Andalgalornis
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''Andalgalornis'' is a
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 ...
of flightless
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
y
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 ...
s of the extinct
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Phorusrhacidae 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 ...
(often called "terror birds") that lived in
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 ...
. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
and only
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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 429,556 as per the , and covers an area of 102,602 km2. Its literacy rate is 95.5%. Neighbouring provinces are (clockwise, f ...
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 () 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 Chasicoan and precedes the Montehermosan age. Etymology T ...
in the SALMA classification)
Ituzaingó Formation The Ituzaingó Formation (), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive formation (geology), geological formation of Late Miocene (Tortonian, or Huayquerian in the South American land mammal age, SA ...
of the
Paraná Basin The Paraná Basin (, ) 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 do Sul states. The remainder area is distribu ...
. The following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
shows the position of ''Andalgalornis'' following a 2015 analysis:


Description

Around tall in height, ''Andalgalornis'' is estimated to have weighed around , on the basis of the methodology proposed by Campbell and Marcus (1992). 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 bird, flightless terror birds that inhabited South America during the Miocene epoch. ''Phorusrhacos'' was one of the dominant land predators in South America at the time it existed. It ...
'', 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 thought to have been ground predators or scavengers, and have often been considered
apex predators An apex predator, also known as a top predator or superpredator, 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 hi ...
that dominated Cenozoic South America in the absence of mammalian predators, though they did co-exist with some large, carnivorous borhyaenid 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 (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...
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
hawk Hawks are birds of prey of the family Accipitridae. They are very widely distributed and are found on all continents, except Antarctica. The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks, and others. This ...
s) 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 Finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical models, mathematical modeling. Typical problem areas of interest include the traditional fields of structural ...
using
CT scan A computed tomography scan (CT scan), formerly called computed axial tomography scan (CAT scan), is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers or ...
s, estimated its
bite force Bite force quotient (BFQ) is a numerical value commonly used to represent the bite force of an animal adjusted for its body mass, while also taking factors like the allometry effects. The BFQ is calculated as the regression of the quotient of an ...
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 polyphyletic circumscription ...
and
white-tailed eagle The white-tailed eagle (''Haliaeetus albicilla''), sometimes known as the 'sea eagle', is a large bird of prey, widely distributed across temperate Eurasia. Like all eagles, it is a member of the family Accipitridae (or accipitrids) which also ...
, 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 an "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, an ...
(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. * 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. {{Taxonbar, from=Q760382 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