''Procariama'' is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

phorusrhacid Phorusrhacids, colloquially known as terror birds, are an extinct family of large carnivorous, mostly flightless birds that were among the largest apex predators in South America during the Cenozoic era. Their definitive fossil records range from ...

, which lived from the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

to the

Late Pliocene Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...

(11-2 million years ago) of

Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...

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 preserve ...

s of the animal have been found in six places, in the Cerro Azul and Andalhuala Formations. More specifically in the Andagalá department and in the north of the

Belén department Belén is a department of Catamarca Province in Argentina. The provincial subdivision has a population of about 12,000 inhabitants in an area of , and its capital city is Belén, which is located around 1,465 kilometres from Buenos Aires City ...

of the

Catamarca province Catamarca () is a province of Argentina, located in the northwest of the country. The province had a population of 429,556 as per the , and covers an area of 102,602 km2. Its literacy rate is 95.5%. Neighbouring provinces are (clockwise, f ...

, with a single location in the

La Pampa province La Pampa () is a sparsely populated province of Argentina, located in the Pampas in the center of the country. Neighboring provinces are from the north clockwise San Luis, Córdoba, Buenos Aires, Río Negro, Neuquén and Mendoza. History In ...

. The type and only species, ''Procariama simplex'', is the largest member of the subfamily Psilopterinae.

History of Discovery

The lectotype of ''Procariama'' (MACN-8225) is a partial skeleton consisting of an incomplete

skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...

, a

pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...

proximal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

and

distal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...

parts of the left

femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...

, distal parts of the right

tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These ...

, proximal and distal parts of the right

tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bird bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) a ...

foot The foot (: feet) is an anatomical structure found in many vertebrates. It is the terminal portion of a limb which bears weight and allows locomotion. In many animals with feet, the foot is an organ at the terminal part of the leg made up o ...

bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...

s and the

nail Nail or Nails may refer to: In biology * Nail (anatomy), toughened protective protein-keratin (known as alpha-keratin, also found in hair) at the end of an animal or human digit, such as fingernail * Nail (beak), a plate of hard horny tissue a ...

bearing

toe Toes are the digits of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plantigrade''; ...

bones of the nearly complete left foot, and fragments of the toe bones of the right foot. The genus name was first published in 1914 by Cayetano Rovereto in ''Anales del Museo Nacional de Historia Natural''.

Description

Reaching about in height and in body mass, ''Procariama'' was one of the smaller phorusrhacids. However, it was larger than ''Psilopterus'', '' Paleopsilopterus'', and other members of the subfamily Psilopterinae. ''Procariama'' is quite similar to ''Psilopterus'', but differs from this genus in both size and a more robust build. This more robust build can be attributed to a slight difference in the ratio between the bones of the legs, with the thigh being comparatively shorter in ''Procariama''. Next to this, the wing bones of ''Procariama'' are also proportionally smaller than those of ''Psilopterus''. In the tarsometatarsus, the hypotarsus has two protrusions, one laterally and one medially, in its most proximal portion. These protrusions look like two crests, which is different from the condition in both ''Psilopterus'' and ''Paleopsilopterus''. The genus is very similar to ''Mesembriornis'' and lived during the same geological period, making fossils of the two easily confused. This happened, for example, in the original publication by Rovereto. The genus name of the animal means “before ''

Cariama The red-legged seriema (''Cariama cristata''), also known as the crested cariama and crested seriema, is a mostly predatory terrestrial bird in the seriema family (Cariamidae), included in the Gruiformes in the old polyphyletic circumscription ...

''”, referring to the genus ''Cariama'', the red-legged seriema, which is a genus of small, carnivorous

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

native Native may refer to: People * '' Jus sanguinis'', nationality by blood * '' Jus soli'', nationality by location of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Nat ...

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

. It is also the closest living relative of the Phorusrhachids.

Known material

Three specimens of ''Procariama'' are known in addition to the lectotype. Firstly, there's MACN-6939, consisting of a femur missing its distal end, the distal part of the left tarsometatarsus and a few pedal phalanges. Secondly we have FM-P 14525, an exquisitely preserved and nearly complete skeleton.

Classification

Like all phorusrhacids, ''Procariama'' is part of the order

Cariamiformes Cariamiformes (or Cariamae) is an order of primarily flightless birds that has existed for over 50 million years. The group includes the family Cariamidae (seriemas) and the extinct families such as Phorusrhacidae, Bathornithidae, Idiornithid ...

, the only modern representatives of which are the

seriema The seriemas are the sole living members of the small bird family Cariamidae (the entire family is also referred to as "seriemas"), which is also the only surviving lineage of the order Cariamiformes. Once believed to be related to cranes, they ...

s. ''Procariama'' is traditionally placed in the subfamily Psilopterinae, but in the description of ''Llallawavis scagliai'', it is placed in the subfamily

Mesembriornithinae Phorusrhacids, colloquially known as terror birds, are an extinct family of large carnivorous, mostly flightless birds that were among the largest apex predators in South America during the Cenozoic era. Their definitive fossil records range from ...

as a sister taxon of ''Llallawavis''. In 2024, it was reclassified as a member of the Psilopterinae.Supplementary Information
/ref>

References

External links

Genus Taxonomy
{{Taxonbar, from=Q2361775 Phorusrhacidae Extinct flightless birds Miocene birds of South America Pliocene birds of South America Huayquerian Montehermosan Neogene Argentina Fossils of Argentina Cerro Azul Formation Fossil taxa described in 1914