Maevarano Formation

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Maevarano Formation
Stratigraphic range: Maastrichtian
~70–65.8  Ma
O
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Pg
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Type Geological formation
Sub-unitsMasorobe, Anembalemba & Miadana Members
Underlies Berivotra Formation
Overlies Marovoay Beds
Thickness>105 m (344 ft)
Lithology
Primary Sandstone
Other Claystone, siltstone
Location
Coordinates 15°54′S46°36′E / 15.9°S 46.6°E / -15.9; 46.6
Approximate paleocoordinates 30°06′S38°24′E / 30.1°S 38.4°E / -30.1; 38.4
Region Mahajanga Province
CountryFlag of Madagascar.svg  Madagascar
Extent Mahajanga Basin
Type section
Named for Maevarano River
Named bySalètes
Year defined1895
Madagascar physical map.svg
Lightgreen pog.svg
Maevarano Formation (Madagascar)

The Maevarano Formation is a Late Cretaceous sedimentary rock formation found in the Mahajanga Province of northwestern Madagascar. It is most likely Maastrichtian in age, [1] and records a seasonal, semiarid environment with rivers that had greatly varying discharges. Notable animal fossils recovered include the theropod dinosaur Majungasaurus , the early bird Vorona , the paravian Rahonavis , the titanosaurian sauropod Rapetosaurus , and the giant frog Beelzebufo .

Contents

Description

The Maevarano Formation is well exposed in the Mahajanga Basin, in particular near the village of Berivotra near the northwestern coast of the island where its outcrops have been heavily dissected by erosion. At the time it was being deposited, its latitude was between 30°S and 25°S as Madagascar drifted northward after splitting from India about 88 million years ago. It is composed of three smaller units or members. The lowest is the Masorobe Member, which is usually reddish and is at least 80 metres (260 ft). Its rocks are mostly poorly sorted coarse-grained sandstones with some finer-grained beds. It is separated by an erosional disconformity from the next member, the Anembalemba Member. The lower portion of the Anembalemba Member is fine to coarse clay-rich sandstone, whitish or light grey in color, with cross-bedding. The upper portion of this member is made of poorly sorted clay-rich sandstone, light olive-grey in color, that lacks cross-bedding. Most vertebrate fossils come from the Anembalemba Member, especially from the upper portion. The Miadana Member, the third and uppermost member, is not always present, and is up to 25 metres (82 ft) in some places. Elsewhere, it is replaced by the marine Berivotra Formation. The Miadana Member is made up of claystone, siltstone, and sandstone, lacks cross-bedding, and has several colors of rock. The Maevarano Formation as a whole is underlain by the Marovoay beds and capped by the Berivotra Formation. [1]

The age of the Maevarano Formation has been debated; the Berivotra Formation, which is partially contemporaneous with the upper portions of the formation, shows that at least the upper part of the Maevarano is Maastrichtian in age. There is no evidence that it is Campanian, [1] despite previous reports to that effect. [2] The Berivotra Formation appears to include near its top a magnetic reversal, interpreted as the shift from Chron 30N to Chron 29R, which occurred approximately 65.8 million years ago (about 300,000 years before the Cretaceous–Paleogene boundary and associated Cretaceous–Paleogene extinction event. This suggests that Maevarano organisms also lived shortly before (geologically speaking) the extinction event. [1]

History of exploration

The Maevarano Formation was first explored by French military physician Dr. Félix Salètes and his staff officer Landillon in 1895, and fossils and geologic data were sent to paleontologist Charles Depéret. [3] He briefly described the formation and named two dinosaurs from the remains ( Titanosaurus madagascariensis and Megalosaurus crenatissimus, now Majungasaurus). [4] Similar collections were made throughout the 20th century, yielding mostly fragmentary fossils; [3] one such specimen, a rough partial skull roof, became the holotype of supposed pachycephalosaur (bonehead dinosaur) Majungatholus in 1979. [5] (This specimen was later shown to be part of the skull ornamentation of a Majungasaurus.) Large-scale expeditions (seven to date), under the banner of the Mahajanga Basin Project, began in 1993. These expeditions, conducted jointly by Stony Brook University and the University of Antananarivo, have greatly expanded knowledge of this formation and the organisms that lived while it was being deposited. [3]

Paleoenvironment

The interpretation of Majungasaurus, Masiakasaurus, and Rapetosaurus during the late Cretaceous Majungasaurus, Masiakasaurus, Rapetosaurus.jpg
The interpretation of Majungasaurus, Masiakasaurus, and Rapetosaurus during the late Cretaceous

The Maevarano Formation is interpreted as a low-relief alluvial plain that over time was covered by a marine transgression. Broad, shallow rivers flowed to the northwest from central highlands; evidence for debris flows suggests that the discharges of the rivers varied greatly, with periods of dilute water flow, and periods of rapid erosion dumping sediment into the channels. Paleosols are reddish and include root casts. The paleosols and other sedimentologic evidence indicate well-drained floodplains with abundant vegetation adapted to a relatively dry climate, strongly seasonal (rainy and dry seasons) and at times semiarid (not unlike the present climate of the area). [1]

Paleofauna

Animals found in the formation include frogs (including Beelzebufo ampinga ), [6] turtles, snakes, lizards, at least seven species of crocodyliforms (including species of Mahajangasuchus and Trematochampsa ), abelisaurid theropods Majungasaurus , noasaurid Masiakasaurus , two types of titanosaurian sauropods ( Rapetosaurus and Vahiny ), and at least five species of bird-like dinosaurs, including Rahonavis . The 6 to 7 metres (20 to 23 ft) long Majungasaurus was likely the apex predator in the terrestrial environment. Crocodyliforms were very diverse and abundant. [1]

Color key
Taxon Reclassified taxonTaxon falsely reported as presentDubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Invertebrates

Invertebrates
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Cubiculum [7] C. ornatusOvoid chambers in dinosaur bonesA trace fossil, possibly puppal chambers of carrion beetles
Ethmosestheria [8] E. mahajangaensisAnembalemba MemberA species of antronstheriid clam shrimp
Osteocallis [7] O. mandibulusCurved grooves on dinosaur bonesA trace fossil, possibly feeding marks by a similar insect as Cubiculum ornatus

Fish

Osteichthyes

Osteichthyes
TaxonSpeciesLocationStratigraphic positionMaterialNotesImages
Albula [9] A. sp.Tooth plates and dentariesA species of bonefish
Characiformes indet. [9] Partial jaw
Coelodus [9] C. sp.A tooth plateA species of pycnodontid
Cypriniformes? [9]
Dipnoi indet. [10] Masorobe and Anembalemba MembersBurrows
Egertonia [9] E. sp.Partial toothplates
Enchodus [9] E. sp.Teeth
Lepisosteus [11] L. sp.Scales, fin rays, teeth, vertebrae, skull bonesA type of gar
Paralbula [9] P. sp.Lac Kinkony MemberPartial tooth platesA species of bonefish
Sciaenidae indet. [9] A species of croaker
Siluriformes indet. [9] VertebraeA type of catfish
Vango [12] V. fahinyLac Kinkony MemberReferred material is hyomandibulae, although other partial remains of gonorynchiform probably belongs to this taxa as wellA chanid gonorynchiform, milkfish

Amphibians

Amphibians
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Beelzebufo Beelzebufo ampingalocality MAD98-25 [13] A large frog
Beelzebufo skeletal.png

Dinosaurs

Indeterminate Lithostrotia remains formerly attributed to Titanosauridae. Undescribed Lithostrotia form. Indeterminate Enantiornithes remains. A rich avifauna with several undescribed taxa are known, including pengornithid enantiornithes and putative omnivoropterygids. [14]

Ornithischians

Ornithischians of the Maevarano Formation
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Stegosaurus [15] S. madagascariensis [15] "Teeth" [16] An indeterminate thyreophoran. [17]

Sauropods

Sauropods of the Maevarano Formation
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Rapetosaurus [15] R. krausei [15] "[Three] skulls, at least [one] postcranial skeleton." [18] A titanosaur
Rapetosaurus BW.jpg
Titanosaurus [15] T. madagascariensis [15]
Vahiny [19] V. depereti [19] "Partial braincase" [19] A titanosaur

Theropods

Theropods of the Maevarano Formation
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Falcatakely [20] F. forsterae"Partial skull"A member of Enantiornithes
Falcatakely restoration.jpg
Majungasaurus [15] M. crenatissimus [15] "Scattered remains leading to nearly most of the animal." [21] An abelisaur
Majungasaurus size chart.png
Masiakasaurus [15] M. knopfleri [15] "Disarticulated remains of at least 6 individuals," as well as other isolated fossils. [22] A noasaurid abelisaur
Masiakasaurus knopfleri skull reconstruction.jpg
Rahonavis [15] R. ostromi [15] "Partial postcranial skeleton" [23] A paravian of unclear phylogenetic placement
Rahonavis NT.jpg
Vorona [15] V. berivotrensis [15] "Partial hindlimbs" [24] An ornithuromorph

Crocodylomorphs

Crocodylomorphs
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Mahajangasuchus M. insignisdisarticulated postcranial skeleton, multiple skull remainsA mahajangasuchid
Mahajangasuchus FMNH.jpg
Miadanasuchus M. oblitaA peirosaurid. Formerly known as Trematochampsa oblita
Simosuchus S. clarkimultiple specimens representing most of the skeletonA ziphosuchian
Simosuchus.jpg
Araripesuchus A. tsangatsanganaAnembalemba Membermultiple specimen including several skulls and one almost complete specimenA notosuchian
Araripesuchus wegeneri.jpg

Squamates

Squamates
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Kelyophis K. hechtiA nigerophiid snake
Konkasaurus K. mahalanaA cordylid lizard
Madtsoia M. madagascariensisA madtsoiid snake
Menarana M. nosymena Vertebrae and rib fragmentsA madtsoiid snake

Turtles

Turtles
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Kinkonychelys K. rogersiA side-necked turtle
Sahonachelys S. mailakavavaA sahonachelyid side-necked turtle
Sahonachelys life restoration.jpg
Sokatra S. antitraA sahonachelyid side-necked turtle

Mammals

Mammal remains include an undescribed gondwanathere, [25] a broken tooth UA 8699, which has been interpreted both as metatherian and as eutherian, a non-gondwanathere multituberculate tooth fragment, a non-gondwanathere multituberculate femur, [26] and a yet undescribed mammal known from an articulated skeleton. [27] Some taxa are particularly large sized herbivores, exemplifying the diversity of Mesozoic mammals. [28]

Mammals
GenusSpeciesLocationStratigraphic positionMaterialNotesImages
Adalatherium A. huiA gondwanatherian
Adalatherium skull.svg
Lavanify L. miolakateethA gondwanatherian
Vintana V. sertichiA gondwanatherian
Vintana NT small.jpg

See also

Related Research Articles

<span class="mw-page-title-main">Gondwanatheria</span> Extinct group of Mammaliaformes that lived during the Upper Cretaceous through the Miocene

Gondwanatheria is an extinct group of mammaliaforms that lived in parts of Gondwana, including Madagascar, India, South America, Africa, and Antarctica during the Upper Cretaceous through the Paleogene. Until recently, they were known only from fragmentary remains. They are generally considered to be closely related to the multituberculates and likely the euharamiyidians, well known from the Northern Hemisphere, with which they form the clade Allotheria.

Lavanify is a mammalian genus from the late Cretaceous of Madagascar. The only species, L. miolaka, is known from two isolated teeth, one of which is damaged. The teeth were collected in 1995–1996 and described in 1997. The animal is classified as a member of Gondwanatheria, an enigmatic extinct group with unclear phylogenetic relationships, and within Gondwanatheria as a member of the family Sudamericidae. Lavanify is most closely related to the Indian Bharattherium; the South American Sudamerica and Gondwanatherium are more distantly related. Gondwanatheres probably ate hard plant material.

<i>Rapetosaurus</i> Extinct genus of dinosaurs

Rapetosaurus is a genus of titanosaurian sauropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period. Only one species, Rapetosaurus krausei, has been identified.

<span class="mw-page-title-main">Allotheria</span> Extinct subclass of mammals

Allotheria is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower molariform teeth equipped with longitudinal rows of cusps and enlarged incisors. Typically, the canine teeth are also lost. Allotheria includes Multituberculata, Gondwanatheria, and probably Haramiyida, although some studies have recovered haramiyidans to be basal mammaliaforms unrelated to multituberculates. Allotherians are often placed as crown group mammals, more closely related to living marsupials and placentals (Theria) than to monotremes or eutriconodonts, though some studies place the entirety of Allotheria outside of crown Mammalia.

<i>Rajasaurus</i> Abelisaurid dinosaur genus from Late Cretaceous India

Rajasaurus is a genus of carnivorous abelisaurid theropod dinosaur from the Late Cretaceous of India, containing one species: Rajasaurus narmadensis. The bones were excavated from the Lameta Formation in the Gujarat state of Western India, probably inhabiting what is now the Narmada River Valley. It was formally described by palaeontologist Jeffrey A. Wilson and colleagues in 2003 based on a partial skeleton comprising the braincase, spine, hip bone, legs, and tail–a first for an Indian theropod. The dinosaur likely measured 6.6 metres (22 ft), and had a single horn on the forehead which was probably used for display and head-butting. Like other abelisaurids, Rajasaurus was probably an ambush predator.

<i>Masiakasaurus</i> Noasaurid theropod dinosaur genus from the Late Cretaceous period

Masiakasaurus is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, masiaka means "vicious"; thus, the genus name means "vicious lizard". The type species, Masiakasaurus knopfleri, was named after the musician Mark Knopfler, whose music inspired the expedition crew. It was named in 2001 by Scott D. Sampson, Matthew Carrano, and Catherine A. Forster. Unlike most theropods, the front teeth of M. knopfleri projected forward instead of straight down. This unique dentition suggests that they had a specialized diet, perhaps including fish and other small prey. Other bones of the skeleton indicate that Masiakasaurus were bipedal, with much shorter forelimbs than hindlimbs. M. knopfleri was a small theropod, reaching 1.8–2.1 m (5.9–6.9 ft) long and weighing 20 kg (44 lb).

<i>Rugops</i> Genus of dinosaur

Rugops is a monospecific genus of basal abelisaurid theropod dinosaur from Niger that lived during the Late Cretaceous period in what is now the Echkar Formation. The type and only species, Rugops primus, is known only from a partial skull. It was named and described in 2004 by Paul Sereno, Jeffery Wilson and Jack Conrad. Rugops has an estimated length of 4.4–5.3 metres (14–17 ft) and weight of 410 kilograms (900 lb). The top of its skull bears several pits which correlates with overlaying scale and the front of the snout would have had an armour-like dermis.

<i>Rahonavis</i> Extinct genus of dinosaurs

Rahonavis is a genus of bird-like theropod from the Late Cretaceous of what is now northwestern Madagascar. It is known from a partial skeleton found by Catherine Forster and colleagues in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province. Rahonavis was a small predator, at about 70 centimetres (2.3 ft) long and 0.45-2.27 kg, with the typical dromaesaurid-like raised sickle claw on the second toe. It was originally the first African coelurosaur until the discovery of Nqwebasaurus in 2000.

<span class="mw-page-title-main">Kristina Curry Rogers</span> American paleontologist

Kristina "Kristi" Curry Rogers is an American vertebrate paleontologist and a professor in Biology and Geology at Macalester College. Her research focuses on questions of dinosaur paleobiology, bone histology, growth, and evolution, especially in a subgroup of sauropods called Titanosauria. She has named two dinosaur species from Madagascar, Rapetosaurus, the most complete Cretaceous sauropod and titanosaur found to date, and Vahiny, so far known only from a partial skull. She and Jeffrey A. Wilson co-authored The Sauropods, Evolution and Paleobiology, published in December 2005. Her research includes field work in Argentina, Madagascar, Montana, South Africa, and Zimbabwe.

<i>Majungasaurus</i> Abelisaurid theropod dinosaur from the Late Cretaceous period

Majungasaurus is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 66 million years ago, at the end of the Cretaceous Period, making it one of the last-known non-avian dinosaurs that went extinct during the Cretaceous–Paleogene extinction event. The genus contains a single species, Majungasaurus crenatissimus. This dinosaur is also called Majungatholus, a name which is considered a junior synonym of Majungasaurus.

<i>Simosuchus</i> Extinct genus of reptiles

Simosuchus is an extinct genus of notosuchian crocodyliforms from the Late Cretaceous of Madagascar. It is named for its unusually short skull. Fully grown individuals were about 0.75 metres (2.5 ft) in length. The type species is Simosuchus clarki, found from the Maevarano Formation in Mahajanga Province, although one isolated multicuspid tooth of this genus was discovered in Kallamedu Formation of India.

<span class="mw-page-title-main">Berivotra Formation</span> Geological formation in Madagascar

The Berivotra Formation is a Maastrichtian sedimentary formation of the Mahajanga Basin in Boeny, Madagascar. The claystones of the formation were deposited in a shallow marine environment. The Berivotra Formation overlies the fossil-rich Maevarano Formation, in which more fossils of Beelzebufo, also recovered from the Berivotra Formation, have been found. Many shark and ray teeth were collected by surface prospecting on outcrops of the Berivotra Formation, which is readily distinguished by its yellowish grey to pale olive colour, as opposed to the white and green fluvial sandstones that form the upper 15 to 20 metres of the underlying Maevarano Formation.

<i>Madtsoia</i> Extinct genus of snakes

Madtsoia is an extinct genus of madtsoiid snakes. It is known from the Eocene of Argentina, the Paleocene of Brazil, the Late Cretaceous (Maastrichtian) of India, and the Late Cretaceous (Maastrichtian) of Madagascar. The type species was the largest with an estimated length of 9–10 m (30–33 ft), and the other three species were smaller. A 5.1 m (17 ft) long M. madagascariensis would have weighed 50 kg (110 lb), but an isolated specimen suggests that this species reached 8 m (26 ft) in maximum length.

The Ankazomihaboka Formation is a Coniacian geologic formation in the Mahajanga Basin of northwestern Madagascar. The formation comprises claystones and sandstones deposited in a fluvial to lacustrine environment. The formation is overlain by the Marovoay Beds and overlies basalt.

Kinkonychelys is an extinct genus of side-necked turtle which existed in Madagascar during the Late Cretaceous period. It contains the single species Kinkonychelys rogersi, named in honor of its discoverer, Raymond R. Rogers. The genus and species are based on UA 9748, a nearly complete skull, which represents the first turtle skull described from the pre-Holocene era in Madagascar. A number of isolated skull and jaw bones have also been assigned to K. rogersi. These specimens were found in rocks of the Maastrichtian-age Maevarano Formation in the Mahajanga Basin of northwestern Madagascar. Another specimen, FMNH PR 2446, is speculated to represent another species, currently known as Kinkonychelys sp., but consensus on its distinction from K. rogersi remains unclear.

UA 8699 is a fossil mammalian tooth from the Cretaceous of Madagascar. A broken lower molar about 3.5 mm (0.14 in) long, it is from the Maastrichtian of the Maevarano Formation in northwestern Madagascar. Details of its crown morphology indicate that it is a boreosphenidan, a member of the group that includes living marsupials and placental mammals. David W. Krause, who first described the tooth in 2001, interpreted it as a marsupial on the basis of five shared characters, but in 2003 Averianov and others noted that all those are shared by zhelestid placentals and favored a close relationship between UA 8699 and the Spanish zhelestid Lainodon. Krause used the tooth as evidence that marsupials were present on the southern continents (Gondwana) as early as the late Cretaceous and Averianov and colleagues proposed that the tooth represented another example of faunal exchange between Africa and Europe at the time.

Several mammals are known from the Mesozoic of Madagascar. The Bathonian Ambondro, known from a piece of jaw with three teeth, is the earliest known mammal with molars showing the modern, tribosphenic pattern that is characteristic of marsupial and placental mammals. Interpretations of its affinities have differed; one proposal places it in a group known as Australosphenida with other Mesozoic tribosphenic mammals from the southern continents (Gondwana) as well as the monotremes, while others favor closer affinities with northern (Laurasian) tribosphenic mammals or specifically with placentals. At least five species are known from the Maastrichtian, including a yet undescribed species known from a nearly complete skeleton that may represent a completely new group of mammals. The gondwanathere Lavanify, known from two teeth, is most closely related to other gondwanatheres found in India and Argentina. Two other teeth may represent another gondwanathere or a different kind of mammal. One molar fragment is one of the few known remains of a multituberculate mammal from Gondwana and another has been interpreted as either a marsupial or a placental.

<span class="mw-page-title-main">Majungasaurinae</span> Extinct subfamily of reptiles

Majungasaurinae is a subfamily of large carnivorous theropods from the Upper Cretaceous, found in Madagascar, India, and France. It is a subgroup within the theropod family Abelisauridae, a Gondwanan clade known for their thick and often horned skulls and vestigial arms. The two subfamilies of Abelisauridae are Carnotaurinae, best known from the South American Carnotaurus, and Majungasaurinae, consisting of Madagascar’s Majungasaurus and its closest relatives. Their ancestors emerged in the Middle Jurassic, and the clade lasted until the Upper Cretaceous.

<i>Vintana</i> Extinct species of mammal

Vintana sertichi is an early gondwanatherian mammal dating from the Late Cretaceous, approximately 66 million years ago. Scientists found the lone fossil, a skull, on Madagascar's west coast in the Maastrichtian Maevarano Formation.

<i>Adalatherium</i> Extinct genus of mammals

Adalatherium is an extinct gondwanatherian that lived on Madagascar during the Maastrichtian stage of the Late Cretaceous. The discovery of the first nearly-complete Adalatherium skeleton from the Maevarano Formation was announced in April 2020.

References

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  2. Weishampel et al., 2004
  3. 1 2 3 Krause et al., 2007b
  4. Depéret, 1896
  5. Sues & Taquet, 1979
  6. Evans, Susan E.; Jones, Marc E. H.; Krause, David W. (2008). "A giant frog with South American affinities from the Late Cretaceous of Madagascar". Proceedings of the National Academy of Sciences of the United States of America . 105 (8): 2951–2956. Bibcode:2008PNAS..105.2951E. doi: 10.1073/pnas.0707599105 . PMC   2268566 . PMID   18287076.
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  9. 1 2 3 4 5 6 7 8 9 Ostrowski, S.A. (2012). "The teleost ichthyofauna from the Late Cretaceous of Madagascar: systematics, distributions, and implications for Gondwanan biogeography" (PDF). Michigan State University. Geological Sciences.
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  12. Murray, Alison M.; Brinkman, Donald B.; Friedman, Matt; Krause, David W. (2023-10-17). "A large, freshwater chanid fish (Ostariophysi: Gonorynchiformes) from the Upper Cretaceous of Madagascar". Journal of Vertebrate Paleontology. doi:10.1080/02724634.2023.2255630. ISSN   0272-4634.
  13. Evans et al., 2014, p.5
  14. O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
  15. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 "83.2 Faritany Majunga, Madagascar; 3. Maevarano Formation," in Weishampel et al., 2004, p.605
  16. "Table 14.1," in Weishampel, et al. (2004). Page 326.
  17. Maidment, Susannah; Norman, David; Barrett, Paul; Upchurch, Paul (2008). "Systematics and phylogeny of Stegosauria (Dinosauria: Ornithischia)". Journal of Systematic Palaeontology. 6 (4): 367–407. Bibcode:2008JSPal...6..367M. doi:10.1017/S1477201908002459. S2CID   85673680.
  18. "Table 13.1," in Weishampel, et al. (2004). Page 270.
  19. 1 2 3 Rogers & Wilson, 2014
  20. Patrick M. O’Connor; Alan H. Turner; Joseph R. Groenke; Ryan N. Felice; Raymond R. Rogers; David W. Krause; Lydia J. Rahantarisoa (2020). "Late Cretaceous bird from Madagascar reveals unique development of beaks". Nature. 588 (7837): 272–276. Bibcode:2020Natur.588..272O. doi:10.1038/s41586-020-2945-x. PMID   33239782. S2CID   227174405.
  21. "Table 3.1," in Weishampel et al., 2004, p.50
  22. "Table 3.1," in Weishampel et al., 2004, p.49
  23. "Table 11.1," in Weishampel et al., 2004, p.211
  24. "Table 11.1," in Weishampel et al., 2004, p.212
  25. Krause et al, 2014
  26. Krause, David W.; Hoffmann, Simone; Werning, Sarah (December 2017). "First postcranial remains of Multituberculata (Allotheria, Mammalia) from Gondwana". Cretaceous Research. 80: 91–100. Bibcode:2017CrRes..80...91K. doi: 10.1016/j.cretres.2017.08.009 .
  27. Krause, David W.; O'Connor, Patrick M.; Rogers, Kristina Curry; Sampson, Scott D.; Buckley, Gregory A.; Rogers, Raymond R. (23 August 2006). "Late Cretaceous terrestrial vertebrates from Madagascar: Implications for Latin American biogeography". Annals of the Missouri Botanical Garden. 93 (2): 178–208. doi:10.3417/0026-6493(2006)93[178:LCTVFM]2.0.CO;2. JSTOR   40035721. S2CID   9166607.
  28. Krause et al., 2020

Bibliography