The , or '''madtsoiid snakes''', 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 . This grouping of fossil species includes Very Primitive snakes, which like extant boas and pythons would likely dispatch their prey by constriction, such as ''Gigantophis'', the longest snake known at an estimated 10.7 meters, and the Australian Aboriginal Mythology -themed ''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). Further study and new finds allowed ranking the group as a distinct family in Linnaean Systems . 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 Python s and Boa s, 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 Vertebra 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 based on diagnostic vertebral characters described by Apesteguía and Zaher (2006).
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'', raising the possibility of misassociation. The type material of ''Najash'' is the only 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. The latter condition 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 Squamate s (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.
- '' Madtsoia '' Simpson, 1933
- ---''Madtsoia bai'' Simpson, 1933 (Paleogene, Early Eocene; Argentina )
- ---''Madtsoia'' cf. ''M. bai'' (Simpson 1935, Hoffstetter 1960; Paleogene, Late Paleocene ; Argentina)
- ---''Madtsoia madagascariensis'' Hoffstetter, 1961a (Piveteau 1933; Cretaceous , Santonian or Campanian ; Madagascar )
- ---''Madtsoia'' aff. ''madagascariensis'' (de Broin et al. 1974; Cretaceous, Coniacian or Santonian, Niger )
- ---''Madtsoia laurasiae'' Rage, 1996 (Astibia et al. 1990; Cretaceous, Campanian or Maastrichtian ; Spain)
- ---''Madtsoia camposi'' Rage, 1998 (Paleogene, middle Paleocene; Brazil )
- '' Wonambi '' Smith, 1976
- ---''Wonambi naracoortensis'' Smith, 1976 (Scanlon and Lee 2000, Scanlon 2005; Neogene, Pliocene to Pleistocene; Australia)
- ---''Wonambi barriei'' Scanlon in Scanlon and Lee, 2000 ( Neogene , early Miocene ; Australia)
- '' Patagoniophis '' Albino, 1986
- ---''Patagoniophis parvus'' Albino, 1986 (Cretaceous, Campanian or Maastrichtian; Argentina)
- ---''Patagoniophis australiensis'' Scanlon, 2005 (Scanlon 1993; Paleogene, early Eocene; Australia)
- '' Alamitophis '' 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 to late Pleistocene; Australia)
- '' Herensugea '' Rage, 1996
- ---''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)
- Madtsoiidae indet. (Rage 1987; Paleogene, Paleocene; Morocco )
- Madtsoiidae indet. (Werner and Rage 1994, Rage and Werner 1999; Cretaceous, Cenomanian ; Sudan )
- ?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 )
- Madtsoiidae nov. (Gomez and Baez 2006; Cretaceous, late Campanian or early Maastrichtian; Argentina)
According to a cladistic analysis by Scanlon (2006), ''Wonambi'' and ''Yurlunggur'' as representative genera of Madtsoiidae form a Monophyletic assembly.
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''Pachyrhachis''†
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''Haasiophis''†
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'''''Wonambi'''''†
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'''''Yurlunggur'''''†
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''Dinilysia''†
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'''Scolecophidia'''
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'''Alethinophidia'''
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