''Epapatelo'' (meaning "wing") is a genus of

pteranodontian Pteranodontia is an extinct group of ornithocheiroid pterodactyloid pterosaurs that lived during the Late Cretaceous period (Coniacian to Maastrichtian stages) of North America and Africa. They were some of the most advanced pterosaurs, and ...

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the Order (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...

from the late

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...

Mocuio Formation of

Namibe Province Namibe Province is a province of Angola. Under Portuguese rule it was the Moçâmedes District. It has an area of 57,091 km2 and had a 2014 census population of 495,326. The port and city of Moçâmedes is the capital of the province with a ...

Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordina ...

. The genus contains a single species, ''E. otyikokolo'', known from partial left limb bones.

Discovery and naming

Since 2005, the Projecto PaleoAngola carried out excavations at Bentiaba. For the first time in history, remains of pterosaurs were found in Angola. The recovered fossils consist of fourteen specimens of at least eleven individuals. Two of these represent the new taxon ''Epapatelo''. In 2022, the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...

''Epapatelo otyikokolo'' was named and described by Alexandra E. Fernandes, Octávio Mateus, Brian Andres, Michael J. Polcyn, Anne S. Schulp, António Olímpio, Gonçalves and Louis L. Jacobs. The scientific name is derived from Nhaneca, the language of the local nomadic population. The generic name, ''Epapatelo'', means "wing", and the specific name, ''otyikokolo'', means "lizard". 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 seve ...

, MGUAN-PA650 was found in a sandstone layer of the Mocuio Formation, a marine deposit dating from the early

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 interv ...

, between 71.64 and 71.4 million years old. It consists of a partial left

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...

articulated with a left

ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...

. The

paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...

, MGUAN-PA661, represents an articulated left ulna and

radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...

. All of these bones are preserved three-dimensionally, without significant compression. They were recovered from a surface area of fifteen hectares. The plesiosaur ''

Cardiocorax ''Cardiocorax'' is an extinct genus of elasmosaurid known from the Late Cretaceous (early Maastrichtian stage) Mocuio Formation of Namibe Province, southern Angola. It contains a single species, ''Cardiocorax mukulu''. Its length is estimate ...

'' was also found in the Mocuio Formation.

Description

The wingspan of the holotype was estimated at . The paratype would have been a larger individual, at . Some of the material found at Bentiaba suggests an even larger size. The fourth metacarpal of the holotype was extrapolated at about in length. Elsewhere in Angola, a partial manual digit IV has been found that is at least .

Classification

In their phylogenetic analyses, Fernandes ''et al''. (2022) recovered ''Epapatelo'' as the

sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

to ''

Simurghia ''Simurghia'' is a genus nyctosaurid pterosaur from the Ouled Abdoun Basin of Morocco, a basin that dates to the Maastrichtian stage of the Late Cretaceous period, about 66 million years ago. It was published in 2018 by paleontologists Nicholas ...

'' and '' Alcione''. These taxa, along with the

Nyctosauridae Nyctosauridae (meaning "night lizards" or "bat lizards") is a family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly Europe. It was named in 1889 by Henry Alleyne Nicholson and Richard Lyde ...

, comprise the new clade,

Aponyctosauria Pteranodontia is an extinct group of ornithocheiroid pterodactyloid pterosaurs that lived during the Late Cretaceous period (Coniacian to Maastrichtian stages) of North America and Africa. They were some of the most advanced pterosaurs, and ...

References

{{Pterosauria, P. Pteranodontians Late Cretaceous pterosaurs of Africa Fossil taxa described in 2022