the notably unsolved issues of this compiled by Spiekman et al. (2021). Petrolacosaurus
analysis is that the three species of Macrocnemus have kansensis was designated as outgroup. The analysis was
not been recovered as a monophyletic clade. Since the conducted using the ‘Traditional Search’ option in TNT
phylogenetic position of some tanystropheids is still
1.5 (Goloboff & Catalano, 2016). We performed 10,000
uncertain, this also hinders the understanding of the bio-
replicates of Wagner trees using random addition
geographical origin and early evolution of
sequences, followed by Tree Bisection Reconnection
Tanystropheidae (De-Oliveira et al., 2020).
branch swapping (10 trees held per replicate). To
In recent years, many basal archosauromorphs have
explore branch support, we used the ‘Bootstrap support’
been found in the marine Triassic of South China,
option for 1000 iterations in TNT to calculate decay
including both tanystropheids and dinocephalosaurids.
indices. Bremer support values and consistency and
Dinocephalosauridae (Spiekman et al., 2021) from
South China are all known from the Upper Member of retention indices were calculated using the TNT scripts
the Anisian Guanling Formation, including Pectodens Bremer.run and Stats.run.
zhenyuensis (C. Li et al., 2017) and Dinocephalosaurus
orientalis (Liu et al., 2017; Rieppel et al., 2008).
Tanystropheids from South China are all known from Systematic palaeontology
the late Ladinian–early Carnian Zhuganpo Formation,
including Tanystropheus (C. Li et al., 2007; Rieppel Subclass Diapsida Osborn, 1903
et al., 2010), Macrocnemus (Jiang et al., 2011; C. Li Clade Archosauromorpha Huene, 1946
et al., 2007; Scheyer et al., 2020; Zhang et al., 2010) Family Tanystropheidae Camp, 1945
and Fuyuansaurus (Fraser et al., 2013). Luxisaurus gen. nov.
The Upper Member of the Anisian Guanling
Formation is widely distributed in Guizhou and Yunnan Type species. Luxisaurus terrestris sp. nov.
provinces in SW China. It is comprised of nodular
micritic limestone, marl, and arenaceous limestone with Diagnosis. As for the type species below.
dolomite. While all known Chinese dinocephalosaurids Etymology. The genus name is derived from the
were reported from this member (C. Li et al., 2017; Liu Chinese Pinyin of Luxi County, where the holotype was
et al., 2017; Spiekman et al., 2021), no tanystropheids collected.
had previously been found. Here, we report a new taxon
of Tanystropheidae from the Upper Member of the Luxisaurus terrestris sp. nov.
Anisian Guanling Formation (Fig. 1), representing the
(Figs 2–6)
earliest occurrence of the group in the eastern Tethys. In
addition to the description of the skeletal anatomy of
the new taxon and the interpretation of its lifestyle, we Holotype. HFUT SML-21-08-001, an incomplete skel-
also attempt to clarify the phylogenetic interrelationships eton preserving most of the limbs, the proximal caudal
of Tanystropheidae and discuss the palaeobiogeographi- vertebrae, and some elements of the pelvic girdle. The
cal origin of the clade. specimen is now catalogued in the Geological Museum
of Hefei University of Technology.
Institutional abbreviations Horizon. Upper Member, Guanling Formation, Anisian,
GMPKU, Geological Museum of Peking University, Middle Triassic. Conodont analysis suggested a
Beijing, China; HFUT, Hefei University of Technology, Pelsonian age for the fossil assemblages preserved in
 A new archosauromorph from the Triassic of China 3

Figure 1. Geological map showing the quarry where Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001) was discovered
(updated from Y. W. Hu & Liu, 2022). Inset is the map of China. Abbreviations: E, Palaeogene; T2f, Falang Formation, Ladinian,
Middle Triassic; T2y, Yangliujing Formation, Anisian–Ladinian, Middle Triassic; T2g2, Upper Member of Guanling Formation,
Anisian, Middle Triassic; T2g1, Lower Member of Guanling Formation, Anisian, Middle Triassic; T1, Lower Triassic.

Luxi (Wen et al., 2020), the same as those found in a weak hook shape; the first phalanx of digit 5 on the
Luoping and Panzhou (Liu, 2015; Sun et al., 2006). pes approximately equal to metatarsal 5; tibia and fibula
more than 20% longer than femur; fibula much thinner
Locality. The specimen was collected from a quarry
than tibia; manual phalangeal formula of 2-3-4-4-3;
about 2 km east of Suomeiluo, Luxi County, Yunnan
presence of a longitudinal ridge along the lateral surface
Province, SW China (Fig. 1).
of the pedal unguals; length of the longest metatarsal
Etymology. The species name refers to the lifestyle of less than twice that of the longest metacarpal; length of
the specimen, which is terrestrial. metatarsal 3 more than twice that of metatarsal 1.
Diagnosis. Unguals slightly longer than the last non-
ungual phalanx of the same digit in the manus; lateral Description
side of metatarsal 5 approximately straight; medial side HFUT SML-21-08-001 is an incomplete skeleton,
of metatarsal 5 concave; medial side of the proximal including most of the limbs and 14 proximal caudal ver-
end of metatarsal 5 with a triangular protrusion, forming tebrae, and some elements of the pelvic girdle and
4 Y.-T. Lu and J. Liu

Figure 2. Photograph and anatomical interpretation of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001). A, photograph;
B, line drawing. Abbreviations: as, astragalus; ca, calcaneum; cal, carpal; dt, distal tarsal; fe, femur; fi, fibula; il, ilium; is, ischium;
mc, metacarpal; mt, metatarsal; pu, pubis; r/g, ribs and gastralia; ra, radius; ti, tibia; ul, ulna.

gastralia (Fig. 2). HFUT SML-21-08-001 shares many and the remaining caudals are exposed in lateral view.
synapomorphies of Tanystropheidae (see below) that The size of caudal ribs gradually decreases posteriorly.
have been previously recognized (Jaquier et al., 2017; The distal ends of the first and second caudal ribs are
Liu et al., 2017; Nopcsa, 1930; Peyer, 1937; Rieppel, sharp (especially in the first caudal), but more posterior
1989; Scheyer et al., 2020; Spiekman et al., 2021). caudal ribs have a broad distal end. There are seven chev-
rons preserved, and the first appear on the seventh caudal.
Axial skeleton. HFUT SML-21-08-001 has two sacral
vertebrae with articulated sacral ribs (Fig. 2). On the Forelimb. The forelimbs are not completely preserved,
right side, the first sacral rib is lost, and only the prox- and the left and right forelimbs cannot be differentiated
imal part of the second sacral rib is preserved. Left (Fig. 3). In addition, the proximal ends of the ulna and
sacral ribs are not completely exposed, and the distal radius, as well as the complete humeri, are missing. The
ends of the sacral ribs are hidden underneath the ilium. manus is slender. The distal ends of the radius and ulna
The sacral ribs are narrow proximally and broadened are straight.
distally. The second sacral rib is broader than the first A single carpal is preserved in each manus. The car-
sacral rib. The second sacral rib bifurcates distally into pal of one manus is round, and located distal to the ulna
anterior and posterior processes, as in Langobardisaurus and proximal to metacarpals 4 and 5. The carpal of the
and Macrocnemus (Scheyer et al., 2020; Spiekman other manus is approximately round, and located imme-
et al., 2021). The sacral ribs are not fused with the cor- diately distal to the spatium interosseum between the
responding vertebrae, a feature that is also seen in the radius and ulna and proximal to metacarpal 4.
adult Dinocephalosaurus (Liu et al., 2017) and M. The metacarpals are closely arranged. They increase
fuyuanensis (PIMUZ T 1559: see Jaquier et al., 2017; in length from the first to the fourth metacarpal, with
Scheyer et al., 2020). There are a few dark brown areas metacarpal 4 being the longest while metacarpal 5 is the
that display a mottled pattern near the proximal part of shortest (Table 1). The second phalanx of digit 2 is
the left femur and in the region of the pelvis, which shorter than the first phalanx of digit 2 in the manus.
may be coprolites. The terminal phalanx of each digit forms an ungual,
There are 14 caudal vertebrae preserved and exposed, which is slightly longer than the last phalanges of the
and the distal-most part of the tail is missing. The first same digit (Table 1). The manus is completely pre-
to tenth caudal vertebrae are exposed in dorsal view, served, and the phalangeal formula is 2-3-4-4-3.
 A new archosauromorph from the Triassic of China 5

Figure 3. Forelimb of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001). A, photograph; B, line drawing.
Abbreviations: cal, carpal; d, digit; mc, metacarpal; ra, radius; ul, ulna.

Table 1. Length (mm) of the metacarpals and phalanges in the manus Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-
001). A and B represent different manus.a, ungual phalanx.
A B
Digit 1 2 3 4 5 1 2 3 4 5
Metacarpal 5.35 9.89 12.40 14.09 4.31 5.40 9.82 12.35 13.55 4.39
Phalanx 1 3.14 2.95 4.18 4.88 2.27 2.82 2.81 4.21 5.16 2.25
Phalanx 2 2.72a 2.41 2.25 2.81 2.32 2.80a 2.13 2.37 2.53 2.03
Phalanx 3 2.7a 2.30 2.01 2.48a 2.89a 2.05 2.10 2.67a
Phalanx 4 2.86a 2.49a 2.74a 2.78a

Pelvic girdle. The left pelvic region is completely pre- HFUT SML-21-08-001 has a well-preserved ankle
served (Fig. 4). Except for the left ilium, their outlines and pes, and elongated metatarsals and phalanges. The
are difficult to discern. The left ilium is well exposed in three ossified tarsals are located in close articulation
lateral view. The preacetabular margin of the ilium is with each other in both ankles. The largest tarsal is
smooth without a distinct process or tuber, and the iliac located distal to the tibia and represents the astragalus
blade is rather elongated and directed posteriorly. The (Fig. 6). The second-largest tarsal is located distal to the
dorsal iliac blade has a preacetabular process, but it fibula and represents the calcaneum. The foramen for
does not project beyond the level of the anterior margin the passage of the perforating artery between the astrag-
of the acetabulum portion. The ilium has a dorsally alus and calcaneum does not exist, which is unique
rimmed caudifemoralis brevis muscle origin on the later- among tanystropheids, but convergently evolved in the
oventral surface of the postacetabular process. The left Chinese dinocephlosaurids (C. Li et al., 2017; Rieppel
ischium is covered by the ilium, and the left pubis is et al., 2008). The smallest tarsal ossification is the distal
covered by the second sacral vertebra and the ilium. tarsal 4, which is approximately circular.
Hindlimb. The right hindlimb is completely preserved All metatarsals are completely preserved (Fig. 5).
except for the femur, while the left hindlimb is com- There is a distinct asymmetry in the length of the meta-
pletely preserved (Fig. 5). The gracile hindlimb resembles tarsals, with metatarsal 4 being the longest, followed
that of Macrocnemus and Tanystropheus, as described in by metatarsals 3, 2, 1 and 5 (Table 3). Metatarsals are
earlier studies (Fraser & Furrer, 2013; Nopcsa, 1930; similar to Macrocnemus, Tanystropheus, Elessaurus
Nosotti, 2007; Scheyer et al., 2020). The femur is a rela- gondwanoccidens, Amotosaurus rotfeldensis and
tively slender element, slightly twisted into an ‘S’ shape. Langobardisaurus in that they are tightly bunched
The tibia and fibula are slender elements, subequal in (Casey et al., 2007; De-Oliveira et al., 2020; Fraser &
length, 23% longer than the femur (Table 2). The trans- Rieppel, 2006; Nosotti, 2007; Rieppel, 1989; Saller
verse width at mid-length of the fibula is distinctly nar- et al., 2013; Spiekman et al., 2021). The most character-
rower than the transverse width of the tibia. istic element of the pes is metatarsal 5. It is thick, short,
6 Y.-T. Lu and J. Liu

Figure 4. The pelvic region of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001). A, photograph; B, line drawing.
Abbreviations: ac, acetabulum; cav, caudal vertebra; dv, dorsal vertebra; il, ilium; ilb, iliac blade; is, ischium; prap, preacetabular
process; pu, pubis; pup, pubic peduncle; sr, sacral rib; sv, sacral vertebra.

Figure 5. Photograph and anatomical interpretation of the posterior part of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-
001). A, photograph; B, line drawing. Abbreviations: as, astragalus; ca, calcaneum; car, caudal rib; cav, caudal vertebra; d, digit;
dt, distal tarsal; dv, dorsal vertebra; fe, femur; fi, fibula; ha, haemal arch; il, ilium; is, ischium; mt, metatarsal; pu, pubis; r/g, ribs
and gastralia; sr, sacral rib; sv, sacral vertebra; ti, tibia.
 A new archosauromorph from the Triassic of China 7

and robustly built. The proximal and distal ends are phalanges represent pointed unguals. The lateral surface
expanded. Metatarsal 5 has a unique morphology, with of the pedal unguals has a clear longitudinal ridge. The
an approximately straight lateral margin and a concave length of the metatarsal is close to that of the most
medial margin. A triangular protrusion on the medial proximal phalanx in digit 5. The phalanges are well pre-
margin of the proximal end of metatarsal 5 forms a served, with a phalangeal formula of 2-3-4-5-4.
weakly hooked shape.
All phalanges are preserved intact. Phalanx 1 of digit
5 on the pes is short, similar to Macrocnemus (Fraser &
Phylogenetic analysis
Furrer, 2013); the length of this phalanx 1 is approxi-
mately equal to that of metatarsal 5. The distal-most
To test the phylogenetic position of Luxisaurus terrest-
ris, we conducted a phylogenetic analysis based on the
Table 2. Measurements (mm) of the hindlimb elements in data matrix of Spiekman et al. (2021). The data matrix
Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001). used in our phylogenetic analysis includes the original
Abbreviations: DW, width of distal end; L, length; PW,
307 characters of Spiekman et al. (2021), but an add-
width of proximal end.
itional state 2 is added for character 262: [manual dig-
Left Right
its, unguals length: about the same length as or shorter
Bone L DW PW L DW PW than the last non-ungual phalanx of the same digit (0);
Femur 52.84 6.06 7.89 – – – slightly longer than the last non-ungual phalanx of the
Tibia 65.19 4.75 5.81 65.06 5.09 6.17 same digit (1); distinctly longer than the last non-
Fibula 64.64 2.96 1.9 64.81 2.67 1.66
ungual phalanx of the same digit (2)]. In addition to

Figure 6. The photograph of the right pes of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-001). Abbreviations: as,
astragalus; ca, calcaneum; d, digit; dt, distal tarsal; fi, fibula; mt, metatarsal; ti, tibia.

Table 3. Length (mm) of the metatarsals and phalanges in the pes of Luxisaurus terrestris gen. et sp. nov. (HFUT SML-21-08-
001). a, ungual phalanx.
Left Right
Digit 1 2 3 4 5 1 2 3 4 5
Metatarsal 10.93 19.28 22.69 25.09 7.47 10.57 18.81 22.74 26.07 7.31
Phalanx 1 4.37 4.64 6.51 8.82 7.33 4.98 4.67 6.42 8.45 7.46
Phalanx 2 4.23a 3.35 3.57 4.9 3.58 4.23a 3.65 3.48 5.01 3.47
Phalanx 3 4.54a 3.31 3.57 3.42 4.53a 3.64 3.38 3.36
Phalanx 4 4.16a 3.23 3.93a 4.27a 3.65 4.09a
Phalanx 5 3.58a 3.92a
8 Y.-T. Lu and J. Liu

Figure 7. Phylogenetic hypothesis for Luxisaurus terrestris gen. et sp. nov. Bremer values above 1 and bootstrap frequencies above
50% are provided above and below the nodes, respectively.
 A new archosauromorph from the Triassic of China 9

Luxisaurus terrestris, the operational taxonomic units The remaining tanystropheids form a clade, which is sup-
included in our data matrix are the same as those in ported by three unambiguous synapomorphies: ratio of
analysis 4 of Spiekman et al. (2021), the final analysis length of the longest metacarpal to length of the longest
accepted by them. The analysis (see Supplemental metatarsal between 0.36 and 0.41 (ch. 258: 2 ! 1); the
material) generated 54 most parsimonious trees, with distal condyles of the femur are prominent and show
1246 steps, a consistency index of 0.357, and a reten- strong dorsoventral expansion (in sprawling orientation)
tion index of 0.535. Our analysis recovers monophy- restricted to the distal end (ch. 281: 1 ! 0); ratio of
letic Sauria, Lepidosauromorpha, Archosauromorpha, length of metatarsal 1 to metatarsal 3 between 0.54 and
Crocopoda, Rhynchosauria, Allokotosauria and 0.63 (ch. 297: 1 ! 2).
Archosauriformes, as in the previous phylogenetic
study (Spiekman et al., 2021).
Protorosaurus speneri is recovered as the sister Discussion
taxon to all remaining Archosauromorpha, consisting
of Jesairosaurus lehmani, Tanystropheidae, Ontogenetic stage of the holotype of Luxisaurus
Dinocephalosauridae and Crocopoda (Fig. 7). terrestris
Jesairosaurus lehmani is recovered as the sister taxon to HFUT SML-21-08-001 (the holotype of Luxisaurus ter-
the unnamed clade consisting of Tanystropheidae and restris) only preserves part of the limbs and 14 proximal
Dinocephalosauridae. Dinocephalosaurus orientalis and caudal vertebrae, and some elements of the pelvic gir-
Pectodens zhenyuensis comprise the monophyletic dle, which makes the evaluation of ontogenetic stage
Dinocephalosauridae, which is supported by six unam- difficult. However, there are still several morphological
biguous synapomorphies: posterior process of jugal absent features that are useful in assessing the ontogenetic
(ch. 42.1); glenoid fossa considerably displaced ventrally stage of the specimen. The distal ends of radius, ulna,
compared to the tooth row (ch. 161.1); anterior free-end- tibia and fibula are straight, indicating that the holotype
ing process (¼ accessory process) on anterior surface of likely represents an osteologically immature individual
anterior cervical ribs present and long, extending anterior (Hugi & Scheyer, 2012). Although the metacarpals are
to the prezygapophyses of the corresponding vertebra tightly articulated with the phalangeal elements, there is
when in articulation (ch. 200.2); ratio of total length of a large gap between the carpals and metacarpals. In add-
the humerus to total length of the femur between 0.84 ition, there is only one ossified carpal and three ossified
and 1.05 (ch. 248.2/3); hook-shaped proximal end absent tarsals. The sacral rib is also not fused with the corre-
on metatarsal 5 (ch. 300.0); ratio of length of metatarsal 4 sponding vertebra. All these morphological features sug-
to proximodistal length of metatarsal 5 between 1.25 and gest that the holotype is a juvenile (Hugi & Scheyer,
1.90 (ch. 305.0). 2012; C. Li, 2007; Premru, 1991; Rieppel, 1992, 1993).
Tanystropheidae is supported by four unambiguous Although establishing a new taxon based on a juvenile
synapomorphies: distal margin of anterior and mid-post- is not ideal, Luxisaurus differs from dinocephalosaurids
axial cervical neural spines completely straight along and other tanystropheids in many respects, as noted
anteroposterior length in lateral view (ch. 179.1); anterior below.
margin of the neural spine of anterior and mid-postaxial
cervical vertebrae is anterodorsally inclined at an angle of Morphological comparison between Luxisaurus
less than 60 from the horizontal plane (ch. 194.2); prox- terrestris, other tanystropheids and
imal end of anterior dorsal ribs is dichocephalous (two dinocephalosaurids
facets) (ch. 213.1); caudifemoralis brevis muscle origin The manual phalangeal formula of Luxisaurus is 2-3-4-
on the lateroventral surface of the postacetabular process 4-3. This differs from the manual phalangeal formula of
of ilium, dorsally rimmed by a low brevis shelf (ch. Macrocnemus, Langobardisaurus and Pectodens, which
269.1). Our phylogenetic analysis shows that the three is 2-3-4-5-3, but is the same as that of Tanystropheus
species of Macrocnemus are recovered as a monophyletic and Tanytrachelos (see table 9 of Nosotti, 2007).
clade. Macrocnemus is supported by two unambiguous Metacarpal 4 is longer than metacarpal 3, which is dif-
synapomorphies: maxillary teeth are distinctly recurved ferent from Tanystropheus hydroides, Tanystropheus
(ch. 171.1); maximum height of postaxial anterior or mid- longobardicus and Dinocephalosaurus (Nosotti, 2007;
cervical neural spines is approximately equal in height to Rieppel et al., 2008). The tibia is longer than the femur,
the posterior articular surface of the centrum (ch. 178.1). which is different from the condition in Tanystropheus,
Fuyuansaurus acutirostris occupies the most basal pos- Langobardisaurus and Tanytrachelos ahynis (Casey
ition in Tanystropheidae, and Luxisaurus terrestris com- et al., 2007; Nosotti, 2007; Saller et al., 2013). The
prises the sister taxon to all remaining tanystropheids. femur shaft exhibits sigmoidal curvature, being different
10 Y.-T. Lu and J. Liu

Figure 8. Time-scaled phylogenetic tree of Dinocephalosauridae and Tanystropheidae. The grey box highlights the basal Chinese
dinocephalosaurids and tanystropheids.

from Macrocnemus obristi, Langobardisaurus and Langobardisaurus, Tanytrachelos ahynis, Elessaurus and
Tanytrachelos ahynis (Casey et al., 2007; Fraser & Dinocephalosaurus (Casey et al., 2007; De-Oliveira et al.,
Furrer, 2013; Saller et al., 2013). Metatarsals 1–4 are 2020; Nosotti, 2007; Rieppel et al., 2008; Saller et al.,
tightly bunched, which differs from Dinocephalosaurus 2013). The second sacral rib bifurcates distally, which is
and Tanytrachelos ahynis (Casey et al., 2007; Rieppel different from Tanystropheus, Raibliania calligarisi and
et al., 2008). Metatarsal 5 forms a weak hook shape, which Tanytrachelos ahynis (Casey et al., 2007; Dalla Vecchia,
is different from Macrocnemus, Dinocephalosaurus and 2020; Nosotti, 2007).
Pectodens (C. Li et al., 2017; Nosotti, 2007; Rieppel et al., Luxisaurus also shows several autapomorphies among
2008). The ratio of the length of metatarsal 4 to the length tanystropheids, including unguals slightly longer than
of metatarsal 3 is higher than 1.13. This is different from the last non-ungual phalanx of the manus, ratio of the
Tanystropheus hydroides, Tanystropheus longobardicus, length of the longest metacarpal to the longest
 A new archosauromorph from the Triassic of China 11

Figure 9. Triassic palaeogeographical map showing the distribution of Dinocephalosauridae and Tanystropheidae. The colour code of
taxa in the palaeogeographical map is same as the colour code used in the geological time scale (produced from the Paleobiology
Database, https://paleobiodb.org/#/). 1. Sanga do Cabral Formation, Early Triassic, Brazil (De-Oliveira et al., 2018); 2. Jilh
Formation, Middle Triassic, Saudi Arabia (Vickers-Rich et al., 1999); 3. Moenkopi Formation, Middle Triassic, Arizona and New
Mexico (Formoso et al., 2019); 4. Middle–Late Triassic, Italy (Dalla Vecchia, 2000, 2005); 5. Middle Triassic, Spain and France
(Spiekman & Scheyer, 2019); 6. Late Triassic, Canada (Sues & Olsen, 2015) and Chinle Formation, New Mexico (Pritchard et al.,
2015).

metatarsal higher than other tanystropheids, ratio of the gondwanoccidens) are older than the tanystropheids
length of metatarsal 1 to metatarsal 3 lower than other from South China (Fig. 8), our phylogenetic results
tanystropheids, and the presence of a longitudinal ridge show that Fuyuansaurus acutirostris and Luxisaurus ter-
along the lateral surface of the unguals. In addition, the restris from South China comprise the consecutive sister
phylogenetic results (Fig. 7) show Luxisaurus forms the groups to all other tanystropheids. In particular,
sister group to a clade consisting of all other tanystrop- Dinocephalosauridae, the sister taxon of
heids except Fuyuansaurus. Tanystropheidae, is almost exclusively known from
South China (see Fig. 9, but note that Tanystropheus
antiquus from the Lower Muschelkalk was tentatively
Palaeobiogeographical implications recovered as a dinocephalosaurid in analyses 1 and 3
Previously, tanystropheids from the eastern Tethys were
of Spiekman et al., 2021). Based on the above phylo-
all known from the late Ladinian–early Carnian
genetic hypothesis, we put forward a palaeobiogeo-
Zhuganpo Formation in SW China (Fraser et al., 2013;
graphical hypothesis that Tanystropheidae originated
C. Li, 2007; Rieppel et al., 2010; Zhang et al., 2010).
in the Early Triassic of South China and then dispersed
Luxisaurus terrestris from the eastern Tethys is of inter-
globally. New tanystropheids from the Early Triassic of
est because, in addition to extending the stratigraphical
South China and sampling from other regions of the
and geographical distribution of Tanystropheidae, it rep-
world (e.g. South America) are needed to test this
resents the first tanystropheid from the Anisian
hypothesis.
Guanling Formation and also the earliest tanystropheid
from the eastern Tethys. Although some species of
Tanystropheidae from other regions (Olenekian Palaeoecological implications
Augustaburiania vatagini, early Anisian Amotosaurus By the end of the Early Triassic, tanystropheids had diver-
rotfeldensis and the Early Triassic Elessaurus sified both morphologically and ecologically, including
12 Y.-T. Lu and J. Liu

terrestrial, semi-aquatic, entirely aquatic, and even gliding tanystropheid in the eastern Tethys. The morphological
forms (Benton, 2016; Dzik & Sulej, 2016; Ezcurra, 2016; characteristics of the specimen and phylogenetic results
Renesto, 2005). Some tanystropheids are considered terres- unambiguously recover Luxisaurus terrestris as a tany-
trial, including Macrocnemus, Elessaurus gondwanocci- stropheid. Based on the phylogenetic results of this
dens and Langobardisaurus. The structure of the limb, study, we hypothesize that Tanystropheidae originated
such as the hooked metatarsal 5 and elongate digits, in the Early Triassic of South China after the PTME,
together with the claw-like distal phalanges and the distally and then dispersed globally. In addition, the limb struc-
bifurcating pleurapophy on the second sacral vertebra, ture suggests that Luxisaurus terrestris is a terrestrial
indicates that the limb morphology conforms to the terres- reptile, further confirming the existence of terrestrial
trial habit (De-Oliveira et al., 2020; Renesto & Avanzini, ecosystems in the marine Guanling Formation.
2002; Rieppel, 1989). Some tanystropheids are considered
to be semi-aquatic or even completely aquatic, including
Fuyuansaurus, Augustaburiania vatagini, Tanytrachelos Acknowledgements
ahynis and Tanystropheus hydroides. Evidence supporting
We thank Y. W. Hu, Q. Li, A. S. Wolniewicz and J. He
an aquatic lifestyle includes the extreme length of the
for the discussion of the specimen, other members of the
neck, the flattened shape of the snout, and the placement
Paleontological Lab of HFUT for their help in the field,
of the external nares on its dorsal surface (Fraser &
T. Sato and L. Y. Li for preparation of the specimen, and
Furrer, 2013; Nosotti, 2007; Olsen, 1979; Sennikov, 2011;
S. N. F. Spiekman for help in the phylogenetic analysis.
Spiekman et al., 2020; Tschanz, 1986). Ozimek volans has
Constructive comments from S. N. F. Spiekman and
the elongated appendages, a flight membrane extending to
N. C. Fraser helped improve the manuscript.
the tips of the hind appendages, an enlarged coracoid, and
possibly an ossified sternum, indicating that it may even
glide between trees (Dzik & Sulej, 2016).
Luxisaurus has elongated and flexible limbs. The long, Supplemental material
slender limbs with pronounced articular ends suggest that
the joint is flexible, which enables Luxisaurus to perform Supplemental material for this article can be accessed
terrestrial locomotion with flexible limb joints (C. Li here: https://doi.org/10.1080/14772019.2023.2250778.
et al., 2017). In addition, the fore- and hindlimbs retain
the claw-shaped terminal phalanges, and the configuration
of the metatarsal-like proportions of the first phalanx of Funding
digit 5 indicates that the pes is likely adapted for the pro-
pulsive phase of a stride on land (Rieppel, 1989). The This work was supported by the National Natural
hook shape of metatarsal 5 can provide leverage for pedal Science Foundation of China (42172026 and 41772003)
plantarflexion, resulting in a propulsive force (Rieppel, and the Department of Natural Resources of Anhui
1989). In conclusion, the limb structure of Luxisaurus Province (2021-g-2-16).
suggests that it is a terrestrial reptile. The vertebrate fau-
nas of the Upper Member of the Anisian Guanling
Formation are mainly composed of marine reptiles and ORCID
fishes, but occasionally terrestrial reptiles are also recov-
ered, such as Pectodens zhenyuensis (C. Li et al., 2017). Jun Liu http://orcid.org/0000-0001-7859-5209
Luxisaurus terrestris is the second terrestrial reptile
reported from the Guanling Formation. The occurrence of
terrestrial reptiles, as well as the terrestrial plants and References
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