A Late Cretaceous (Maastrichtian) snake assemblage from the Maevarano Formation, Mahajanga Basin, Madagascar

Figures & data

FIGURE 1 Late Cretaceous and Paleocene strata of the Mahajanga Basin, northwestern Madagascar. The Berivotra and Masiakakoho study areas are indicated by rectangular outlines.

FIGURE 2 Trunk vertebrae of Madtsoia madagascariensis from the Late Cretaceous of Madagascar in l, lateral; a, anterior; p, posterior; d, dorsal; and v, ventral views. A, vertebra from anterior part of anterior trunk region with well-developed hypapophysis, FMNH PR 2546; B, vertebra from posterior part of anterior trunk region, FMNH PR 2549; C, mid-trunk vertebra, FMNH PR 2551; D, vertebra from anterior part of posterior trunk region, FMNH PR 2554; E, vertebra from middle part of posterior trunk region, FMNH PR 2555.

FIGURE 3 Cloacal and postcloacal vertebrae of Madtsoia madagascariensis from the Late Cretaceous of Madagascar in l, lateral; a, anterior; p, posterior; d, dorsal; and v, ventral views. A, cloacal vertebra, FMNH PR 2556; B, postcloacal vertebra, FMNH PR 2557. Articular facets on ventral aspect of postcloacal vertebra for chevron bone enlarged at bottom right.

FIGURE 4 Ribs and rib fragments of Madtsoia madagascariensis from the Late Cretaceous of Madagascar. A, anterior; and B, posterior views of UA 9764, nearly complete left rib exhibiting a pathological lesion (indicated by arrows). C, anterior; D, posterior; and I, stereophotographic proximal views of UA 9746, proximal half of right rib (reversed to facilitate comparison). E, anterior; F, posterior; and J, stereophotographic proximal views of FMNH PR 2571, proximal fragment of left rib. G, anterior; and H, posterior views of UA 9763, nearly complete left rib.

TABLE 1 Measurements of vertebral specimens of Madtsoia madagascariensis. See text for list of abbreviations. ? = vertebral fragment not assigned to region.

Specimen	Position	CL	NAW	PRW	POW	ZSW	COW	CNW	NSH	PR-PO
FMNH PR 2545	ATV	12.9	20.6	27.6	28.1	11.6	12.2	11.3	12.0	16.2
FMNH PR 2546	ATV	17.1	19.2	24	25.5	12.6	11.8	10.8	23.0	19.6
FMNH PR 2547	ATV	17.0	23.8	31.9	31.4	14.3	11.8	10.9	19.3	21.1
FMNH PR 2548	ATV	19.8	25	31.8	32.6	15.7	12.8	11.8	—	22.5
FMNH PR 2549	ATV	18	26	35.4	35.7	16.1	13.2	12.1	19.7	20.5
FMNH PR 2558	ATV	—	—	—	—	—	—	10.6*	—	—
FMNH PR 2702	ATV	16.4	21.7	—	30.0*	—	12.8	11.3	19.4	19.6
UA 9688	ATV	—	—	—	—	—	—	9.8*	—	—
UA 9689	ATV	15.7*	—	—	—	14.7*	14.3*	12.2*	—	18.4*
UA 9690	ATV	—	—	—	—	—	—	10.4	—	—
UA 9691	ATV	13.8*	—	—	—	—	11.7	10.4	—	—
UA 9692	ATV	16.1*	—	—	—	—	—	11.1*	—	—
UA 9693	ATV	—	—	—	—	—	—	9.7	—	—
UA 9703	ATV	—	—	—	—	—	—	12.0	—	—
UA 9728	ATV	—	—	—	—	—	—	11.7*	—	—
FMNH PR 2550	MTV	17.8*	27.2	—	36.0	—	15.5*	14.0	—	23.3
FMNH PR 2551	MTV	17.7	28.1	39.5	39.2	14.4	14.9	14.1	16.2	22.5
FMNH PR 2552	MTV	16.8*	25.9	38.7	—	—	14.3	13.6	14.6	22.7
FMNH PR 2553	MTV	18.6	29.2	41.2	41.6	15.3	15.4	14.6*	17.7	22.6
UA 9695	MTV	—	28.1	—	—	15.5*	—	—	—	—
UA 9697	MTV	—	—	—	33.2	—	12.1	—	—	—
UA 9698	MTV	—	—	—	—	12.4	—	—	—	22.0
UA 9745	MTV	16.4*	—	35.8	—	13.4*	14.9	13.3	—	21.1
FMNH PR 2554	PTV	15.6*	24.4	34.2	33.7	12.8	14.2	12.2	10.4	20.1
FMNH PR 2555	PTV	14.3	19.6	—	26.7	—	10.5	9.7*	10.1	18.2
UA 9700	PTV	—	—	—	—	—	—	14.0	—	23.8
FMNH PR 2556	CV	7.7*	13.5	19.4*	18.8*	8.1*	6.0*	5.2*	—	11.8*
FMNH PR 2557	PCV	—	9.9	—	—	5.2*	4.7	—	—	—
FMNH PR 2560	?	11.0	15.2	—	—	—	8.6*	7.7*	—	—
FMNH PR 2561a	?	—	—	—	—	17.1	—	—	—	—
FMNH PR 2561b	?	—	—	—	—	15.1	—	—	—	—
FMNH PR 2561c	?	—	—	—	—	15.9	—	—	—	—
FMNH PR 2562	?	—	—	—	—	—	—	17.8	—	—
FMNH PR 2564	?	—	—	—	—	19.4*	—	—	—	—
FMNH PR 2565	?	—	—	—	—	—	—	15.7*	—	—
FMNH PR 2566	?	—	—	—	—	—	—	18.2	—	—
FMNH PR 2567	?	—	—	—	27.6*	11.7*	—	—	—	—
FMNH PR 2584	?	—	—	—	—	16.3	—	—	—	—
FMNH PR 2585	?	—	—	—	—	10.8	—	—	—	—
UA 9694	?	—	19.4	28.2	—	—	10.8	—	—	17.4
UA 9696	?	11.6*	18.7	—	—	10.0*	11.0*	10.3*	9.9*	—
UA 9705	?	—	—	—	—	—	—	12.2	—	—
UA 9706	?	—	—	—	—	—	—	12.3	—	—
UA 9707	?	—	—	—	—	11.3	—	—	13.2	—
UA 9709	?	—	—	—	—	—	—	12.1	—	—
UA 9710	?	—	—	—	—	—	—	15.7*	—	—
UA 9712	?	—	—	—	—	—	—	14.9*	—	—
UA 9715	?	13.1*	—	—	—	9.5	—	7.0*	—	—
UA 9717	?	11.4*	—	—	—	—	—	8.0*	—	—
UA 9718	?	14.6*	—	—	—	—	—	10.1	—	—
UA 9726	?	15.8*	—	—	—	—	—	14.4*	—	—
UA 9727	?	—	—	—	—	12.0*	—	—	12.7	—
UA 9730	?	—	—	—	—	—	15.6*	—	—	—
UA 9731	?	—	—	—	—	—	12.1	—	—	—
UA 9735	?	16.1	—	—	—	—	—	10.9	—	—
UA 9736	?	—	—	—	—	19.3	—	—	16.5	—
UA 9738	?	20.7	—	—	—	—	—	15.3	—	—
UA 9739	?	—	—	—	—	—	—	9.9	—	—
UA 9740	?	—	—	—	—	—	—	—	—	14.5
UA 9741	?	15.4*	—	—	—	—	—	10.0	—	—
UA 9743	?	—	—	—	—	—	—	9.7	—	—

FIGURE 5 Braincase fragment, UA 9684-3 (part of holotype), of Menarana nosymena, gen. et sp. nov., from the Late Cretaceous of Madagascar. Stereophotographs (left and center) and interpretive drawings (right) of A, dorsal; B, ventral; C, left lateral; and D, right lateral views. Abbreviations: bbs, basisphenoid-basioccipital suture; ci, crista interfenestralis; ct, crista tuberalis; ds, dorsum sellae; eap, exoccipital ascending process; lr, lagenar recess; pbs, prootic-basisphenoid suture; pip, inferior process of prootic; rst, recessus scalae tympani; sot, spheno-occipital (basal) tubercle; vc, Vidian canal; VII, facial canal; VII h, foramen for hyomandibular branch of facial nerve; VII p, foramen for palatine branch of facial nerve; and XII, hypoglossal canal.

FIGURE 6 Braincase fragment, UA 9684-3 (part of holotype), of Menarana nosymena, gen. et sp. nov., from the Late Cretaceous of Madagascar in dorsolateral view (image obtained from HRXCT dataset). Abbreviations: bbs, basisphenoid-basioccipital suture; ci, crista interfenestralis; ds, dorsum sellae; eap, exoccipital ascending process; lc, lagenar crest; lr, lagenar recess; rst, recessus scalae tympani; VII, facial canal; and XII, hypoglossal canal.

FIGURE 7 Atlas, UA 9684-4 (part of holotype), of Menarana nosymena, gen. et sp. nov., from the Late Cretaceous of Madagascar. Stereophotographs of A, anterior; and B, posterior views.

FIGURE 8 Trunk vertebrae of Menarana nosymena, gen. et sp. nov., from the Late Cretaceous of Madagascar in l, lateral; a, anterior; p, posterior; d, dorsal; and v, ventral views. A, anterior trunk vertebra, UA 9687-1 (lateral view reversed); B, mid-trunk vertebra, UA 9684-1 (part of holotype); C, posterior trunk vertebra, UA 9684-2 (part of holotype).

TABLE 2 Measurements of vertebral specimens of Menarana nosymena, gen. et sp. nov. See text for list of abbreviations. ? = vertebral fragment not assigned to region.

Specimen	Position	CL	NAW	PRW	POW	ZSW	COW	CNW	NSH	PR-PO
UA 9687-1	ATV	8.1	7.5	11.0	—	4.5	4.3	4.1	—	9.8
UA 9687-2	ATV	8.5*	—	—	—	—	4.4*	—	—	—
FMNH PR 2543	MTV	8.7*	12.2	17.6	—	6.7	7.1	6.6	3.1	11.7
UA 9686	MTV	7.4*	7.4	11.1	—	—	4.7	4.0*	2.4	8.8
UA 9684-1	MTV	11.5	14.8*	—	18.9	6.9	8.3	7.6	3.2	15.1
UA 9684-7	MTV	10.4	—	—	—	—	—	5.1	—	—
UA 9694-8	MTV	12.6	14.5	21.2	—	7.4	8.4	7.9	3.3	16.0*
UA 9694-9	MTV	—	14.8*	20.9	—	7.2*	7.6	—	3.5	15.8
UA 9694-10	MTV	11.4	12.0	17.4	—	5.8	7.6	6.9	3.1	13.7
UA 9694-11	MTV	11.7	—	20.7	—	—	8.6	7.6	—	—
UA 9694-12	MTV	10.4	—	—	—	—	—	5.2	—	—
UA 9694-13	MTV	10.4	—	—	—	—	—	5.0	—	—
FMNH PR 2544	MTV	7.4	11.8	—	17.2	6.4*	—	5.2	3.5	—
FMNH PR 2703	MTV	8.2	9.8	14.2	—	5.5	6.2	5.3	2.5	9.8
UA 9684-2	PTV	11.3	12.4	17.8	—	—	8.0	7.2	3.5	14.1
UA 9684-5	PTV	—	11.5	17.6	—	—	7.0	—	3.3	—
FMNH PR 2542	PTV	11.4	—	—	—	—	—	7.7	—	—
UA 9684-6	?	—	—	—	—	7.0	—	—	—	—
UA 9685	?	—	—	—	—	8.0	—	—	3.8	—
UA 9733	?	9.0	—	—	—	—	—	6.4	—	—

FIGURE 9 Proximal fragment of right rib, UA 9684-5 (part of holotype), of Menarana nosymena, gen. et sp. nov., from the Late Cretaceous of Madagascar in A, anterior; B, posterior, and C, stereophotographic proximal views.

FIGURE 10 Trunk vertebrae of Kelyophis hechti, gen. et sp. nov., from the Late Cretaceous of Madagascar in l, lateral; a, anterior; p, posterior; d, dorsal; and v, ventral views. A, anterior trunk vertebra, FMNH PR 2539 (lateral view reversed); B, mid-trunk vertebra, UA 9682 (holotype).

TABLE 3 Measurements of vertebral specimens of Kelyophis hechti, gen. et sp. nov. See text for list of abbreviations.

Specimen	Position	CL	NAW	PRW	POW	ZSW	COW	CNW	NSH	PR-PO
FMNH PR 2540	ATV	5.7	4.1	6.7	—	—	2.8	2.4	—	6.6
UA 9683	ATV	7.6	—	—	—	—	—	3.4	—	—
FMNH PR 2539	MTV	3.6	2.2	3.7	3.3	1.6	1.7	—	—	3.8
FMNH PR 2541	MTV	3.4*	—	—	—	—	—	1.9*	—	—
UA 9682	MTV	3.0	2.1	3.5	3.5	1.5	1.5	1.3	0.3	3.7
UA 9725	?	7.0*	—	—	—	—	—	3.2*	—	—

FIGURE 11 Temporal and geographic distribution of madtsoiid (localities indicated by solid symbols, numbers) and nigerophiid (open symbols, letters) snakes. See Tables 4 and 5 for more detailed information.

TABLE 4 Temporal and geographic distribution of madtsoiid and questionably madtsoiid snakes (arranged alphabetically by genus and species, and then by more uncertain family-level attributions). : = Early; = Middle; = Late.

Taxon	Locality	Age	Reference
1. Alamitophis argentinus	Los Alamitos, Argentina	Campanian/Maastrichtian	Albino, 1986
2. Alamitophis argentinus	La Colonia, Argentina	Campanian/Maastrichtian	Albino, 2000
3. Alamitophis argentinus	Salinas de Trapalcó, Argentina	Campanian/Maastrichtian	Albino, 1994; Albino, 2007
4. Alamitophis argentinus	El Palomar, Argentina	Campanian/Maastrichtian	Albino, 1994
5. Alamitophis argentinus	Salitral de Santa Rosa, Argentina	Campanian/Maastrichtian	Martinelli and Forasiepi, 2004; Albino, 2007
6. Alamitophis elongatus	Salinas de Trapalcó, Argentina	Campanian/Maastrichtian	Albino, 1994; Albino, 2007
7. Alamitophis elongatus	Los Alamitos, Argentina	Campanian/Maastrichtian	Albino, 1994
8. Alamitophis tingamarra	Tingamarra, Australia	E. Eocene	Scanlon, 2005b
9. Gigantophis garstini	Dor et Talha, Libya	L. Eocene	Hoffstetter, 1961b
10. Gigantophis garstini	Fayum, Egypt	L. Eocene*	Andrews, 1901, 1906
11. Herensugea caristiorum	Laño, Spain	L. Campanian	Rage, 1996a, 1999
12. Madtsoia bai	Cañadón Vaca, Argentina	E. Eocene	Simpson, 1933
13. Madtsoia cf. M. bai	Gaiman, Argentina	L. Paleocene	Hoffstetter, 1959; Simpson, 1935
14. Madtsoia camposi	Itaborai, Brazil	M. Paleocene	Rage, 1998
15. Madtsoia laurasiae (= Menarana laurasiae in this paper)	Laño, Spain	L. Campanian	Rage, 1996a, 1999
16. Madtsoia madagascariensis	Mahajanga Basin, Madagascar	Maastrichtian	Hoffstetter, 1961a; LaDuke et al., this paper
17. Madtsoia aff. M. madagascariensis (= ?Madtsoia sp. in this paper)	In Beceten, Niger	E. Senonian	de Broin et al., 1974; Rage, 1981
18. ?Madtsoia sp.	Cañadón Hondo, Argentina	L. Paleocene	Albino, 1993
19. ?Madtsoia sp.	Cañadón Vaca, Argentina	E. Eocene	Albino, 1993
20. ?Madtsoia sp.	Gran Barranca, Argentina	E. Eocene	Albino, 1993
21. Menarana nosymena	Mahajanga Basin, Madagascar	Maastrichtian	LaDuke et al., this paper
22. Najash rionegrina (?Madtsoiidae)	La Buitrera, Argentina	E. Cenomanian	Apesteguía and Zaher, 2006
23. Nanowana godthelpi	Riversleigh, Australia	E. Miocene	Scanlon, 1997
24. Nanowana schrenki	Riversleigh, Australia	E. Miocene	Scanlon, 1997
25. Patagoniophis australiensis	Tingamarra, Australia	E. Eocene	Scanlon, 2005b
26. Patagoniophis parvus	Los Alamitos, Argentina	Campanian/Maastrichtian	Albino, 1986
27. Patagoniophis parvus	Salitral de Santa Rosa, Argentina	Campanian/Maastrichtian	Martinelli and Forasiepi, 2004; Albino, 2007
28. Rionegrophis madtsoioides	Los Alamitos, Argentina	Campanian/Maastrichtian	Albino, 1986, 2007
29. Wonambi barriei	Riversleigh, Australia	L. Oligocene/E. Miocene	Scanlon and Lee, 2000
30. Wonambi naracoortensis	Naracoorte, Australia	Pleistocene	Smith, 1976
31. Wonambi naracoortensis	Mammoth Cave, Australia	Pleistocene	Scanlon, 1995
32. Wonambi naracoortensis	Koala Cave, Australia	Pleistocene	Scanlon, 1995
33. Wonambi naracoortensis	Wellington Caves, Australia	Pleistocene	Scanlon, 1995
34. Wonambi naracoortensis	Corra-Lynn Cave, Australia	E. Pliocene?	Scanlon, 1995; 2005a
35. Wonambi naracoortensis	Tight Entrance Cave, Australia	Pleistocene	Scanlon, 2005a
36. Yurlunggur camfieldensis	Bullock Creek, Australia	M. Miocene	Scanlon, 1992
37. Yurlunggur sp.	Kanunka, Australia	Pliocene	Scanlon, 1995; 2004
38. Yurlunggur sp.	Tarkarooloo, Australia	L. Oligocene	Scanlon, 2004
39. Yurlunggur sp.	Lake Ngapakaldi, Australia	E. or M. Miocene	Scanlon, 2004
40. Yurlunggur sp.	Chinchilla, Australia	E. or M. Pliocene	Mackness and Scanlon, 1999; Scanlon, 2004
41. Yurlunggur sp.	Gogolo-Garnpung, Willandra Lakes, Australia	L. Pleistocene	Scanlon, 2004
42. Yurlunggur sp.	Kangaroo Well, Australia	L. Oligocene	Megirian et al., 2004
43. Yurlunggur sp.	Wyandotte, Australia	L. Pleistocene	Scanlon, 1995; 2004
44. Yurlunggur sp. or spp.	Riversleigh, Australia	L. Oligocene/E. Miocene	Scanlon, 2006
45. Madtsoiidae indet.	Salinas de Trapalcó, Argentina	Campanian/Maastrichtian	Gomez and Baez, 2006
46. Madtsoiidae indet.	Wadi Abu Hashim, Sudan	Cenomanian	Rage and Werner, 1999
47. Madtsoiidae indet.	Adrar Mgorn, Morocco	L. Paleocene	Gheerbrant et al., 1993
48. Madtsoiidae indet.	Yacimiento Las Flores, Argentina	M. Paleocene	Albino, 1993
49. Madtsoiidae indet.	Haţeg Basin, Romania	Maastrichtian	Folie and Codrea, 2005
50. Madtsoiidae indet.	Kelapur, Maharashtra, India	Maastrichtian	Rage et al., 2004
51. Madtsoiidae indet.	Salitral de Santa Rosa, Argentina	Campanian/Maastrichtian	Martinelli and Forasiepi, 2004; Albino, 2007
52. ?Madtsoiidae	Takli, India	Maastrichtian	Gayet et al., 1985
53. ?Madtsoiidae	Champ-Garimond, France	Campanian	Sigé et al., 1997
54. ?Madtsoiidae	Los Alamitos, Argentina	Campanian/Maastrichtian	Albino, 2007
55. ?Madtsoiidae	La Colonia, Argentina	Campanian/Maastrichtian	Albino, 2000
56. ?Madtsoiidae	Ranquil-Có, Argentina	Campanian/Maastrichtian	González Riga, 1999; Albino, 2007
57. ?Madtsoiidae indet.	Salitral de Santa Rosa, Argentina	Campanian/Maastrichtian	Martinelli and Forasiepi, 2004; Albino, 2007
58. ?Madtsoiidae indet.	Vastan Lignite Mine, Gujarat, India	E. Eocene	Rage et al., 2008
59. ?Madtsoiidae indet cf. Madtsoia sp.	Tingamarra, Australia	E. Eocene	Scanlon, 2005b
60. ?Madtsoiidae or Boidae	Tiupampa, Bolivia	E. Paleocene	Rage, 1991
61. ?Madtsoiidae or Boidae	La Colonia, Argentina	Campanian/Maastrichtian	Albino, 2000
62. ?Madtsoiidae or Boidae	Panandhro Mine, Kutch, India	E. Eocene	Rage et al., 2003
63. ?Madtsoiidae or Boidae	Asifabad, India	Maastrichtian	Rage et al., 2004
64. ?Madtsoiidae or Boidae	Pisdura, India	Maastrichtian	Rage et al., 2004

TABLE 5 Temporal and geographic distribution of nigerophiid and questionably nigerophiid snakes (arranged alphabetically).

Taxon	Locality	Age	Reference
A. Kelyophis hechti	Mahajanga Basin, Madagascar	Maastrichtian	LaDuke et al., this paper
B. Nigerophis mirus	Krebb de Sassao & Tillia, Niger	Paleocene	Rage, 1975
C. Nubianophis afaahus	Wadi Abu Hashim, Sudan	Cenomanian	Rage and Werner, 1999
D. Nubianophis cf. N. afaahus	Wadi Abu Hashim, Sudan	Cenomanian	Rage and Werner, 1999
E. Indophis sahnii (?Nigerophiidae)	Naskal, India	Maastrichtian	Prasad and Rage, 1995
F. Indophis sahnii (?Nigerophiidae)	Anjar, Gujarat, India	Maastrichtian	Rage et al., 2004
G. ?Indophis sahnii (?Nigerophiidae)	Kelapur, Maharashtra, India	Maastrichtian	Rage et al., 2004
H. “Nessovophis” zhylga (?Nigerophiidae)	Zhylga, Kazakhstan	Early Eocene	Averianov, 1997; Rage et al., 2003
I. Woutersophis novus (?Nigerophiidae)	Van Pachtenbeke Sand Pit, Belgium	Middle Eocene	Rage, 1980

Albino , A. M. 1986 . “ Nuevos Boidae Madtsoiinae en el Cretacico tardio de Patagonia (Formacion Los Alamitos, Rio Negro, Argentina) ” . In Simposio Evolucion de los Vertebrados Mesozoicos , Edited by: Bonaparte , J. F. 15 – 21 . Mendoza : Actas IV Congreso Argentino de Paleontología y Bioestratigrfía .

 Google Scholar

Albino , A. M. 2000 . New record of snakes from the Cretaceous of Patagonia (Argentina) . Geodiversitas , 22 : 247 – 253 .

 Google Scholar

Albino , A. M. 1994 . Una nueva serpiente (Reptilia) en el Cretácico superior de Patagonia, Argentina . Pesquisas , 21 : 58 – 63 .

 Google Scholar

Albino , A. M. 2007 . “ Lepidosauria; pp ” . In Patagonian Mesozoic Reptiles , Edited by: Gasparini , Z. , Salgado , L. and Coria , R. A. 87 – 115 . Indiana : Indiana University Press, Bloomington & Indianapolis .

 Google Scholar

Martinelli , A. G. and Forasiepi , A. M. 2004 . Late Cretaceous vertebrates from Bajo de Santa Rosa (Allen Formation), Río Negro province, Argentina, with the description of a new sauropod dinosaur (Titanosauridae) . Revista del Museo Argentino de Ciencias Naturales , 6 : 257 – 305 .

 Google Scholar

Scanlon , J. D. 2005b . Australia's oldest known snakes: Patagoniophis, Alamitophis, and cf. Madtsoia (Squamata: Madtsoiidae) from the Eocene of Queensland . Memoirs of the Queensland Museum , 51 : 215 – 235 .

 Google Scholar

Hoffstetter , R. 1961b . Nouvelles récoltes de serpents fossiles dans l’Éocène supérieur du désert Libyque , Vol. 33 , 326 – 331 . Paris : Bulletin du Muséum national d’Histoire naturelle .

 Google Scholar

Andrews , C. W. 1901 . Preliminary note on some recently discovered extinct vertebrates from Egypt (Part II) . Geological Magazine , 8 : 434 – 444 .

 Google Scholar

Rage , J.-C. 1996a . Les Madtsoiidae (Reptilia, Serpentes) du Crétacé supérieur d’Europe: témoins gondwaniens d’une dispersion transtéthysienne . Comptes Rendus de l’Académie des Sciences Paris (2) , 322 : 603 – 608 .

 Google Scholar

Simpson , G. G. 1933 . A new fossil snake from the Notostylops beds of Patagonia . Bulletin of the American Museum of Natural History , 67 : 1 – 22 .

 Google Scholar

Hoffstetter , R. 1959 . Un serpent terrestre dans le Cretace inférieur du Sahara . Bulletin de la Societé géologique de France , 7 : 897 – 902 .

 Google Scholar

Simpson , G. G. 1935 . Early and middle Tertiary geology of the Gaiman region, Chubut, Argentina . American Museum Novitates , 775 : 1 – 29 .

 Google Scholar

Rage , J.-C. 1998 . Fossil snakes from the Palaeocene of São José de Itaborai, Brazil. Part 1 . Madtsoiidae, Aniliidae. Palaeovertebrata, Montpellier , 27 : 109 – 144 .

 Google Scholar

Hoffstetter , R. 1961a . Nouveaux restes d’un serpent boïdé (Madtsoia madagascariensis nov. sp.) dans le Crétacé supérieur de Madagascar , Vol. 33 , 152 – 160 . Paris : Bulletin du Muséum national d’Histoire naturelle .

 Google Scholar

de Broin , F. , Buffetaut , E. , Koeniguer , J. C. , Rage , J.-C. , Taquet , P. , Vergnaud-Grazzini , C. and Wenz , S. 1974 . La faune de Vertébrés continentaux du gisement d’In Beceten (Sénonien du Niger) . Comptes rendus de l’Académie des Sciences, Paris , 279 : 469 – 472 .

 Google Scholar

Rage , J.-C. 1981 . Les continents péri-atlantiques au Crétacé supérieur: migrations des faunes continentales et problèmes paléogéographiques . Cretaceous Research , 2 : 65 – 84 .

 Google Scholar

Albino , A. M. 1993 . Snakes from the Paleocene and Eocene of Patagonia, Argentina: paleoecology and coevolution with mammals . Historical Biology , 7 : 51 – 69 .

 Google Scholar

Apesteguía , S. and Zaher , H. 2006 . A Cretaceous terrestrial snake with robust hindlimbs and a sacrum . Nature , 440 : 1037 – 1040 .

 PubMed Web of Science ®Google Scholar

Scanlon , J. D. 1997 . Nanowana gen. nov., small madtsoiid snakes from the Miocene of Riversleigh; sympatric species with divergently specialized dentition . Memoirs of the Queensland Museum , 41 : 393 – 412 .

 Google Scholar

Scanlon , J. D. and Lee , M. S. Y. 2000 . The Pleistocene serpent Wonambi and the early evolution of snakes . Nature , 403 : 416 – 420 .

 PubMed Web of Science ®Google Scholar

Smith , M. J. 1976 . Small fossil vertebrates from Victoria Cave, Naracoorte, South Australia. IV. Reptiles . Transactions of the Royal Society of South Australia , 100 : 39 – 51 .

 Google Scholar

Scanlon , J. D. 1995 . First records from Wellington Caves, New South Wales, of the extinct madtsoiid snake Wonambi naracoortensis Smith, 1976 . Proceedings of the Linnean Society of New South Wales , 115 : 233 – 238 .

 Google Scholar

Scanlon , J. D. 2005a . Cranial morphology of the Plio-Pleistocene giant madtsoiid snake Wonambi naracoortensis . Acta Palaeontologica Polonica , 50 : 139 – 180 .

 Web of Science ®Google Scholar

Scanlon , J. D. 1992 . A new large madtsoiid snake from the Miocene of the Northern Territory . The Beagle, Records of the Northern Territory Museum of Arts and Sciences , 9 : 49 – 60 .

 Google Scholar

Scanlon , J. D. 2004 . First known axis vertebra of a madtsoiid snake (Yurlunggur camfieldensis) and remarks on the neck of snakes . The Beagle: Records of the Museums and Art Galleries of the Northern Territory , 20 : 207 – 215 .

 Google Scholar

Mackness , B. S. and Scanlon , J. D. 1999 . The first Pliocene record of the madtsoiid snake genus Yurlunggur Scanlon, 1992 from Queensland . Memoirs of the Queensland Museum , 43 : 783 – 785 .

 Google Scholar

Megirian , D. , Murray , P. , Schwartz , L. and von der Borch , C. 2004 . Late Oligocene Kangaroo Well Local Fauna from the Ulta Limestone (new name), and climate of the Miocene oscillation across Central Australia . Australian Journal of Earth Sciences , 51 : 701 – 741 .

 Web of Science ®Google Scholar

Scanlon , J. D. 2006 . Skull of the large non-macrostomatan snake Yurlunggur from the Australian Oligo-Miocene . Nature , 439 : 839 – 842 .

 PubMed Web of Science ®Google Scholar

Gomez , R. O. and Baez , A. M. 2006 . A new madtsoiid snake (Squamata, Ophidia) from the Upper Cretaceous of Patagonia , San Juan, , Argentina : XXII Jornadas Argentinas de Paleontología de Vertebrados . (2006):21

 Google Scholar

Rage , J.-C. and Werner , C. 1999 . Mid-Cretaceous (Cenomanian) Snakes from Wadi Abu Hashim, Sudan: the earliest snake assemblage . Palaeontologia Africana , 35 : 85 – 110 .

 Google Scholar

Gheerbrant , E. , Cappetta , H. , Feist , M. , Jaeger , J.-J. , Sudre , J. , Vianey-Liaud , M. and Sigé , B. 1993 . La succession des faunes de vertebrés d’âge paléocène supérieur et éocène inférieur dans le basin d’Ouarzazate, Maroc. Contexte géologique, portée biostratigraphique et paléogéographique . Newsletters in Stratigraphy , 28 : 33 – 58 .

 Google Scholar

Folie , A. and Codrea , V. 2005 . New lissamphibians and squamates from the Maastrichtian of Haţeg Basin, Romania . Acta Palaeontologica Polonica , 50 : 57 – 71 .

 Web of Science ®Google Scholar

Rage , J.-C. , Prasad , G. V. R. and Bajpai , S. 2004 . Additional snakes from uppermost Cretaceous (Maastrichtian) of India . Cretaceous Research , 25 : 425 – 434 .

 Web of Science ®Google Scholar

Gayet , M. , Rage , J.-C. and Rana , R. S. 1985 . “ Nouvelles ichthyofaune et herpétofaune de Gitti Khadan, le plus ancient gisement connu du Deccan (Crétacé/Paléocène) à microvertébrés. Implications paléogéographiques ” . In Paléogéographie de l’Inde, du Tibet et du Sud-Est Asiatique , Edited by: Jaeger , J.-J. , Rage , J.-C. and Buffetaut , E. 55 – 65 . Mémoire de la Societé géologique de France 147 .

 Google Scholar

Sigé , B. , Buscalioni , A. D. , Dugaud , S. , Gayet , M. , Orth , B. , Rage , J.-C. and Sanz , J. L. 1997 . Etat des données sur le gisement Crétacé supérieur continental de Champ-Garimond (Gard, Sud de la France) . Münchner Geowissenschaftliche Abhandlungen A , 34 : 111 – 130 .

 Google Scholar

González Riga , B. J. 1999 . Hallazgo de vertebrados fósiles en la Formación Loncoche, Cretácico Superior de la provincia de Mendoza, Argentina . Ameghiniana , 36 : 401 – 410 .

 Google Scholar

Rage , J.-C. , Folie , A. , Rana , R. S. , Singh , H. , Rose , K. D. and Smith , T. 2008 . A diverse snake fauna from the early Eocene of Vastan Lignite Mine, Gujarat, India . Acta Palaeontologica Polonica , 53 : 391 – 403 .

 Web of Science ®Google Scholar

Rage , J.-C. 1991 . “ Squamate reptiles from the early Paleocene of the Tiupampa area (Santa Lucia Formation), Bolivia ” . In Fosiles y Facies de Bolivia—Volume 1 Vertebrados , Edited by: Suarez-Soruco , R. 503 – 508 . Revista Técnica de YPFB 12 .

 Google Scholar

Rage , J.-C. , Bajpai , S. , Thewissen , J. G. M. and Tiwari , B. N. 2003 . Early Eocene snakes from Kutch, Western India, with a review of the Palaeophiidae . Geodiversitas , 25 : 695 – 716 .

 Web of Science ®Google Scholar

Seiffert , E. 2006 . Revised age estimates for the later Paleogene mammal faunas of Egypt and Oman . Proceedings of the National Academy of Science , 103 : 5000 – 5005 .

 PubMed Web of Science ®Google Scholar

Rage , J.-C. 1975 . Un serpent du Paléocène du Niger. Etude préliminaire sur l’origine des Caenophidiens (Reptilia, Serpentes) . Comptes Rendus de l’Académie des Sciences, Paris (D) , 281 : 515 – 518 .

 Google Scholar

Prasad , G. V. R. and Rage , J.-C. 1995 . Amphibians and squamates from the Maastrichtian of Naskal, India . Cretaceous Research , 16 : 95 – 107 .

 Web of Science ®Google Scholar

Averianov , A. O. 1997 . Paleogene sea snakes from the eastern part of Tethys . Russian Journal of Herpetology , 4 : 128 – 142 .

 Google Scholar

Rage , J.-C. 1980 . Un serpent marin nouveau de l’Éocene de Belgique. Le problème des serpents marine du Paléogène . Comptes Rendus de l’Académie des Seances, Paris (D) , 291 : 469 – 471 .

 Google Scholar