Andalgalornis
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''Andalgalornis'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of flightless
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
y
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 ...
s of the extinct
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Phorusrhacidae Phorusrhacids, colloquially known as terror birds, are an extinct clade of large carnivorous flightless birds that were one of the largest species of apex predators in South America during the Cenozoic era; their conventionally accepted temporal ...
(often called "terror birds") that lived in
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. The type and only
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 ...
is ''A. steulleti''.


Taxonomy

''Andalgalornis'' is known from an incomplete skeleton and some single bones found from sites in the Entre Ríos and
Catamarca Province Catamarca () is a province of Argentina, located in the northwest of the country. The province had a population of 334,568 as per the , and covers an area of 102,602 km2. Its literacy rate is 95.5%. Neighbouring provinces are (clockwise, fr ...
s of northeast and northwest Argentina. The fossils were uncovered 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 ...
(
Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age. ...
in the SALMA classification)
Ituzaingó Formation The Ituzaingó Formation ( es, Formación Ituzaingó), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive geological formation of Late Miocene (Tortonian, or Huayquerian in the SALMA classification) age in ...
of the
Paraná Basin The Paraná Basin ( pt, Bacia do Paraná, es, Cuenca del Paraná) is a large cratonic sedimentary basin situated in the central-eastern part of South America. About 75% of its areal distribution occurs in Brazil, from Mato Grosso to Rio Grande d ...
.Paraná, Pueblo Brugo to Diamante, Ituzaingó Fm
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
shows the position of ''Andalgalornis'' following a 2015 analysis:


Description

''Andalgalornis'' stood around one and a half meters tall. The subfamily to which the genus belonged, the Patagornithinae, contained species that were of quite slender build; it looked much like the larger phorusrhacid ''
Phorusrhacos ''Phorusrhacos'' ( ) is an extinct genus of giant flightless terror birds that inhabited Argentina during the Miocene epoch. ''Phorusrhacos'' was one of the dominant land predators in South America at the time it existed. It is thought to have ...
'', but it was more elegant and smaller and had a proportionally higher beak, the most massive in proportion to body size of all phorusrhacids.


Paleobiology

Phorusrhacids are though to have been ground predators or scavengers, and have often been considered
apex predators 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 ...
that dominated Cenozoic South America in the absence of mammalian predators, though they did co-exist with some large, carnivorous
borhyaenid Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and o ...
mammals. Earlier hypotheses of phorusrhacid feeding ecology were mainly based on them possessing large skulls with hooked beaks rather than through detailed hypotheses and biomechanical studies, and such studies of their running and predatory adaptations were only tested from the beginning of the 21st century. Alvarenga and Elizabeth Höfling made some general remarks about phorusrhacid habits in a 2003 article. They were flightless, as evidenced by the proportional size of their wings and body mass, and wing-size was more reduced in larger members of the group. They pointed out that the narrowing of the pelvis, upper maxilla, and
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
may have been adaptations for hunting in regions with high vegetation, which would permit greater agility when moving between vertical obstacles. The narrow upper maxilla would also help catching small animals hidden among tree trunks or stones. The large expansions above the eyes formed by the lacrimal bones (similar to what is seen in modern
hawks Hawks are birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica. * The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. This subfamily a ...
) would have protected the eyes against the sun, and enabled keen eyesight, which indicates they hunted by sight in open, sunlit areas, and not shaded forests. A 2010 study by Degrange and colleagues of ''Andalgalornis'', based on
Finite Element Analysis The finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical modeling. Typical problem areas of interest include the traditional fields of structural analysis, heat ...
using CT scans, estimated its
bite force Bite force quotient (BFQ) is a numerical value commonly used to represent the bite force of an animal, while also taking factors like the animal's size into account. The BFQ is calculated as the regression of the quotient In arithmetic, a ...
and stress distribution in its skull, and showed that it had lost a large degree of intracranial immobility (mobility of skull bones in relation to each other), as was also the case for other large phorusrhacids. These researchers interpreted this loss as an adaptation for enhanced rigidity of the skull, and compared to the modern
red-legged seriema 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 paraphyletic circumscriptio ...
and
white-tailed eagle The white-tailed eagle (''Haliaeetus albicilla'') is a very large species of sea eagle widely distributed across temperate Palearctic, Eurasia. Like all eagles, it is a member of the family Accipitridae (or accipitrids) which includes other diur ...
, the skull of the phorusrhacid showed relatively high stress under sideways loadings, but low stress where force was applied up and down, and in simulations of “pullback”. Due to the relative weakness of the skull at the sides and middle, these researchers considered it unlikely that ''Andalgalornis'' engaged in potentially risky behavior that involved using its beak to subdue large, struggling prey. Instead, they suggested that it either fed on smaller prey that could be killed and consumed more safely, by for example swallowing it whole, or that when targeting large prey, it used a series of well-targeted repetitive strikes with the beak, in a "attack-and-retreat" strategy. Struggling prey could also be retained with the feet, despite the lack of sharp talons. A 2012 follow up study by Tambussi and colleagues analyzed the flexion abilities of the neck of ''Andalgalornis'', based on the morphology of its neck vertebrae, finding the neck to be divided into three sections. They concluded that the neck musculature and skeleton of ''Andalgalornis'' was adapted to carrying a large head, and for helping it rise after a maximum downwards strike, and the researchers assumed the same would be true for other large, big-headed phorusrhacids. A 2020 study of phorusrhacid skull morphology by Degrange found that there were two main morphotypes within the group, derived from a seriema-like ancestor; the "Psilopterine Skull Type", which was
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
(more similar to the ancestral type), and the "Terror Bird Skull Type", which included ''Andalgalornis'' and other large members, that was more specialized, with more rigid and stiff skulls. Despite the differences, studies have shown the two types handled prey similarly, while the more rigid skulls and resulting larger bite force of the "Terror Bird" type would have been an adaptation to handling larger prey.


References


Further reading

* Kraglievich, Lucas (1931): Contribución al conocimiento de las aves fósiles de la época araucoentrerriana. ''Physis'' 10: 304–315. rticle in Spanish* Patterson, B. & Kraglievich, Lucas (1960): Sistemática y nomenclatura de las aves fororracoideas del Plioceno Argentino. ''Publicacion del Museo Municipal Ciencias Naturales y Tradicionales de Mar del Plata'' 1: 1-51. rticle in Spanish Patagornithinae Extinct flightless birds Prehistoric bird genera Miocene birds of South America Huayquerian Neogene Argentina Fossils of Argentina Ituzaingó Formation Fossil taxa described in 1960 {{paleo-bird-stub