Sanajeh Indicus
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''Sanajeh'' (meaning "ancient gape" in
Sanskrit Sanskrit (; attributively , ; nominally , , ) is a classical language belonging to the Indo-Aryan branch of the Indo-European languages. It arose in South Asia after its predecessor languages had diffused there from the northwest in the late ...
) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
madtsoiid
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
from western
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. A fossil described in
2010 File:2010 Events Collage New.png, From top left, clockwise: The 2010 Chile earthquake was one of the strongest recorded in history; The Eruption of Eyjafjallajökull in Iceland disrupts air travel in Europe; A scene from the opening ceremony of ...
from the
Lameta Formation The Lameta Formation, also known as the Infratrappean Beds, is a sedimentary geological formation found in Madhya Pradesh, Gujarat, and Maharashtra, India, associated with the Deccan Traps. It is of Maastrichtian age (Late Cretaceous), and is nota ...
was found coiled around an egg and an adjacent skeleton of a 50 cm (19 in) long
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
hatchling. This suggests that the snake preyed on hatchling sauropods at nesting sites.


Description

The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
specimen, known as GSI/GC/2901–2906, consists of a nearly complete skull and lower jaws, and 72 precloacal vertebrae and ribs preserved in five articulated sections. It was found from
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
-age
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an adje ...
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
s outcropping in the village of Dholi Dungri in
Gujarat Gujarat (, ) is a state along the western coast of India. Its coastline of about is the longest in the country, most of which lies on the Kathiawar peninsula. Gujarat is the fifth-largest Indian state by area, covering some ; and the ninth ...
. ''Sanajeh'' was around in length based on the length of the skull, which is . On the side of the skull there is an opening called the juxtastapedial recess, which is characteristically rectangular. The juxtastapedial recess would have contained
cranial nerves Cranial nerves are the nerves that emerge directly from the brain (including the brainstem), of which there are conventionally considered twelve pairs. Cranial nerves relay information between the brain and parts of the body, primarily to and f ...
associated with the ear, while another opening located in front of the recess, the trigeminal foramen, housed cranial nerves associated with the jaws. The jaw joint of ''Sanajeh'' is located to the side of the posterior margin of the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
, which is characteristic of basal snakes. A
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
runs along the ventral surface of the braincase and served as an attachment for ''protractor pterygoidei'' muscles that moved the toothed bones of the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
. Articulations between the vertebrae are well developed in ''Sanajeh'' in a similar way to other madtsoiids and the genus ''
Najash ''Najash'' is an extinct genus of basal snake from the Late Cretaceous Candeleros Formation of Patagonia. Like a number of other Cretaceous and living snakes it retained hindlimbs, but ''Najash'' is unusual in having well-developed legs that e ...
''. The
neural spine The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
s are thin and angled posteriorly. In ''Sanajeh'', the synapophyses, or rib articulations, extend outward past the margins of the
prezygapophyses The articular processes or zygapophyses (Greek language, Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "-physis, process") of a vertebra are projections of the vertebra that serve the purpose of fitting ...
. This is a characteristic of all madtsoiids.


Classification

''Sanajeh'' is a member of the Madtsoiidae, an extinct
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of snakes that includes the
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
n genera ''
Wonambi ''Wonambi'' is an extinct genus of madtsoiid snakes that lived in late Neogene to late Quaternary Australia. Species of ''Wonambi'' were constrictor snakes unrelated to Australian pythons. Description ''Wonambi'' was a fairly large snake, ...
'' and ''
Yurlunggur ''Yurlunggur'' is a genus of fossil snake in the extinct family Madtsoiidae. This genus was a part of the extinct megafauna of Australia, and contains the species ''Yurlunggur camfieldensis''. ''Y. camfieldensis'' was a large apex predator t ...
''. Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
showing the phylogenetic relations of ''Sanajeh'': In 2022, a morphological study found Madtsoiidae as previously defined to be paraphyletic, with ''Sanajeh'' being the most basal member of the
Ophidia __FORCETOC__ Ophidia (also known as Pan-Serpentes) is a group of squamate reptiles including modern snakes and reptiles more closely related to snakes than to other living groups of lizards. Ophidia was defined as the "most recent common ancesto ...
, while the Australian "madtsoiids" from the Cenozoic are basal alethinophidians.


Paleobiology


Feeding

Like many early snakes, ''Sanajeh'' did not have the wide gape seen in
boids Boids is an artificial life program, developed by Craig Reynolds in 1986, which simulates the flocking behaviour of birds. His paper on this topic was published in 1987 in the proceedings of the ACM SIGGRAPH conference. The name "boid" corr ...
,
pythons The Pythonidae, commonly known as pythons, are a family of nonvenomous snakes found in Africa, Asia, and Australia. Among its members are some of the largest snakes in the world. Ten genera and 42 species are currently recognized. Distribution ...
, and caenophidians. Therefore, it could not consume prey as large as that which many modern snakes can. Living snakes that have narrow gapes, including uropeltids, ''
Anomochilus The Anomochilidae, or anomochilids, are a monotypic family of snakes, created for the genus ''Anomochilus'', which currently contains three species. It is commonly called the dwarf pipe snake. Description Anomochilids are small snakes, with mus ...
'', ''
Cylindrophis The Cylindrophiidae are a monotypic family of secretive, semifossorial, non-venomous snakes containing the genus ''Cylindrophis'' found in southeastern Asia. These are burrowing snakes and most have a banded pattern on the belly. Currently, thir ...
'', and ''
Anilius The Aniliidae are a monotypic family created for the monotypic genus ''Anilius'' that contains the single species ''A. scytale''. Common names include American pipe snake and false coral snake. It is found in South America. This snake posses ...
'', have diets that are limited to smaller animals such as
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s,
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
larvae,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s, and
amphisbaenian Amphisbaenia (called amphisbaenians or worm lizards) is a group of usually legless squamates, comprising over 200 extant species. Amphisbaenians are characterized by their long bodies, the reduction or loss of the limbs, and rudimentary eyes. As ...
s and
caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
s. The short
supratemporal The supratemporal bone is a paired cranial bone present in many tetrapods and tetrapodomorph fish. It is part of the temporal region (the portion of the skull roof behind the eyes), usually lying medial (inwards) relative to the squamosal and latera ...
and broad, short
quadrate Quadrate may refer to: * Quadrate bone * Quadrate (heraldry) * Quadrate lobe of liver * Quadrate tubercle The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...
indicate that the oral gape of ''Sanajeh'' was narrow. The gape is thought to have been similar to that of the extant genus ''
Xenopeltis ''Xenopeltis'', the sunbeam snakes, are the sole genus of the monotypic family Xenopeltidae, the species of which are found in Southeast Asia. Sunbeam snakes are known for their highly iridescent scales. Two species are recognized, each one wit ...
''. However, the presence of strong ''m. protractor pterygoidei'' muscles inferred from the sagittal crest of ''Sanajeh'' indicates that it was able to manipulate prey in its mouth like modern macrostomatans. The intramandibular joint was able to flex greatly, which would allow for the consumption of larger prey. The presence of these features in ''Sanajeh'' shows that increased oral kinesis (movement of the mouth) and intraoral mobility (the ability to move the bones of the palate) preceded the development of wide gapes in snakes. Therefore, reduced cranial kinesis in basal living snakes may be a
fossorial A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees. Prehistoric eviden ...
adaptation rather than the retention of a
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
trait. The holotype of ''Sanajeh'' was found in association with sauropod eggs belonging to the
oospecies Egg fossils are the fossilized remains of eggs laid by ancient animals. As evidence of the physiological processes of an animal, egg fossils are considered a type of trace fossil. Under rare circumstances a fossil egg may preserve the remains of t ...
'' Megaloolithus dhoridungriensis'' and one incompletely preserved sauropod hatchling, likely a
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still th ...
. The snake was coiled around a crushed egg, which the hatchling may have exited from. The eggs were laid in a nest structure that was not preserved, but was likely covered in loose sediment or vegetation based on the porosity of the eggs. The rigid eggs were probably too large for ''Sanajeh'' to consume, but the snake would have been able to break an egg open and consume its contents in a way similar to the living ''
Loxocemus :''Common names: Mexican python, Mexican burrowing python, Mexican burrowing snake.'' ''Loxocemus bicolor'',the sole member of the monotypic family Loxocemidae, is a species of python-like snake found in Mexico and Central America. No subspecies ...
''. ''Sanajeh'' is likely to have had a nest-plundering feeding strategy, and it is possible that the snake consumed a large variety of prey items including the eggs of
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s and smaller reptiles, which are common in the Lameta Formation. Accelerated growth rates and large numbers of hatchlings would have enabled titanosaurs such as ''
Isisaurus ''Isisaurus'' (named after the Indian Statistical Institute) is a genus of titanosaurian dinosaur from the Late Cretaceous Lameta Formation of India. The genus contains a single species, ''Isisaurus colberti''. Discovery and Naming The type spe ...
'' and ''
Jainosaurus ''Jainosaurus'' is a genus of titanosaurian sauropod dinosaur of India and wider Asia, which lived in the Maastrichtian (approximately 68 mya (unit), million years ago). No accurate estimate of the length, height, or weight has yet been mad ...
'' to overcome the losses of predation by animals such as ''Sanajeh''. Titanosaurs would have been free of the risk of predation by their first year as a result of their rapid growth rate. The size of adult titanosaurs, which could be in length, would have deterred nearly all predators.


References

{{Taxonbar, from=Q135632 Fossils of India Cretaceous snakes Fossil taxa described in 2010