Madtsoiidae are an extinct group of mostly
GondwanaGondwana , originally Gondwanaland, is the name given to a southern precursor-supercontinent. Its final joining occurred between ca. 570 and 510 Ma ago, joining East Gondwana to West Gondwana. It later separated from Laurasia 180- during the breakup of the Pangaea supercontinent that existed about...
n snakes with a fossil record extending from early
CenomanianThe Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...
(Upper Cretaceous) to late
PleistoceneThe Pleistocene is the epoch from 2.588 million to 12 000 years BP covering the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
strataIn geology and related fields, a stratum is a layer of rock or soil with internally consistent characteristics that distinguishes it from contiguous layers. Each layer is generally one of a number of parallel layers that lie one upon another, laid down by natural forces. They may extend over...
located in
South AmericaSouth America is the southern continent of the Americas, situated entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere...
,
AfricaAfrica is the world's second-largest and second most-populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area. With a billion people in 61 territories, it accounts for about 14.8% of the...
,
IndiaIndia, officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the south, the Arabian Sea on the west, and the Bay of Bengal...
,
AustraliaAustralia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the continental mainland , the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans...
and Southern
EuropeEurope is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally divided from Asia to its east by the water divide of the Ural Mountains, the Ural River, the Caspian Sea, the Caucasus Mountains , and the Black Sea to the southeast...
.
Madtsoiid snakes include
very primitiveIn phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...
snakes, which like extant boas and pythons would likely dispatch their prey by
constriction-Snakes:Constriction is a method used by various snake species to kill their prey. Although some species of venomous and mildly-venomous snakes do use constriction to subdue their prey, most snakes which use constriction lack venom. The snake initially strikes at its prey and holds on, pulling the...
, such as
Gigantophis, one of the longest snakes known at an estimated 10.7 meters, and the
Australian Aboriginal mythologyAustralian Aboriginal myths are the stories traditionally performed by Aboriginal peoples within each of the language groups across Australia....
-named
Wonambi and
Yurlunggur.
As a grouping of basal forms the composition and even the validity of Madtsoiidae is in a state of flux as new pertinent finds are described.
Madtsoiidae was first classified as a subfamily of Boidae, Madtsoiinae, in Hoffstetter (1961a).
Madtsoiidae are an extinct group of mostly
GondwanaGondwana , originally Gondwanaland, is the name given to a southern precursor-supercontinent. Its final joining occurred between ca. 570 and 510 Ma ago, joining East Gondwana to West Gondwana. It later separated from Laurasia 180- during the breakup of the Pangaea supercontinent that existed about...
n snakes with a fossil record extending from early
CenomanianThe Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...
(Upper Cretaceous) to late
PleistoceneThe Pleistocene is the epoch from 2.588 million to 12 000 years BP covering the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
strataIn geology and related fields, a stratum is a layer of rock or soil with internally consistent characteristics that distinguishes it from contiguous layers. Each layer is generally one of a number of parallel layers that lie one upon another, laid down by natural forces. They may extend over...
located in
South AmericaSouth America is the southern continent of the Americas, situated entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere...
,
AfricaAfrica is the world's second-largest and second most-populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area. With a billion people in 61 territories, it accounts for about 14.8% of the...
,
IndiaIndia, officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the south, the Arabian Sea on the west, and the Bay of Bengal...
,
AustraliaAustralia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the continental mainland , the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans...
and Southern
EuropeEurope is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally divided from Asia to its east by the water divide of the Ural Mountains, the Ural River, the Caspian Sea, the Caucasus Mountains , and the Black Sea to the southeast...
.
Madtsoiid snakes include
very primitiveIn phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...
snakes, which like extant boas and pythons would likely dispatch their prey by
constriction-Snakes:Constriction is a method used by various snake species to kill their prey. Although some species of venomous and mildly-venomous snakes do use constriction to subdue their prey, most snakes which use constriction lack venom. The snake initially strikes at its prey and holds on, pulling the...
, such as
Gigantophis, one of the longest snakes known at an estimated 10.7 meters, and the
Australian Aboriginal mythologyAustralian Aboriginal myths are the stories traditionally performed by Aboriginal peoples within each of the language groups across Australia....
-named
Wonambi and
Yurlunggur.
As a grouping of basal forms the composition and even the validity of Madtsoiidae is in a state of flux as new pertinent finds are described.
Description
Madtsoiidae was first classified as a subfamily of Boidae, Madtsoiinae, in Hoffstetter (1961a). Further study and new finds allowed ranking the group as a distinct family in
Linnaean systemsLinnaean taxonomy is either# the particular classification of Linnaeus, as set forth in his Systema Naturae and subsequent works...
. With the recent use of
cladisticsCladistics is a form of biological systematics which classifies living organisms on the basis of shared ancestry...
to unravel phylogeny, various analyses have posited Madtsoiidae as a likely clade within Serpentes, or possible paraphyletic stem group outside Serpentes and within a more inclusive Ophidia.
Madtsoiid snakes ranged in size from less than 1 m (estimated total length) to over 9 m, and are thought to have been constrictors analogous to modern pythons and boas, but with more primitive
jawThe jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term jaws is also broadly applied to the whole of the structures constituting the vault of the mouth and serving to open and close it and is part of the body plan of...
structures less highly adapted for swallowing large prey. There are specific anatomical features that diagnose members of this family, such as the presence of hypapophyses only in anterior trunk, that the middle and posterior trunk vertebrae possess a moderately or well developed haemal keel, except for a few near the cloacal region, often with short laterally paired projections on the posterior part of the keel. Also, all trunk and caudal vertebrae have at least a parazygantral foramen, sometimes several of them, located in a more or less distinct fossa that is lateral to each zygantral facet. Addition features are the prezygapophyseal processes' absence while the paracotylar foramina are present and that the diapophyses are relatively wide, exceeding width across prezygapophyses at least in the posterior trunk vertebrae. (Scanlon 2005)
Like most fossil snakes the majority of madtsoiids are known only from isolated
vertebraA vertebra is an individual bone in the flexible column that defines vertebrate animals, e.g. humans. The vertebral column encases and protects the spinal cord, which runs from the base of the cranium down the dorsal side of the animal until reaching the pelvis. From there, vertebra continue into...
e, but several (
Madtsoia bai,
M. camposi,
Wonambi naracoortensis,
Nanowana spp., unnamed
Yurlunggur spp.,
Najash rionegrina) have associated or articulated parts of skeletons. Of the genera listed below, all have been referred to Madtsoiidae in all recent classifications except
Najash rionegrina, which is included here based on diagnostic vertebral characters described by Apesteguía and Zaher (2006). These authors didn't include Najash among madtsoiids because they consider that madtsoiids are a paraphyletic assemblage of basal macrostomatans related to Madtsoia bai and consequently, not related to the Cretaceous alethinophidians from southern continents.
Rieppel et al. (2002) classified
Wonambi naracoortensis within the extant radiation, (
crown groupA crown group is the smallest monophyletic group, or "clade", to contain the last common ancestor of all members, and all of that ancestor's descendants...
), of snakes as Macrostomata
incertae sedis, but many of their character state attributions for this species have been criticised or refuted by Scanlon (2005a) and the better-preserved
skullThe skull is a bony structure found in the head of many animals. The skull supports the structures of the face and protects the head against injury....
s of
Yurlunggur sp./spp. have numerous characters apparently more plesiomorphic than any macrostomatans (Scanlon 2006). The partial skull attributed to
Najash rionegrina (Apesteguía and Zaher 2006) resembles that of the non-madtsoiid
DinilysiaDinilysia is an extinct genus of snake from the Late Cretaceous of South America.-In Popular Culture:The BBC website confirms Dinilysia as appearing in the last episode of the famous 1999 documentary series Walking with Dinosaurs as the snake encountered by two infant Tyrannosaurus, although the...
patagonica, and vertebrae support that they are related. The type material of
Najash is the only possible madtsoiid specimen retaining evidence of pelvic and hindlimb elements, which are claimed to be more plesiomorphic than other Cretaceous limbed snakes, such as
PachyrhachisPachyrhachis is an extinct genus of snake with well developed hind legs known from fossils discovered Ein Yabrud, near Ramallah, in the central West Bank. It is one of three genera of Cenomanian snakes with hindlimbs. Many of today's modern pythons and boas still have small bone spurs, where limbs...
,
HaasiophisHaasiophis, consisting of the sole species Haasiophis terrasanctus, is an extinct genus of snakes with hind limbs. It is one of three genera of Cenomanian snakes known to have possessed hindlimbs.-Etymology:...
or
EupodophisEupodophis is an extinct genus of snake from the Late Cretaceous period. The only species , E. descouensi, had two legs, and is a "transitional form" between Cretaceous lizards and limbless snake....
, in retaining a sacro-iliac contact and well developed limbs, with a huge and well defined trochanter. The sacro iliac contact is perhaps misleadingly described by Apesteguía and Zaher as unique possession of a
sacrumThe sacrum is a large, triangular bone at the base of the spine and at the upper and back part of the pelvic cavity, where it is inserted like a wedge between the two hip bones. Its upper part connects with the last lumbar vertebra, and bottom part with the coccyx...
, whereas it has rarely been questioned that the
cloacaIn zoological anatomy, a cloaca is the posterior opening that serves as the only such opening for the intestinal, reproductive and urinary tracts of certain animal species. The word comes from Latin, and means sewer...
l vertebrae in snakes are
homologousIn evolutionary biology, homology refers to any similarity between characteristics of organisms that is due to their shared ancestry. The word homologous derives from the ancient Greek ομολογειν, 'to agree'. There are examples in different branches of biology...
to the sacrals of limbed squamates (i.e. the sacrum is present but has lost contact with the reduced ilia in other taxa). It would be unsurprising if other madtsoiids also possessed hindlimbs as complete as those of
Najash.
Several madtsoiid genera have been named using indigenous words for legendary
Rainbow SerpentThe Rainbow Serpent is a common motif in the art & mythology of Aboriginal Australia.The Rainbow Serpent is seen as the inhabitant of permanent waterholes and is in control of life's most precious resource, water. He is the underlying Aboriginal mythology for the famous Outback "bunyip"...
s or
dragonDragons are legendary creatures, typically with serpentine or otherwise reptilian traits, that feature in the myths of worldwide cultures.The two most familiar interpretations of dragons are European dragons, derived from various European folk traditions, and the unrelated Oriental dragons, such as...
s, including
Wonambi (Pitjantjatjara),
Yurlunggur (
YolnguThe Yolngu are an Indigenous Australian people inhabiting north-eastern Arnhem Land in the Northern Territory of Australia. Yolngu literally means “person” in the language spoken by the people.-Yolŋu Law:...
) and
Nanowana (
Ancient GreekAncient Greek is the historical stage in the development of the Greek language spanning across the Archaic , Classical , and Hellenistic periods of ancient Greece and the ancient world. It is predated in the 2nd millennium BC by Mycenaean Greek...
nano-, 'dwarf' +
WarlpiriThe Warlpiri language is spoken by about 3000 of the Warlpiri people in Australia's Northern Territory. It is one of the Ngarrkic languages of the large Southwest branch of the Pama-Nyungan family, and is one of the largest aboriginal languages in Australia in terms of number of speakers.-...
WanaWana is the largest town of South Waziristan Agency in Pakistan's FATA . It is the summer headquarters for the Agency's administration, Tank located in neighbouring Tank District being the winter HQ...
) in Australia, and
Herensugea (
BasqueBasque is the ancestral language of the Basque people, who inhabit the Basque Country, a region spanning an area in northeastern Spain and southwestern France. It is the mother tongue of approximately one fifth of Basques, 632,000 out of nearly 3,000,000...
) in Europe. G.G. Simpson (1933) apparently started this trend by compounding
Madtsoia from indigenous roots. In this particular case these originated from the
Tehuelche languageTehuelche is a nearly extinct Chon language spoken by four people in Patagonia out of an ethnic group of 200. They were originally nomadic hunters from Chile. It is also known as Aonikenk or Aonek'o 'ajen. The decline of the language started with the Araucanization of Patagonia, when many Tehuelche...
, although the reference made was geographic rather than mythological, the derivation being from that language's terms
mad, "valley" and
tsoi, "cow" as a rough translation from
SpanishSpanish or Castilian is a Romance language in the Ibero-Romance group that originated in northern Spain and gradually spread in the Kingdom of Castile, evolving into the principal language of government and trade in the Iberian peninsula...
name of the type locality, Cañadón Vaca.
Classification
- Gigantophis Andrews, 1901
- Gigantophis garstini
Gigantophis garstini was a prehistoric snake which may have measured more than 10 meters , larger than any living species of snake. It once took the mantle of largest snake before Titanoboa, which was recently discovered in Colombia...
Andrews, 1901 (Andrews 1906, Hoffstetter 1961b; PaleogeneThe Paleogene is a geologic period and system that began 65.5 ± 0.3 and ended 23.03 ± 0.05 million years ago and comprises the first part of the Cenozoic era...
, Late EoceneThe Eocene epoch, lasting from 55.8 ± 0.2 to 33.9 ± 0.1 Ma , is a major division of the geologic timescale and the second epoch of the Palaeogene period in the Cenozoic era. The Eocene spans the time from the end of the Paleocene epoch to the beginning of the Oligocene epoch. The start of the...
; EgyptEgypt , officially the Arab Republic of Egypt, is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Western Asia...
, LibyaLibya , officially the Great Socialist People's Libyan Arab Jamahiriya , is a country located in North Africa...
)
- Madtsoia
Madtsoia is an extinct genus of madtsoiid snake. It is known from the Eocene of Argentina , the Paleocene of Brazil , the Late Cretaceous of Spain , and the Late Cretaceous of Madagascar and possibly Niger Madtsoia is an extinct genus of madtsoiid snake. It is known from the Eocene of...
Simpson, 1933
- Madtsoia bai Simpson, 1933 (Paleogene, Early Eocene; Argentina
Argentina, officially the Argentine Republic , is the second largest country in South America, constituted as a federation of 23 provinces and an autonomous city, Buenos Aires. It is the eighth largest country in the world by land area and the largest among Spanish-speaking nations, though Mexico,...
)
- Madtsoia cf. M. bai (Simpson 1935, Hoffstetter 1960; Paleogene, Late Paleocene
The Paleocene or Palaeocene, the "early recent", is a geologic epoch that lasted from 65.5 ± 0.3 Ma to 55.8 ± 0.2 Ma . It is the first epoch of the Palaeogene Period in the modern Cenozoic era...
; Argentina)
- Madtsoia madagascariensis Hoffstetter, 1961a (Piveteau 1933; Cretaceous
The Cretaceous , Latin language for "chalky", usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago . In the geologic timescale, the Cretaceous follows on the Jurassic period and is followed by the Paleogene period of the...
, SantonianThe Santonian is an age in the geologic timescale or achronostratigraphic stage. It is a subdivision of the Late Cretaceous epoch or Upper Cretaceous series. It spans the time between 85.8 ± 0.7 mya and 83.5 ± 0.7 mya...
or CampanianThe Campanian is, in the ICS' geologic timescale, the fifth of six ages of the Late Cretaceous epoch . The Campanian spans the time from 83.5 ± 0.7 Ma to 70.6 ± 0.6 Ma ...
; MadagascarMadagascar, or Republic of Madagascar , is an island nation in the Indian Ocean off the southeastern coast of Africa. The main island, also called Madagascar, is the fourth-largest island in the world, and is home to 5% of the world's plant and animal species, of which more than 80% are endemic to...
)
- Madtsoia aff. madagascariensis (de Broin et al. 1974; Cretaceous, Coniacian
The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous epoch or Upper Cretaceous series and spans the time between 89.3 ± 1 Ma and 85.8 ± 0.7 Ma...
or Santonian, NigerNiger , officially the Republic of Niger is a landlocked country in Western Africa, named after the Niger River. It borders Nigeria and Benin to the south, Burkina Faso and Mali to the west, Algeria and Libya to the north and Chad to the east...
)
- Madtsoia laurasiae Rage, 1996 (Astibia et al. 1990; Cretaceous, Campanian or Maastrichtian
The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...
; Spain)
- Madtsoia camposi Rage, 1998 (Paleogene, middle Paleocene; Brazil
Brazil , officially the Federative Republic of Brazil , is a country in South America. It is the fifth largest country by geographical area, occupying nearly half of South America, the fifth most populous country, and the fourth most populous democracy in the world. Bounded by the Atlantic Ocean...
)
- Wonambi
Wonambi is a genus currently consisting of two species of very large snakes. These species are not pythons, like Australia's other large constrictors of the genus Morelia, but a member of a now extinct family Madtsoiidae...
Smith, 1976
- Wonambi naracoortensis Smith, 1976 (Scanlon and Lee 2000, Scanlon 2005; Neogene, Pliocene
The Pliocene epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present....
to Pleistocene; Australia)
- Wonambi barriei Scanlon in Scanlon and Lee, 2000 (Neogene
The Neogene is a geologic period and system starting 23.03 ± 0.05 million years ago and lasting either until today or ending 2.588 million years ago with the beginning of the Quaternary. The Neogene Period follows the Paleogene Period of the Cenozoic Era...
, early MioceneThe Miocene is a geological epoch of the Neogene period and extends from about 23.03 to 5.33 million years before the present . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the...
; Australia)
- Patagoniophis Albino, 1986
- Patagoniophis parvus Albino, 1986 (Cretaceous, Campanian or Maastrichtian; Argentina)
- Patagoniophis australiensis Scanlon, 2005 (Scanlon 1993; Paleogene, early Eocene; Australia)
- Alamitophis
Alamitophis is a genus of fossil snake in the extinct family of Madtsoiidae. Its length is 80 cm and it fed on frogs, lizards, and small mammals. It is found in Australia and Argentina.-References:*...
Albino, 1986
- Alamitophis argentinus Albino, 1986 (Cretaceous, Campanian or Maastrichtian; Argentina)
- Alamitophis elongatus Albino, 1994 (Cretaceous, Campanian or Maastrichtian; Argentina)
- Alamitophis tingamarra Scanlon, 2005 (Scanlon 1993; Paleogene, early Eocene; Australia)
- Rionegrophis Albino, 1986
- Rionegrophis madtsoioides Albino, 1986 (Cretaceous, Campanian or Maastrichtian; Argentina)
- Yurlunggur Scanlon, 1992
- Yurlunggur camfieldensis Scanlon, 1992 (Neogene, middle Miocene; Australia)
- Yurlunggur spp. (Scanlon 2004, 2006; Paleogene-Neogene, late Oligocene
The Oligocene is a geologic epoch of the Paleogene period and extends from about 33.9 million to 23 million years before the present. As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
to late Pleistocene; Australia)
- Herensugea Rage, 1996
- Herensugea caristiorum Rage, 1996 (Cretaceous, Campanian or Maastrichtian; Spain
Spain , officially the Kingdom of Spain , is a country located in southwestern Europe on the Iberian Peninsula.
[The Spanish constitution does not establish any official denomination of the country, even though España , Estado español and Nación española are used interchangeably...]
)
- Nanowana Scanlon, 1997
- Nanowana godthelpi Scanlon, 1997 (Neogene, early to middle Miocene; Australia)
- Nanowana schrenki Scanlon, 1997 (Neogene, early to middle Miocene; Australia)
- Najash Apesteguía and Zaher, 2006
- Najash rionegrina Apesteguía and Zaher, 2006 (Cretaceous, Cenomanian; Argentina)
Unnamed specimens
- Madtsoiidae indet. (Rage 1987; Paleogene, Paleocene; Morocco
Morocco, officially the Kingdom of Morocco, is a country located in North Africa with a population of nearly 32 million and an area just under . Its capital is Rabat, and its largest city is Casablanca. Morocco has a coast on the Atlantic Ocean that reaches past the Strait of Gibraltar into the...
)
- Madtsoiidae indet. (Werner and Rage 1994, Rage and Werner 1999; Cretaceous, Cenomanian
The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...
; SudanSudan is a country in northeastern Africa. It is the largest country in Africa and in the Arab World, and tenth largest in the world by area...
)
- ?Madtsoiid (Rage and Prasad 1992; Cretaceous, Maastrichtian; India)
- ?Madtsoiid (Rage 1991; Paleogene, early Paleocene; Bolivia)
- ?Madtsoiidae indet. cf. Madtsoia sp. (Scanlon 2005; Paleogene, early Eocene; Australia)
- Madtsoiidae indet. (Folie and Codrea 2005; Cretaceous, Maastrichtian; Romania
Romania is a country located in Southeastern and Central Europe, North of the Balkan Peninsula, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea. Almost all of the Danube Delta is located within its territory...
)
- Madtsoiidae nov. (Gomez and Baez 2006; Cretaceous, late Campanian or early Maastrichtian; Argentina)
Phylogeny
According to a cladistic analysis by Scanlon (2006),
Wonambi and
Yurlunggur as representative genera of Madtsoiidae form a monophyletic assembly. However, as
Madtsoia is not included, its grouping in the same family is questionable.