The Triassic, as the first period of the Mesozoic era, lasted for 51 million years (251.2 - 201.3 mya) (table 1). It represents a severe reduction in species, following the mass extinction at the end of the Permian Period. Only about 4 % of the world's marine species, and an estimated 30% of terrestial species, survived the Permian-Triassic Mass Extinction (Benton 2005; Sahney and Benton 2008). Also occurring was the only known mass extinction of insects, with 57% of all insect families and 83% of all insect genera becoming extinct (Labandeira and Sepkoski 1993).
majority of dominant land plant species also disappeared. Glossopterids
were one of the major casualties of the end-
of-Permian, mass extinction event. The
Glossopterid family, which arose in the Southern Hemisphere by the
Early Permian period, became the dominant elements of the
southern flora through the rest of the Permian. Glossopteris
woody, seed-bearing tree, growing as high as 30 meters. They
are interpreted to have thrived in very wet soil
have been found in
all of the southern continents, with over 70 fossil species recognized
Fig.1: Fossilized leaves and stem from Glossopteris, a Permian tree.
across several, now detached,
Austrian geologist Eduard
Suess to infer that these areas had once been connected by a land
named this large land mass Gondwanaland, after the district in
Direct effects of the Permian-Triassic
extinction lasted at least through
the Early Triassic, a period of five million years. The mass
extinction was probably
by a combination of volcanic activity, global warming, and increasingly
arid climate in
the mega-continent Pangea (fig.2). At the
the start of the Early Triassic, during the Induan stage (252-251 mya),
plates were joined into Pangea except China, Indonesia, and the
zone including Thailand and parts of Turkey, the latter areas being
islands in the Tethys Sea.
Regarding volcanism as a likely cause of radical atmospheric changes, much attention has been paid to evidence provided by the extensive lava fields known as the Siberian Trapss, a vast area in Russian Siberia extending for about 3000 km. The Siberian Traps indicate massive volcanism which occurred between 252.6 and 250.3 mya, starting at the very end of the Permian, and lasting through the initial Induan Stage and part of the Olenikian stage of the Early Triassic (Kump et al. 2005; Knoll et al. 2007; Ogdena and Sleep 2011).
Fig.2: Pangaea at the start of the Early Triassic (Induan Stage).
The Siberian Traps consist primarily of basalt and tuff formations. The term "traps" is derived from "trappa", the Swedish word for stairs, referring to the step-like hills of the region, which were created by massive outpourings of lava that hardened into basalt. The major eruptive events which formed the traps, the largest known over about 500 million years since the Cambrian period, are thought to have resulted not only in the extensive outpourings of lava forming the basalt traps, but also frequent explosions and coal/gas fires with significant impact on the atmosphere. In combination, these released enough acid and carbon dioxide over an extended period, to have critically damaged both the marine environment and the Earth's atmosphere and its vegetation.
Some evidence suggests that equatorial temperatures exceeded the thermal tolerance for many marine vertebrates over a long period, spanning at least two thermal maxima , whereas terrestrial equatorial temperatures were sufficiently severe to suppress plant and animal abundance during most of the Early Triassic (Sun et al 2013). Consequences included atmospheric and oceanic warming by 5–10° C, anoxia in the oceans, and acid rain leading to massive killing of forests. Most damaging to marine life forms were acidification and anoxia (Knoll et al., 2007); while on land, most harm was caused by acid rain and an increasingly arid climate (Benton and Newell, in press; Benton et al. 2013)
Table 1: Time chart of the Triassic Period, showing stages.
The Early Triassic fossil record
are best represented in the Beaufort Group of
Other components of the Early
Triassic fossil record occur
Early Triassic vertebrates
are also recorded in the Puesto Viejo
formation of Argentina, equjivalent to
During the Early Triassic, Induan and Ikenikian stages (252-247 mya; table 1) some entirely new groups of large aquatic vertebrates appeared, as evidenced by findings in the Yunnan and Ghouzou deposits in south China and elsewhere (Benton et al. 2013). This included first major radiation of marine reptiles, including ichthyosaurs, sauropterygians, and thalattosaurs. In the Permian, there had been only a short-lived expansion of marine reptiles around the southern shores of Gondwanaland. These late Paleozoic marine reptiles are collectively called mesosaurs, a mixed grouping which includes various unrelated taxa (McGowan and Motani, 2003).
While many families of fish disappeared in the massive marine extinction, during the Early Triassic there was a large radiation of bony fish, especially the actinopterygians or spiny-fin groups. Marine invertebrates also saw major changes. The most common Paleozoic groups such as brachiopods, trilobites, and rugose and tabulate corals all became extinct, to be replaced during the Early and Middle Triassic (252-235 mya) by echinoderms, mollusks (both bivalves and gastropids), arthropods, ammonites, and scleractinian corals (Benton et al. 2013).
many groups of
during the end-of-Permian extinction,
to be replaced by new
Early Triassic land vertebrates, two main patterns occurred throughout
the continental land mass called Pangea
(Colbert 1973; Sun 1980).
mainly of labyrinthodont amphibians, with little evidence of reptiles
except occasional trace
fossils (i.e., footprints) of smaller reptiles such as Chilotherium.
existed in a portion of Laurasia from
second pattern, much more widely distributed in the southern
Pangean region of Gondwanaland, comprising
Fig.3: Lystrosaurus, a small herbivorous dicynodont common throughout Pangea in the Early Triassic.
Procolophonoids (fig.4) also have a very wide Early Triassic
distribution, their fossils having
been found in
Fig.4: Procolophon, a small anapsid reptile that was widespread in Pangea during the Early Triassic.
of the anapsid Procolophonis
in the Early Triassic Fremouw Formation of Antarctica has helped
faunal distribution in Gondwanaland,
southern half of
Pangea. The Antarctic fossils were all identified as Procolophon
trigoniceps Owen, typically
also found in the Lystrosaurus Zone in
Another successful Early triassic reptile group were the archosaurs, diapsids 1-3 meters in length who were present in modest numbers in the latest Permian, then expanded considerably in the Early Triassic. Protorosaurus ("first lizard"), a lizard-like reptile about two meters long who had lived in the area of Germany during the Late Permian, is the earliest known archosauromorph.
of the most common archosaurs in the Early
Triassic, and one of the largest, was Proterosuchus,
associated with the Lystrosaurus faunal complex in
distinctive trait of P. fergosi
is its an expanded premaxilla with sharp, conical
teeth, which extends downward like a scoop, and effectively extends
reach of the upper jaw.
spent most of its time in rivers or
estuaries, but could also walk on land. At first it was considered to
have been a direct ancestor of modern crocodiles, but it is now
understood to be more basal (i.e., it is ancestral to a series of
archosaurs, that eventually led to crocodiles, dinosaurs, and
A Chinese variant of Proterosuchus is named Chasmatosaurus, which is also linked with the Lystrosaurus fauna in the Early Triassic Heshankou formation in Shanxi Province, in northeast China (Sun 1980). The Triassic sequences in Shanxi, Xianjiag, and Yunnan provinces will be described below.
Another related archosaur is Tasmaniasaurs triassicus, first discovered in 1960 in Hobart, Tasmania on the southeast coast of Australia, and recently identified as a probable member of the Proterosuchidae family, with similarities in its premaxilla and dentition to P. fergosi (Ezcurra 2014). Recovered were skeletal parts and a partial skull, which has provided a partial cranial endocast, the first for Proterosuchidae. This revealed structures related to the olfactory bulb, the brain section associated with the nasal passages and the sense of smell.
three zones of
exposed Triassic formations containing vertebrate
One of these regions
is in the south, located around the boundary of Yunnan and
Gouzhou provinces. These are
China has three zones of exposed Triassic formations containing vertebrate fossils (fig.5).
One of these regions is in the south, located around the boundary of Yunnan and Gouzhou provinces. These aredeposits of ancient marine or estuarine formations with an abundance of Triassic marine reptiles, fish, and invertebrates, but few terrestial animals (Benton et al 2013).
other two regions
are in the north, in the Ordos and Jungghar Basins. These
contain a variety of fossils of synapsids and
well as bony fish, with a much scarcer representation of labyrinthodont
amphibians. In the northeast, the provinces of Shanxi and Shaanxi
Yellow River, along whose valley many fossil outcrops are exposed
(fig.5). In northwestern China, significant Triassic exposures
occur in Xinjiang
(Young 1946; Sun 1980). Lucas
(1993) and Sun (1980) summarize the Triassic tetrapod fauna in northern
while Benton et al.
(2013) review findings from southern China.
The other two regions are in the north, in the Ordos and Jungghar Basins. These contain a variety of fossils of synapsids and reptiles, as well as bony fish, with a much scarcer representation of labyrinthodont amphibians. In the northeast, the provinces of Shanxi and Shaanxi straddle the Yellow River, along whose valley many fossil outcrops are exposed (fig.5). In northwestern China, significant Triassic exposures occur in
Xinjiang province (Young 1946; Sun 1980).Together, these two regions contain virtually all of the known land vertebrates or tetrapod fossils from the Triassic.
Lucas (1993) and Sun (1980) summarize the Triassic tetrapod fauna in northern China
, while Benton et al. (2013) review findings from southern China.
Fig.5: Locations of Chinese provinces with Triassic outcrops.
Mesozoic land vertebrate
fauna in China are in western Xinjiang,
in the Early
Formation. The faunal assemblage there at the Permian-Triassic boundary
is called the Jimsarian fauna (Yang et al. 1992; Spencer 1993). This is
equivalent to the Lystrosaurus fauna from South Africa, as discussed by
Sun (1980). Synapsids in the Jimsarian fauna include two
and the contemporary Dicynodon
; and the
("beast-headed") carnivore Urumchia
(Young 1952). Therocephalians
are advanced synapsids who survived from the Late Permian into the
Early Triassic (ca.
265-245 mya). These were numerous in South Africa, and
spread to Antarctica as well as China, Russia, and Eastern Europe.
the Jimsarian fauna include the procolophon Santisaurus
(Koh 1940); the proterosuchus Chasmatosaurus (Young
(Young 1973). All are typical Early Triassic fauna.
as 1991, no
fossils were known in China,
although plant fossils definitely existed from that period across
northern China (Lucas 1993).
As late as 1991, noLate Triassic
tetrapod fossils were known in China, although plant fossils definitely existed from that period across northern China (Lucas 1993).This apparent gap in the Triassic vertebrate fossil record was resolved in 1992, when the medium sized archosaur Yonghesuchus sangiensis was found in Upper Triassic strata in Shanxi province (Shao-Chun et al. 2001), as discussed below.
The equivalent Early Triassic faunal assemblage from Shanxi and Shaanxi provinces (fig.6) is called the Fugguan fauna, named for the town of Fugu which lies amid the Heshangou Formation (Lucas 1993; Sun 1980). Present are various reptilian taxa, including procolophonids (two species of Eumatabolodon); the protosucherian Xilosuchus, and Fugusuchus, a large Erythrosuchid ("red crocodile"), the largest archosaur predators in the Cygnognathus zone in South Africa. Also from the Fugguan fauna were a few unidentfied remains of labyrinthodont amphibians; and the lungfish Ceratodus ("horned tooth"), similar to examples found in the Bashunchank Formation in Russia.
Fig.6: Late Permian and Triassic vertebrate sites in Shanxi and Shaanxi provinces (after Lucas 1993, fig.3).
Synapsid remains found in Fugguan fauna in Shanxi included the large therocephalian Hazhenia, and smaller therocephalians named Scapulosaurus, also typical of the earliest Triassic of South Africa (Sun 1980; Huttenlocker 2014). Table 2 gives a brief comparison of Early Triassic animal types found in both Xinjiang and Shanxi/Shaanxi.
|Table 2: Comparative Early Triassic fauna in northern China|
|Period:||Early Triassic||Early Triassic|
|Fish: Ostyichthyes||Sarcoptygerian||Dipnoi (lungish)||Cerotodus|
Conspicuous by its absence in the Shanxi/Shaanxi fauna is the small synapsid dicynodont Lystrosaurus, otherwise found in Xinjiang, and throughout most of Pangea in the Early Triassic. This discrepancy was reviewed by Sun (1980), who provides a useful summary of the distribution of Lystrosaurus-related fauna in Pangea. Differences in the faunal assemblages between Shanxi and Xinjiang fossil deposits may be linked to geological processes such as variations in micro-plate formation and the physical linkage of different regions of Pangea, something which has been documented for these provinces (Xiong and Coney 1985). These geological processes may have caused differing environments, as well as relative isolation of some areas.Fig.7: Skull of Parakannemeyeria: A) lateral view, showing position of tusks in maxilla; B) basal view, indicating stapes (after Sun 1960, figs. 2 and 4).
Fossil vertebrates from this
component, called the Ordonian fauna by Lucas (1993), consists
large dicynodonts. Among these is Parakannemeyeria
found at the top of the section , together with a variety of Kannemeyeria. The
latter animal was widespread
Pangea, found in South Africa, Tanzania, Zambia,
Argentina, India, China, and Russia, always in strata at the boundary
of the Early and Middle Triassic (Lucas 1993). Kannemeyeria's
diagnostic role in this
time period (ca. 247-240 mya) is
reflected in its use
as type species for the Middle Triassic fauna in Xingiangi
an long, narrow, and highly projected
crest, as well as tusks of larger dimensions in the
related Parakannemeyeria (Sun
The cynodont Ordosiodon is also represented in the Lower Ermaying Fauna, which is equated to the the Cynognathus Zone of South Africa.
Reptiles from the Lower Ermaying included a moderate sized procolophonid similar to Neoprocolophon, a later form which is larger than Procolophon.
Fig.8: Skull of Shansiodon wuhsiangensis, showing large tusks that comprise the two teeth of this dicynodont (after Yuh 1961).
Upper Ermaying Formation:
Fauna from this formation, named after the city of Ningwuan on the
Sanggan River in the Ordos Valley,
to the Sinokannemeyeria
fauna originally defined by C.C. Young. Young discovered the dicynodont synapsid Sinokannemeyeria
first identification of a Triassic tetrapod in China
Subsequent work in the Ordos basin took place in the 1950s and 60s
Dicynodonts, representing therapsids who were non-mammalian synapsids, are the most numerous taxa in these late Early Triassic levels, consisting of the three genera Kannemeyeria, Parakannemeyeria, and Shansiodon. The former two are large individuals, while the latter, represented by three different species, is small. One of the Shansiodon species, S. wuhsiangensis, shows the large twin tusks characteristic of the genus (fig.8) . Another, S. wagni (Yuh 1959), showed good preservation of the stapes, a hearing ossicle attached to the quadrate bone in the occipital region at the lower rear of the skull (fig.9).
Fig.9: The skull of Shansiodon wagni viewed from behind, showing the stapes, a small tube-shaped bone used as an ossicle in hearing (after Yuh 1959).
Two Cynodonts (a Therapsid group ancestral to mammals) were also recovered from the Upper Ermaying deposits. One was diagnosed as Sinognathus (fig.10), and another identified as Traversodontoides , a bauriidae therapsid found most frequently in the Middle Triassic of South America (Young 1959, l974; Sun 1980; Lucas 1993.
Sinognathus belongs to the family Cynognathidae and is regarded as similar to Cynognathus from South Africa. Sun (1980), however, notes that, while the South African cynodont, Cynognathus is a relatively large individual with a skull that closely resembles a dog, displaying long and narrow post canine teeth, Sinognathus appears dissimilar. It has an more complex maxilla or upper jaw area, and more closely resembles Traversodon or Belesodon.
Fig.10: The skull of Sinognathus gracilis, showing its maxilla or upper jaw with a surmaxilla (after Young 1959 fig.3)
Fossils representing the primitive thecodont reptile, Shansisuchus are also abundant, as exemplified by massive fossiliferous block with the vast majority of elements belonging to this genus (Sun 1980). Other thecodont taxa include Fenhosuchus and Wangisuchus. A single skull of the small, temporally diagnostic reptile Procolophonia is also represented in the Upper Ermaying fauna, in the form of Neoprocolophon.
Kelemayi Formation in Xinjiang:
The Middle Triassic Kelamayi Formation in Xinjiang contains the Kannemeyeria Fauna (Lucas 1993), which correlates with the entire upper section of the Ermaying Formation exposed along the Yellow River in Shanxi and Shanaxi provinces (Table 3). In more distant regions, according to Sun (1980), the Kelamayi Formation may also be appropriately correlated to equivalent fauna in the Indian Yerrapalli and Dongus System in Russia.
|Table 3: Comparative Early and Middle Triassic fauna in northern China|
|Period:||Middle Triassic||Late Early Triassic||Middle Triassic|
|Formation:||Upper Ermaying||Lower Ermaying||Kelemayi|
||Parakannemeyeria (3 species)||Parakannemeyeria||Parakannemeyeria|
|Synapsid||Therapsid||Dicynodont||Sinokannemayeria (2 species)||Kannemayeria|
|Synapsid||Therapsid||Dicynodont||Shansiodon (3 species)|
|Synapsid||Therapsid||Therocephalian||Ordosiodon (3 species)|
|Reptile||Diapsid||Erythysuchian||Shansisuchus (2 species)||Vjushkovia|
A Late Triassic Archosaur from Shanxi Province
In 1992 came the first
record of Late Triassic tetrapod from
fossils, embedded in a
sandstone block, were
found on the
north side of Sangbi
Creek, about 1.5 km southwest of
Fig.11: Reconstructed view of the skull of Yonghesuchus sangiensis, with bones labelled (after Shao-Chun et al. 2001, fig.4).
On the basis of stratigraphy, plus associated fossil fish taxa from the Late Triassic found in the same strata as Yonghesuchus, and some aspects of the more derived traits of Yonghesuchus compared to Middle Triassic archosaurs from China, Chao-Chun et al (2001) identified the fossils as dating from the earliest part of the Late Triassic period. This find thus represents the uppermost stratigraphic record of a tetrapod in the China Triassic geological formations.
Regarding its phylogeny, Y. sangiensis (Anapsida, Archosauromorpha, Archosauriformes) represents a new species of archosaur with some similarities to Turfanosuchus, found in the late Early Triassic, Ermaying formation in Shanxi province, and the Middle Triassic, Kelemayi Formation in Xianjing province (Table 2). Y. sangiensis, however, is considerered more derived than both Turfanosuchus, and the early Late Triassic archosaur Proterochampsa.
Triassic marine reptiles in southern China
zone of fossil-rich
formations spanning the Permian-Triassic
in the adjacent Yunnan and
Ghozou provinces in southern
The South China Basin occupies a section of ancient lithosphere known as the South China Block. At the beginning of the Triassic, South China was an island in the Tethys Sea, located just southwest of northern China, at the eastern end of Pangaea (fig.12).
Chinese paleontologists including C.C. Young began intensive work in the Triassic deposits of South China in the 1930s to 70s. Within the past 30 years, several fossil-rich areas have been studied more intensively, with relatively precise chronology now obtained from the strata (Young 1946; Yin et al., 2001; Shen et al., 2011; Benton et al. 2013).
Fig.12: Location of South China in the Tethys sea in the Triassic (after Benton et al. 2013, fig.2A)
Fig.13: Map of Triassic fossil deposits in southern China, showing Luoping, Panxian, and Guanling biotas (Benton et al. 2013, fig.2B)
from the Luoping biota
include medium-sized (1-5 m -long) sea reptiles such as Mixosaurus,
Triassic vertebrate fossils in India
India contains regions with a continuous succession of Early, Middle, and Late Triassic sediments, containing well preserved terrestial faunas/ In
Temnospondyl amphibians in the Panchet Formation include the trematosaurid Gonioglyptus, a widespread taxa found from Antarctica to Russia. Very similar trematosaurids are in the Prionolobus beds of the Salt Range, of Dienarian age. Four other temnospondyls are known. These include Indobrachiops, of the Indobrachyopid group, which is restricted to India and Australia. Also found are Lydekkerina, the type genus of the widespread Lyderkkerinid group, found from Antarctica to Russia; Pachygonia of the Captitosaurids, another widespread group found from Antarctica to Russia; and Indobenthosuchus, which appears related to Lydekkerina.
Synapsids from the Panchet
Formation include the widespread dicynodont Lystrosaurus,
Synapsids from the Panchet Formation include the widespread dicynodont Lystrosaurus, representingtetrapods of lowland habitats living in fluvial and lacustrine settings (Bandyopadhyay 1999, p.294). Also found is the cynodont Thrinaxodon, associated with the Lystrosaurus Zone in South Africa, representing the initial phase of the Early Triassic. Table 4 lists Early and Middle Triassic vertebrate fauna from three regions of India.
|Table 4: India: Early and Middle Triassic fauna|
|Region:||Damodar Valley||Pranhita-Godovari Basin||Satpura Basin|
|Period:||Early Triassic||Mid Triassic- Mid Anisian||Mid Triassic - Early Anisian|
most common taxa
|Synapsid||Therapsid||Dicynodont : stahleckeriid||Rechnisaurus|
|Synapsid||Therapsid||Dicynodon||unid. genus- small|
|Synapsid||Therapsid||Dicynodon||unid. genus- large|
|Reptile||Diapsid||archosaur||unid. genus- small|
|Amphibian||Temnospondyl||Captitosaurid||Pachygonia||Parotosuchus||Parotosuchus (2 species)|
|Amphibian||Temnospondyl||from Mangli beds of
represets the upper Kamthi formation} Brachyopid
|Brachiops laticeps||unid. genus|
A herd of Triassic
Wadiasaurus indicus, a small Middle Triassic dicynodont herbivore about 50 cm long, has been identifed as a kannemeyeriid, so far the only example of this family of therapsid grazers known in India. The Wadiasaurus type specimen is a skull found in an Anisian (247-242 mya) fluvial claystone deposit in the Yerrapalli Formation of the Pranhita- Godavari valley, about 2 km east-south-east of the town of Yerrapalli. The genus is named for the Indian geologist Darashaw Nosherwan Wadia (1883-1969).
More recent findings of a larger sample of Wadiasaurus fossils, in a locality near that of the original type skull, indicate that some modification of the skull characters given in previous descriptions is necessary. Based on additional information provided by the new samples on the lower jaw and postcranial skeleton, meanwhile, the familial status of W. indicusas as a Kannemeyerid has been confirmed (Bandyopadhyay 1988). A comparative study of Wadiasaurus and other kannemeyeriid genera indicates that it might have been most closely related to Kannemeyeria erithrea. In Wadiasaurus, the parietal crest at the top of the skull is quite specialized, forming two long rounded lobe-like bars separated by a bay. Sexual dimorphism of the species is indicated by maxillary flanges, which in males are triangular, thick and swollen, with stout cylindrical tusks, while similar maxillary flanges of females lack tusks.
Interesting behavioral traits have been revealed by a taphonomic study of the bone assemblage found in the fluvial claystones near Yarapalli. These findings show that a herd of Wadiasaurus, including some juveniles and young animals, was trapped in the soft muds of a floodplain and buried in a small area. The herd wa composed only of females, with some juvenile members. Taphonomic and osteological studies tend to indicate that the female individuals of Wadiasaurus lived in herds, whereas the solitary males joined the herds only during the mating seasons (Bandyopadhyay 1999).
Fig.14: Skull of Rechnisaurus
A second dicynodont from the Pranhita-Gondavari valley is Reichnisaurus (fig.14), with similarities to a South American taxa named Dinodontosaurus, from the Stahlekriid family (Colbert 1984; Bandyopadhyay 1988; Jain 1996).
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