Maevarano Formation
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The Maevarano Formation is a
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
sedimentary Sedimentary rocks are types of rock (geology), rock that are formed by the accumulation or deposition of mineral or organic matter, organic particles at Earth#Surface, Earth's surface, followed by cementation (geology), cementation. Sedimentati ...
rock formation A rock formation is an isolated, scenic, or spectacular surface rock outcrop. Rock formations are usually the result of weathering and erosion sculpting the existing rock. The term ''rock formation'' can also refer to specific sediment ...
found in the
Mahajanga Province Mahajanga was a former province of Madagascar that had an area of 150,023 km². It had a population of 1,896,000 (2004). Its capital was Mahajanga, the second largest city in Madagascar. Except for Fianarantsoa, Mahajanga Province bordere ...
of northwestern
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
. It is most likely
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
in age, and records a seasonal,
semiarid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi- ...
environment with
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of wate ...
s that had greatly varying discharges. Notable animal
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s recovered include the
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
''
Majungasaurus ''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 (geology), Period, making it one of the last known non-avian dinosaurs that went ex ...
'', the early
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
''
Vorona ''Vorona'' ( ; Malagasy for "bird", ''V. berivotrensis'', "from Berivotra") is a monotypic genus of prehistoric birds. It was described from fossils found in a Maevarano Formation quarry near the village of Berivotra, Mahajanga Province, Madag ...
'', the
paravian Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the aviala ...
''
Rahonavis ''Rahonavis'' is a genus of bird-like theropods from the Late Cretaceous (Maastrichtian, about 70 mya) of what is now northwestern Madagascar. It is known from a partial skeleton ( UA 8656) found by Catherine Forster and colleagues in Maevaran ...
'', the
titanosaur Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still th ...
ian
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
''
Rapetosaurus ''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. Like other sauropo ...
'', and the giant
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
''
Beelzebufo ''Beelzebufo ampinga'' ( or ) was a particularly large species of prehistoric frog described in 2008. Common names assigned by the popular media include devil frog, devil toad, and the frog from hell. Fossils of ''Beelzebufo'' have been recover ...
''.


Description

The Maevarano Formation is well exposed in the
Mahajanga Basin Mahajanga (French: Majunga) is a city and an administrative district on the northwest coast of Madagascar. The city of Mahajanga (Mahajanga I) is the capital of the Boeny Region. The district (identical to the city) had a population of 220,629 ...
, in particular near the village of Berivotra near the northwestern coast of the island where its
outcrop An outcrop or rocky outcrop is a visible exposure of bedrock or ancient superficial deposits on the surface of the Earth. Features Outcrops do not cover the majority of the Earth's land surface because in most places the bedrock or superficial ...
s have been heavily dissected by
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is distin ...
. At the time it was being deposited, its
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north pol ...
was between 30°S and 25°S as Madagascar drifted northward after splitting from
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
about 88 million years ago. It is composed of three smaller units or
members Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...
. The lowest is the Masorobe Member, which is usually reddish and is at least . Its rocks are mostly poorly sorted coarse-
grained Graining is the practice of imitating wood grain on a non-wood surface, or on relatively undesirable wood surface, in order to give it the appearance of a rare or higher quality wood, thereby increase that surface's aesthetic appeal. Graining was ...
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
s with some finer-grained beds. It is separated by an erosional
disconformity An unconformity is a buried erosional or non-depositional surface separating two rock masses or strata of different ages, indicating that sediment deposition was not continuous. In general, the older layer was exposed to erosion for an interval ...
from the next member, the Anembalemba Member. The lower portion of the Anembalemba Member is fine to coarse
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4). Clays develop plasticity when wet, due to a molecular film of water surrounding the clay par ...
-rich sandstone, whitish or light grey in color, with
cross-bedding In geology, cross-bedding, also known as cross-stratification, is layering within a stratum and at an angle to the main bedding plane. The sedimentary structures which result are roughly horizontal units composed of inclined layers. The origina ...
. 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 Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
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 in some places. Elsewhere, it is replaced by the
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
Berivotra Formation. The Miadana Member is made up of
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too sm ...
,
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
, 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.Rogers et al., 2007 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 The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...
, despite previous reports to that effect.Weishampel et al., 2004 The Berivotra Formation appears to include near its top a
magnetic reversal A geomagnetic reversal is a change in a planet's magnetic field such that the positions of magnetic north and magnetic south are interchanged (not to be confused with geographic north and geographic south). The Earth's field has alternated b ...
, 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 The Cretaceous–Paleogene (K–Pg) boundary, formerly known as the Cretaceous–Tertiary (K–T) boundary, is a geological signature, usually a thin band of rock containing much more iridium than other bands. The K–Pg boundary marks the end of ...
and associated
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
. This suggests that Maevarano organisms also lived shortly before ( geologically speaking) the extinction event.


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 Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Charles Depéret Charles Jean Julien Depéret (25 June 1854 – 18 May 1929) was a French geologist and paleontologist. He was a member of the French Academy of Sciences, the Société géologique de France
.Krause et al., 2007b He briefly described the formation and named two dinosaurs from the remains (''
Titanosaurus ''Titanosaurus'' (; ) is a dubious genus of sauropod dinosaurs, first described by Richard Lydekker in 1877.R. Lydekker. (1877). Notices of new and other Vertebrata from Indian Tertiary and Secondary rocks. ''Records of the Geological Survey of I ...
madagascariensis'' and ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years a ...
crenatissimus'', now ''Majungasaurus'').Depéret, 1896 Similar collections were made throughout the 20th century, yielding mostly fragmentary fossils; one such specimen, a rough partial skull roof, became the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
of supposed
pachycephalosaur Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachyc ...
(bonehead dinosaur) ''Majungatholus'' in 1979.Sues & Taquet, 1979 (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 Stony Brook University (SBU), officially the State University of New York at Stony Brook, is a public research university in Stony Brook, New York. Along with the University at Buffalo, it is one of the State University of New York system's ...
and the
University of Antananarivo University of Antananarivo (french: Université d'Antananarivo) is the primary public university of Madagascar, located in the capital Antananarivo. History The university traces its founding to 16 December 1955 and the formation of the Instit ...
, have greatly expanded knowledge of this formation and the organisms that lived while it was being deposited.


Paleoenvironment

The Maevarano Formation is interpreted as a low-
relief Relief is a sculptural method in which the sculpted pieces are bonded to a solid background of the same material. The term ''relief'' is from the Latin verb ''relevo'', to raise. To create a sculpture in relief is to give the impression that the ...
alluvial plain An alluvial plain is a largely flat landform created by the deposition of sediment over a long period of time by one or more rivers coming from highland regions, from which alluvial soil forms. A floodplain is part of the process, being the sma ...
that over time was covered by a
marine transgression A marine transgression is a geologic event during which sea level rises relative to the land and the shoreline moves toward higher ground, which results in flooding. Transgressions can be caused by the land sinking or by the ocean basins filling ...
. Broad, shallow rivers flowed to the northwest from central highlands; evidence for
debris flow Debris flows are geological phenomena in which water-laden masses of soil and fragmented rock rush down mountainsides, funnel into stream channels, entrain objects in their paths, and form thick, muddy deposits on valley floors. They generally ...
s suggests that the discharges of the rivers varied greatly, with periods of dilute water flow, and periods of rapid erosion dumping
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand an ...
into the channels.
Paleosol In the geosciences, paleosol (''palaeosol'' in Great Britain and Australia) is an ancient soil that formed in the past. The precise definition of the term in geology and paleontology is slightly different from its use in soil science. In geolo ...
s are reddish and include
root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...
casts. The paleosols and other sedimentologic evidence indicate well-drained
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...
s with abundant vegetation adapted to a relatively dry climate, strongly
season A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
al ( rainy and
dry season The dry season is a yearly period of low rainfall, especially in the tropics. The weather in the tropics is dominated by the tropical rain belt, which moves from the northern to the southern tropics and back over the course of the year. The te ...
s) and at times semiarid (not unlike the present climate of the area).


Paleofauna

Animals found in the formation include
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s (including '' Beelzebufo ampinga''),
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tu ...
s,
snake Snakes are elongated, Limbless vertebrate, limbless, carnivore, carnivorous reptiles of the suborder Serpentes . Like all other Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales. Ma ...
s,
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s, at least seven
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
crocodyliforms Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
(including species of ''
Mahajangasuchus ''Mahajangasuchus'' is an extinct genus of crocodyliform which had blunt, conical teeth. The type species, ''M. insignis'', lived during the Late Cretaceous; its fossils have been found in the Maevarano Formation in northern Madagascar. It was a ...
'' and ''
Trematochampsa ''Trematochampsa'' is a dubious extinct genus of crocodyliform from the Late Cretaceous (Coniacian-Santonian age) In Beceten Formation of Niger. Taxonomy The type species, ''T. taqueti'', was described by Eric Buffetaut in 1974. A second specie ...
''),
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
theropods ''
Majungasaurus ''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 (geology), Period, making it one of the last known non-avian dinosaurs that went ex ...
'',
noasaurid Noasauridae is an extinct family of theropod dinosaurs belonging to the group Ceratosauria. They were closely related to the short-armed abelisaurids, although most noasaurids had much more traditional body types generally similar to other ther ...
''
Masiakasaurus ''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 knopfl ...
'', two types of titanosaurian sauropods (''
Rapetosaurus ''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. Like other sauropo ...
'' and '' Vahiny''), and at least five species of bird-like dinosaurs, including ''
Rahonavis ''Rahonavis'' is a genus of bird-like theropods from the Late Cretaceous (Maastrichtian, about 70 mya) of what is now northwestern Madagascar. It is known from a partial skeleton ( UA 8656) found by Catherine Forster and colleagues in Maevaran ...
''. The long ''Majungasaurus'' was likely the
apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
in the terrestrial environment. Crocodyliforms were very diverse and abundant.


Invertebrates


Osteichthyes


Amphibians


Dinosaurs

Indeterminate
Lithostrotia Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of '' Malawisaurus'' and '' Saltasaurus'' ...
remains formerly attributed to
Titanosauridae Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of ''Malawisaurus'' and ''Saltasaurus'' an ...
. Undescribed
Lithostrotia Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch ''et al.'' in 2004 as the most recent common ancestor of '' Malawisaurus'' and '' Saltasaurus'' ...
form. Indeterminate
Enantiornithes The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teeth and cla ...
remains. A rich avifauna with several undescribed taxa are known, including pengornithid enantiornithes and putative omnivoropterygids.


Crocodylomorphs


Squamates


Turtles


Mammals

Mammal remains include an undescribed gondwanathere,Krause et al, 2014 a broken tooth UA 8699, which has been interpreted both as
metatheria Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
n and as
eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tra ...
n, a non-gondwanathere
multituberculate Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
tooth fragment, a non-gondwanathere multituberculate femur, and a yet undescribed mammal known from an articulated skeleton. Some taxa are particularly large sized herbivores, exemplifying the diversity of Mesozoic mammals.Krause et al., 2020


See also

*
Lists of fossiliferous stratigraphic units in Africa These lists of fossiliferous stratigraphic units in Africa enumerate the rock layers which preserve the fossilized remains of ancient life in Africa by the modern countries wherein they are found. Geographical atlas C ...
**
List of fossiliferous stratigraphic units in Madagascar Several fossiliferous stratigraphic units in Madagascar have provided fossils, most notably the Maastrichtian Maevarano Formation and the Jurassic Isalo III Formation. The oldest fossil-bearing formations date to ages before the break-up of Mada ...
*
Geology of Madagascar The geology of Madagascar comprises a variety of rocks of Precambrian age which make up the larger part of the east and centre of the island. They are intruded by basalts and rhyolites of Mesozoic to Cenozoic age. In contrast, the western part of ...


References


Bibliography

* * * * * * * * * {{ISBN, 0-520-24209-2 Geologic formations of Madagascar Upper Cretaceous Series of Africa Cretaceous Madagascar Maastrichtian Stage Sandstone formations Shale formations Alluvial deposits Deltaic deposits Fluvial deposits Lacustrine deposits Paleontology in Madagascar Formations