Madtsoiidae
Encyclopedia
Madtsoiidae are an extinct group of mostly Gondwana
n snakes with a fossil record extending from early Cenomanian
(Upper Cretaceous) to late Pleistocene
strata
located in South America
, Africa
, India
, Australia
and Southern Europe
. Madtsoiid snakes include very primitive
snakes, which like extant boas and pythons would likely dispatch their prey by constriction
, such as Gigantophis, one of the longest snakes known at an estimated 10.7 meters, and the Australian Aboriginal mythology
-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.
. With the recent use of cladistics
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 jaw
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 vertebrae, 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 group
), 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 skull
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 Dinilysia
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 Pachyrhachis
, Haasiophis
or Eupodophis
, 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 sacrum
, whereas it has rarely been questioned that the cloaca
l vertebrae in snakes are homologous
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 Serpent
s or dragon
s, including Wonambi (Pitjantjatjara), Yurlunggur (Yolngu
) and Nanowana (Ancient Greek
nano-, 'dwarf' + Warlpiri
Wana
) in Australia, and Herensugea (Basque
) 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 language
, 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 Spanish
name of the type locality, Cañadón Vaca.
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...
n snakes with a fossil record extending from early Cenomanian
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...
(Upper Cretaceous) to late Pleistocene
Pleistocene
The Pleistocene is the epoch from 2,588,000 to 11,700 years BP that spans the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....
strata
Stratum
In geology and related fields, a stratum is a layer of sedimentary rock or soil with internally consistent characteristics that distinguish it from other layers...
located in South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
, Africa
Africa
Africa 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...
, India
India
India , 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 with over 1.2 billion people, and the most populous democracy in the world...
, Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...
and Southern Europe
Europe
Europe 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 watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...
. Madtsoiid snakes include very primitive
Basal (phylogenetics)
In 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
Constriction
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 prey...
, such as Gigantophis, one of the longest snakes known at an estimated 10.7 meters, and the Australian Aboriginal mythology
Australian Aboriginal mythology
Australian 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
Linnaean taxonomy can mean either of two related concepts:# the particular form of biological classification set up by Carl Linnaeus, as set forth in his Systema Naturæ and subsequent works...
. With the recent use of cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...
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 jaw
Jaw
The 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 vertebrae, 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 group
Crown group
A crown group is a group consisting of living representatives, their ancestors back to the most recent common ancestor of that group, and all of that ancestor's descendants. The name was given by Willi Hennig, the formulator of phylogenetic systematics, as a way of classifying living organisms...
), 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 skull
Skull
The skull is a bony structure in the head of many animals that supports the structures of the face and forms a cavity for the brain.The skull is composed of two parts: the cranium and the mandible. A skull without a mandible is only a cranium. Animals that have skulls are called craniates...
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 Dinilysia
Dinilysia
Dinilysia is an extinct genus of snake from the Late Cretaceous of South America. The snake reached a length of 6-10 feet and preyed on smaller animals...
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 Pachyrhachis
Pachyrhachis
Pachyrhachis 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...
, Haasiophis
Haasiophis
Haasiophis, 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 Eupodophis
Eupodophis
Eupodophis is an extinct genus of snake from the Late Cretaceous period. It has two small hind legs and is considered a transitional form between Cretaceous lizards and limbless snakes. The feature, described as vestigial, was most likely useless to Eupodophis. The type species Eupodophis...
, 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 sacrum
Sacrum
In vertebrate anatomy the 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 cloaca
Cloaca
In 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...
l vertebrae in snakes are homologous
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
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 Serpent
Rainbow Serpent
The Rainbow Serpent is a common motif in the art and mythology of Aboriginal Australia. It is named for the snake-like meandering of water across a landscape and the colour spectrum caused when sunlight strikes water at an appropriate angle relative to the observer.The Rainbow Serpent is seen as...
s or dragon
Dragon
A dragon is a legendary creature, typically with serpentine or reptilian traits, that feature in the myths of many cultures. There are two distinct cultural traditions of dragons: the European dragon, derived from European folk traditions and ultimately related to Greek and Middle Eastern...
s, including Wonambi (Pitjantjatjara), Yurlunggur (Yolngu
Yolngu
The Yolngu or Yolŋu are an Indigenous Australian people inhabiting north-eastern Arnhem Land in the Northern Territory of Australia. Yolngu means “person” in the Yolŋu languages.-Yolŋu law:...
) and Nanowana (Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...
nano-, 'dwarf' + Warlpiri
Warlpiri language
The 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.-...
Wana
Wana
Wana 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-Colonial history:...
) in Australia, and Herensugea (Basque
Basque language
Basque 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 spoken by 25.7% of Basques in all territories...
) 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 language
Tehuelche language
Tehuelche 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...
, 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 Spanish
Spanish language
Spanish , also known as Castilian , is a Romance language in the Ibero-Romance group that evolved from several languages and dialects in central-northern Iberia around the 9th century and gradually spread with the expansion of the Kingdom of Castile into central and southern Iberia during the...
name of the type locality, Cañadón Vaca.
Classification
- Gigantophis Andrews, 1901
- Gigantophis garstiniGigantophis garstiniGigantophis garstini was a prehistoric snake which may have measured more than , 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; PaleogenePaleogeneThe 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 EoceneEoceneThe Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
; EgyptEgyptEgypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...
, LibyaLibyaLibya is an African country in the Maghreb region of North Africa bordered by the Mediterranean Sea to the north, Egypt to the east, Sudan to the southeast, Chad and Niger to the south, and Algeria and Tunisia to the west....
)
- Gigantophis garstini
- MadtsoiaMadtsoiaMadtsoia is an extinct genus of madtsoiid snake. It is known from the Eocene of Argentina , the Paleocene of Brazil , the Late Cretaceous of Spain , the Late Cretaceous of India , and the Late Cretaceous of Madagascar and possibly Niger Madtsoia is an extinct genus of madtsoiid snake. It is...
Simpson, 1933- Madtsoia bai Simpson, 1933 (Paleogene, Early Eocene; ArgentinaArgentinaArgentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...
) - Madtsoia cf. M. bai (Simpson 1935, Hoffstetter 1960; Paleogene, Late PaleocenePaleoceneThe Paleocene or Palaeocene, the "early recent", is a geologic epoch that lasted from about . It is the first epoch of the Palaeogene Period in the modern Cenozoic Era...
; Argentina) - Madtsoia madagascariensis Hoffstetter, 1961a (Piveteau 1933; CretaceousCretaceousThe Cretaceous , derived from the Latin "creta" , 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 the Jurassic period and is followed by the Paleogene period of the...
, SantonianSantonianThe Santonian is an age in the geologic timescale or a chronostratigraphic 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 CampanianCampanianThe 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 ...
; MadagascarMadagascarThe Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
) - Madtsoia aff. madagascariensis (de Broin et al. 1974; Cretaceous, ConiacianConiacianThe 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, NigerNigerNiger , officially named 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 MaastrichtianMaastrichtianThe 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; BrazilBrazilBrazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
)
- Madtsoia bai Simpson, 1933 (Paleogene, Early Eocene; Argentina
- WonambiWonambiWonambi is a genus that consisted of two species of very large snakes. These species were not pythons, like Australia's other large constrictors of the genus Morelia, and are now a member of an extinct family Madtsoiidae...
Smith, 1976- Wonambi naracoortensis Smith, 1976 (Scanlon and Lee 2000, Scanlon 2005; Neogene, PliocenePlioceneThe Pliocene Epoch is the period in the geologic timescale that extends from 5.332 million to 2.588 million years before present. It is the second and youngest epoch of the Neogene Period in the Cenozoic Era. The Pliocene follows the Miocene Epoch and is followed by the Pleistocene Epoch...
to Pleistocene; Australia) - Wonambi barriei Scanlon in Scanlon and Lee, 2000 (NeogeneNeogeneThe Neogene is a geologic period and system in the International Commission on Stratigraphy Geologic Timescale starting 23.03 ± 0.05 million years ago and ending 2.588 million years ago...
, early MioceneMioceneThe Miocene is a geological epoch of the Neogene Period and extends from about . 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 Pliocene. The Miocene follows the Oligocene...
; Australia)
- Wonambi naracoortensis Smith, 1976 (Scanlon and Lee 2000, Scanlon 2005; Neogene, Pliocene
- Patagoniophis Albino, 1986
- Patagoniophis parvus Albino, 1986 (Cretaceous, Campanian or Maastrichtian; Argentina)
- Patagoniophis australiensis Scanlon, 2005 (Scanlon 1993; Paleogene, early Eocene; Australia)
- AlamitophisAlamitophisAlamitophis 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 OligoceneOligoceneThe Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 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; SpainSpainSpain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...
)
- Herensugea caristiorum Rage, 1996 (Cretaceous, Campanian or Maastrichtian; Spain
- 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)
- SanajehSanajehSanajeh is a genus of late Cretaceous madtsoiid snake from western India. A recently described fossil from the Lameta Formation has been found coiled around an egg and an adjacent skeleton of a 50 cm long sauropod dinosaur hatchling...
Wilson et al., 2010- Sanajeh indicus Wilson et al., 2010 (Cretaceous, Maastrichtian; India)
- KelyophisKelyophisKelyophis is an extinct genus of nigerophiid snake which existed in Madagascar during the Late Cretaceous. The type species is Kelyophis hechti. Trunk vertebrae have been found from the Maastrichtian-age Maevarano Formation in the Mahajanga Basin...
Laduke et al., 2010- Kelyophis hechti Laduke et al., 2010 (Late Cretaceous, Maastrichtian; Madagascar)
- MenaranaMenaranaMenarana is an extinct genus of madtsoiid snake which existed in Madagascar during the Late Cretaceous. The type species is Menarana nosymena...
Laduke et al., 2010- Menarana nosymena Laduke et al., 2010 (Late Cretaceous, Maastrichtian; Madagascar)
Unnamed specimens
- Madtsoiidae indet. (Rage 1987; Paleogene, Paleocene; MoroccoMoroccoMorocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...
) - Madtsoiidae indet. (Werner and Rage 1994, Rage and Werner 1999; Cretaceous, CenomanianCenomanianThe 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...
; SudanSudanSudan , officially the Republic of the Sudan , is a country in North Africa, sometimes considered part of the Middle East politically. It is bordered by Egypt to the north, the Red Sea to the northeast, Eritrea and Ethiopia to the east, South Sudan to the south, the Central African Republic to the...
) - ?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; RomaniaRomaniaRomania is a country located at the crossroads of Central and Southeastern Europe, on the Lower Danube, within and outside the Carpathian arch, bordering on the Black Sea...
) - Madtsoiidae nov. (Gomez and Baez 2006; Cretaceous, late Campanian or early Maastrichtian; Argentina)