1830:
4016:
1958:
670:
1754:. The upper two thirds of its upper surface are concave, while the lower third is flatter. The tarsometatarsus has cotylae (two cup-like cavities at the upper end of the shaft) that are almost oval and deeply concave. The lateral cotyla on the outer side is smaller than the medial cotyla on the inner side, and is slightly below it. The intercotylar eminence between the cotylae is well developed and robust, as in other phorusrhacids. Unique to this genus, there is a round
4187:
2070:
143:
1967:
1413:
502:
1739:
1525:
686:
122:
1536:
1785:
4027:
491:
4022:
3358:
4175:
1763:(the "knuckles" of the tarsometatarsus which articulated with the upper part of the toe phalanges) is much bigger than the two other trochlea (second and fourth), and projects much further down, and the fourth trochlea is larger than the second. The fourth trochlea is irregularly quadrangular, which contrasts with the rectangular trochlea of
1649:, this structure was probably also an extension of the lacrimal. The supraorbital ossification fits within a socket formed by a part of the frontal bone that forms the postorbital process, a configuration unknown in other phorusrhacids. The lower margin of the eye socket is formed by a robust jugal bar which is very tall (larger than that of
1627:
1693:, these bones are separated at the midline by a distinct, longitudinal depression running much of their length, and along the back half of the palate, this depression is flanked by portions of the maxillae. The side margin at the back of the maxilla has a sutured contact with the jugal which is well-defined, similar to
1440:'s skull), making it the largest skull of any known bird. The hind end of the skull is 312 mm (12 in) wide. The tarsometatarsus leg bone is 437 mm (17 in) long. The head height was up to 3 m (9.8 ft), while modern seriemas reach 90 cm (3.0 ft) in height. While the weight of
1385:, and while fossils from Europe have been assigned to the group, their classification is disputed. It is unclear where the group originated; both cariamids and phorusrhacids may have arisen in South America, or arrived from elsewhere when southern continents were closer together or when sea levels were lower.
1721:
enters) is almost triangular, uniquely for this genus, and has a blunt upper apex, and it is slightly smaller than the condyle. Above the foramen magnum is a crest-like prominence, vertically extending from the edge of the foramen to the transverse nuchal crest. A fossa (shallow depression) under the
1586:
The nostrils are small, rectangular, and are located in the upper hind corner of the upper beak as in patagornithines (the size and location of the nostrils is unknown in the larger phorusrhacines and brontornithines). The nostrils appear to be longer from front to back than high, though this may be
1664:
are fully fused. This fusion makes it difficult to identify how these bones were part of the skull roof, but the blunt, robust postorbital processes were probably mainly formed by the frontals. On their lower sides, each frontal forms a large depression where a jaw muscle attached. The postorbital
2035:
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
2060:
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
2051:
was adapted to carrying a large head, and for helping it rise from a maximum extension after a 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
1882:
inside the bones, or by using the legs as kicking weapons (like some modern ground birds do), consistent with the large, curved, and sideways compressed claws known in some phorusrhacids. They also suggested future studies could examine whether they could have used their beaks and claws against
1480:
demonstrated the significant difference between the skulls of large and small members of the group. The holotype skull is very massive, and triangular when viewed from above, with the hind portion compressed from top to bottom. The upper beak is very long, exceeding half the total length of the
1811:
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. These researchers pointed out that the
1636:
While the shape of the eye sockets may be slightly affected by compression from top to bottom, it is likely they were low, almost rectangular in shape, with a concave upper margin and a slightly convex lower border. The upper part of the eye socket is delineated by a thick, rounded edge (a
2031:, 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 midline, these researchers considered it unlikely that
1758:
on the medioplantar corner of the lateral cotyla, lower in height than the intercotylar eminence. The middle of the shaft of the tarsometatarsus is irregularly quadrangular, which is different from that of brontornithines, which are rectangular and very wide. The trochlea of the third
1582:
had a similar morphology on the front part of the palate. Much of the upper beak's side is scarred by small, irregular pits, which functioned as nerve exits. The hindmost two thirds of the upper beak are excavated by a prominent furrow, which runs parallel to the margin of the tomium.
636:(toe bone), and some indeterminate fragments. The describers concluded these bones belonged to a single specimen due to being collected together (and with no other fossils being present), because their general preservation (such as color and texture) was similar, and because they were
1607:(an opening in front of the eye socket) is clear despite it being crushed somewhat on both sides. The front border of this opening is approximately level with the hind margin of the nostril, and its lower margin is straight when viewed from the left side. Robust
1816:
could have been adaptations to enable the birds to search for and take smaller animals in tall plant growth or broken terrain. The large expansions above the eyes formed by the lacrimal bones (similar to what is seen in modern
1913:
may not have been as swift as an ostrich, it could clearly run faster than had previously been assumed for large phorusrhacids, based on the long, slender leg-bones, superficially similar to those of the modern, flightless
1499:
is 0.56, based on the distance between the bony nostril and the front tip. In spite of the crushing from top to bottom, the upper beak is high and very robust, though apparently not as high as in patagornithines, such as
3213:
Genise, Jorge F.; Bellosi, Eduardo S.; Cantil, Liliana F.; González, Mirta G.; Puerta, Pablo (2022). "Middle
Miocene climate transition as reflected by changes in ichnofacies and palaeosols from Patagonia, Argentina".
387:
is the largest-known phorusrhacid, 10% larger than its largest relatives known previously. At 716 mm (2.3 ft) long, the holotype skull is the largest known of any bird, and has been likened to the size of a
1898:
shed doubt on the traditional idea that the size and agility of phorusrhacids correlated, with the larger members of the group being more bulky and less adapted for running. The long and slender tarsometatarsus of
1558:(the frontmost bone of the upper jaw) prominently projects as a sharp, downturned hook. Such a strong downwards projection resembles most closely the condition seen in large to medium sized phorusrhacids such as
1688:
The maxillae form an extensive palate, with the side margins being almost parallel for most of the upper beak's length, and the palate becomes wider from the front back to the region of the eye sockets. Like in
640:
consistent with belonging to a large phorusrhacid. The specimen possessed the most complete skull of a large phorusrhacid known at the time. Previously, such skulls were known only from the fragmentary
724:
most similar to taxa that had traditionally been considered phorusrhacines. Features shared with phorusrhacines include that the hind part of the skull is low and compressed from top to bottom, a wide
523:(or "terror bird", a group of large, predatory birds) were discovered by Argentine high school student Guillermo Aguirre-Zabala between two houses, about 100 m (330 ft) from the railroad of
1574:, rather than the weaker projections of the smaller psilopterines. The underside of the upper beak's front portion forms a pair of prominent ridges that are each separated by a groove from the
1866:, 29 m/s (100 km/h; 65 mph). They found these estimates unlikely due to the large body size of these birds, and instead suggested the strength could have been used to break the
1366:
in the family
Cariamidae. While they are the most speciose group within Cariamiformes, the interrelationships between phorusrhacids are unclear due to the incompleteness of their remains.
577:
The specimen became part of the collection of the Museo
Asociación Paleontológica Bariloche in Río Negro, where it was cataloged as specimen BAR 3877-11. Prior to the animal receiving a
3185:
Tonni, Eduardo P.; Carlini, Alfredo A. (2008). "Neogene vertebrates from
Argentine Patagonia: their relationship with the most significant climatic changes". In Rabassa, Jorge (ed.).
453:
show that large phorusrhacids had very rigid and stiff skulls; this indicates they may have swallowed small prey whole or targeted larger prey with repetitive strikes with the beak.
1653:), and flat from side to side. The jugal bone is about four times taller than thick by the lower center of the eyesocket, and its height is greater than in other phorusrhacids.
716:
were recognized at the time (Brontornithinae, Phorusrhacinae, Patagornithinae, Mesembriornithinae, and
Psilopterinae), though their validity had not then been confirmed through
2036:
well-targeted repetitive strikes with the beak, in a "attack-and-retreat" strategy. Struggling prey could also be restrained with the feet, despite the lack of sharp talons.
1804:
mammals. Earlier hypotheses of phorusrhacid feeding ecology were mainly inferred from them having large skulls with hooked beaks rather than through detailed hypotheses and
1681:, parts of which invaded most of the skull roof at the level of the parietal bones. There is a well developed depression behind the zygomatic process, along the side of the
1930:, and that while there were other large predators in South America at the time, they were limited in numbers and not as fast and agile as the phorusrhacids, and the many
447:
suggests that it could run faster than had previously been assumed for large phorusrhacids, and would have been able to chase down small animals. Studies of the related
1685:, which corresponds to a jaw closing muscle. The subtemporal fossa further behind is broad and its back is defined by a blunt, sidewards extension of the nuchal crest.
1578:, or sharp edge of the beak. These ridges are also separated from a broader central portion of the premaxilla by a longitudinal groove (the rostral premaxillar canal).
2142:
The Collón Curá Formation and the
Colloncuran age of South America represent a time when more open environments with reduced plant covering predominated, similar to
1850:(shin bone) of phorusrhacids to determine their speed, but conceded that such estimates can be unreliable even for extant animals. While the tibiotarsal strength of
4119:
765:, the Argentinian paleontologist Federico J. Degrange and colleagues performed a phylogenetic analysis of Phorusrhacidae, wherein they found Phorusrhacinae to be
1821:) 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.
1393:, about 15 million years ago. Since phorusrhacids survived until the Pleistocene, they appear to have been more successful than for example the South American
2351:
4247:
1656:
The frontal bones appear to have been flat on their upper side. The area where the frontals would have contacted the premaxillae is damaged so that their
736:
in that the supratrochlear surface of the lower end is flat. Further comparison was hampered by the lack of anatomical information about phorusrhacines.
2047:, finding the neck to be divided into three sections. By manually manipulating the vertebrae, they concluded that the neck musculature and skeleton of
358:
clarified the anatomy of large phorusrhacids, as these were previously much less well known. The closest living relatives of the phorusrhacids are the
4106:
4222:
3392:
1456:, the skulls of incompletely known large phorusrhacids were reconstructed as scaled up versions of those of smaller, more complete relatives like
1771:, an opening on the lower front side of the tarsometatarsus, has a centralized position, above the upper ends of the third and fourth trochleae.
4242:
3194:
2818:
2567:
1800:
that dominated
Cenozoic South America in the absence of placental mammalian predators, though they did co-exist with some large, carnivorous
653:. The skull of the latter disintegrated during collection (leaving only the tip of the beak), which hampered comparison between phorusrhacid
4015:
1331:
1829:
1424:
with long hind limbs, narrow pelvises, proportionally small wings, and huge skulls, with a tall, long, sideways compressed hooked beak.
612:
The holotype and only known specimen consists of a nearly complete skull which is somewhat crushed from top to bottom, with most of the
383:
with long hind limbs, narrow pelvises, proportionally small wings, and huge skulls, with a tall, long, sideways compressed hooked beak.
1645:, a similar structure has been suggested to be a process of the lacrimal bone, and while the connection between these is not clear in
2550:
Alvarenga, Herculano; Chiappe, Luis; Bertelli, Sara (2011). "Phorusrhacids: The Terror Birds". In Dyke, Gareth; Kaiser, Gary (eds.).
4212:
2179:
1840:
1382:
130:
3274:
2585:"A new Mesembriornithinae (Aves, Phorusrhacidae) provides new insights into the phylogeny and sensory capabilities of terror birds"
4217:
1808:
studies, and such studies of their running and predatory adaptations were only conducted from the beginning of the 21st century.
2748:"The youngest large carnassial bird (Phorusrhacidae, Phorusrhacinae) from South America (Pliocene-Early Pleistocene of Uruguay)"
2088:) outcrops belonging to the Collón Curá Formation in the southeastern corner of Comallo, Patagonia, an area covered in whitish
1944:, in having gigantic heads, very small forelimbs, and very long legs, and thereby had the same kind of meat-eater adaptations.
2101:
1713:) is round with a vertical groove that originates on its upper surface, and reaches almost to the center of the condyle. The
346:
honors the discoverer. The holotype consists of one of the most complete skulls known of a large phorusrhacid, as well as a
4237:
4165:
3010:
2189:
The Collón Curá Formation of
Argentina has provided a wide assemblage of mammals, including at least 24 taxa such as the
2162:, with its well-developed forests with tree-dwelling animals. Forests would then have been restricted to valleys of the
2052:
morphotypes within the group, derived from a seriema-like ancestor. These were the "Psilopterine Skull Type", which was
1476:. These reconstructions highlighted their assumed very tall beaks, round, high eye sockets, and vaulted braincases, but
1339:
2295:. In addition to the mammals that characterize sediments of this age, there are also a few fossils of birds, reptiles,
3385:
2023:. These researchers interpreted this loss as an adaptation for enhanced rigidity of the skull; compared to the modern
840:
320:
about 15 million years ago. The only known specimen was discovered by high school student Guillermo Aguirre-Zabala in
674:
567:
538:
458:
1432:. The holotype skull is about 716 mm (2.3 ft) long from the tip of the beak to the center of the sagittal
4232:
1854:
and an indeterminate large phorusrhacine suggested a speed of 14 m/s (50 km/h; 31 mph), and that of
1266:
2583:
Degrange, Federico J.; Tambussi, Claudia P.; Taglioretti, Matías L.; Dondas, Alejandro; Scaglia, Fernando (2015).
585:
and Sara Bertelli in a short 2006 article. In 2007, Bertelli, Chiappe, and Claudia Tambussi made the specimen the
1922:
would have been able to chase down small mammals and reptiles. In another 2006 news article, Chiappe stated that
574:
himself, and the discovery led him to shift from studying psychology to studying paleontology and Earth science.
142:
2019:(mobility of skull bones in relation to each other), as was also the case for other large phorusrhacids such as
1167:
2427:
1934:
mammals would have provided ample prey. Chiappe stated that phorusrhacids crudely resembled earlier predatory
1599:, the main bone of the upper jaw). Whether the nostrils are connected to each other at the middle (lacking a
1428:
is the largest known phorusrhacid, about 10% larger than the largest phorusrhacids previously known, such as
4252:
4207:
2016:
2000:
1185:
704:
Phorusrhacidae, based on its enormous size, combined with its sideways compressed, strongly hooked beak (or
606:
396:
is thought to have been about 3 m (9.8 ft) tall and exceeded 100 kg (220 lb) in weight.
343:
3105:
from the Middle Miocene Collón Cura Formation and the Decline of Astrapotheres in Southern South America".
755:, each other's closest relatives. While the analysis supported there being five subfamilies, the resulting
4046:
3378:
2158:(adapted for running) and large animals to occur, contrasting with the earlier conditions during the late
815:
1750:
The shaft of the tarsometatarsus is somewhat slender, with an almost rectangular mid-section, similar to
4145:
2254:
972:
740:
469:(adapted for running) and large animals to occur. The formation has provided fossils of a wide range of
265:
2837:
Degrange, Federico J.; Tambussi, Claudia P.; Moreno, Karen; Witmer, Lawrence M.; Wroe, Stephen (2010).
2166:
mountain ranges, with few tree-dwelling species. This change happened progressively during the earlier
1957:
1400:
predators (which disappeared in the Pliocene), and it is possible that they competed ecologically with
528:
400:
differed from other phorusrhacids in features such as the length of its beak, in having a supraorbital
3304:
Echarri, Sebastian; Ulloa-Guaiquin, Karen S.; Aguirrezabala, Guillermo; Forasiepi, Analia M. (2021). "
2212:
4227:
4093:
4026:
3362:
3223:
3052:
2854:
2759:
2712:
2596:
2263:
2105:
1875:
1698:
1203:
1024:
1006:
865:
637:
3975:
3308:(Metatheria, Sparassodonta) from the Collón Cura Formation (Middle Miocene), Río Negro, Argentina".
1660:(joints between them) cannot be identified, but the sutures between the frontals and the nasals and
1611:
form the hind margins of the antorbital fenestrae, and these bones were recessed in relation to the
759:
did not separate Brontornithinae, Phorusrhacinae, and Patagornithinae. In their 2015 description of
4191:
2291:
1931:
1710:
1604:
954:
729:
669:
409:
3660:
2279:
2069:
1697:. The part of the skull roof behind the eye sockets is flat and scarred by the development of the
3945:
3737:
3667:
3607:
3510:
3335:
3286:
3239:
3164:
3122:
2991:
2783:
2728:
2674:
2649:
2622:
2489:
2431:
2321:
2112:
age of the middle Miocene, about 15 million years ago. The formation was accumulated in a broken
2085:
2028:
2024:
1986:
1982:
1903:
instead shows that this bird may have been much swifter than the smaller, more heavyset and slow
883:
799:
717:
571:
137:
4150:
4021:
3759:
3727:
3681:
3520:
3488:
3803:
3653:
3639:
2285:
2273:
1858:
suggested 27 m/s (97 km/h; 60 mph), the latter is greater than that of a modern
1444:
has not been specifically estimated, it is thought to have exceeded 100 kg (220 lb).
4132:
4124:
3545:
3370:
3278:
3190:
3080:
3035:
Tambussi, Claudia P.; de Mendoza, Ricardo; Degrange, Federico J.; Picasso, Mariana B. (2012).
2983:
2934:
2882:
2814:
2775:
2666:
2563:
2481:
1706:
1666:
1588:
1221:
1110:
705:
4137:
3748:
3690:
3143:
Brandoni, Diego; Ruiz, Laureano González; Bucher, Joaquín (2020). "Evolutive implications of
1619:(main bones of the forehead). The antorbital fenestra is proportionally smaller than that of
3811:
3325:
3317:
3270:
3231:
3156:
3114:
3070:
3060:
2973:
2924:
2916:
2872:
2862:
2806:
2767:
2720:
2658:
2612:
2604:
2555:
2525:
2471:
2423:
2117:
2081:
1989:), and hypothetical up-and-downwards range of movement of the neck in the same genus (right)
1674:
1657:
1630:
1468:, as exemplified by a frequently reproduced 1895 sketch of the destroyed skull of the large
1239:
598:
339:
3878:
3774:
2039:
A 2012 follow-up study by Tambussi and colleagues analyzed the flexibility of the neck of
1966:
1743:
1738:
1421:
1412:
1135:
725:
629:
582:
578:
380:
347:
2325:
3227:
3056:
2858:
2763:
2716:
2600:
501:
4084:
3834:
3786:
3568:
3075:
3036:
2929:
2877:
2838:
2230:
2221:
2194:
2183:
2175:
2121:
2113:
2056:(more similar to the ancestral type), and the "Terror Bird Skull Type", which included
2044:
1797:
1714:
1682:
1670:
1390:
1351:
1347:
780:
413:
317:
220:
207:
30:
2801:
Tambussi, Claudia P.; Degrange, Federico J. (2013). "Neogene Birds of South America".
1524:
685:
4201:
3991:
3961:
3915:
3887:
3717:
3630:
3616:
3478:
3454:
3439:
3402:
3339:
3243:
3168:
3126:
2732:
2700:
2584:
2412:"A new phorusrhacid (Aves: Cariamae) from the middle Miocene of Patagonia, Argentina"
2411:
2235:
2159:
2053:
2012:
1977:
1940:
1846:
In 2005, Rudemar Ernesto Blanco and Washington W. Jones examined the strength of the
1661:
1608:
1512:
1483:
1397:
1359:
1355:
924:
709:
690:
625:
621:
449:
428:
194:
72:
3290:
3258:
2995:
2626:
2530:
2513:
2435:
4179:
3894:
3857:
3819:
3584:
2905:"Terror birds on the run: a mechanical model to estimate its maximum running speed"
2904:
2771:
2493:
2459:
2239:
2226:
2217:
2200:
2093:
1975:
Stress distribution in bird skulls during various movements, including the related
1805:
1638:
1616:
1506:
1433:
766:
649:
633:
602:
520:
401:
351:
305:
2724:
2108:
from different areas of the Collón Curá Formation, it is estimated to date to the
628:
behind the eye sockets are missing. The specimen also includes an associated left
3321:
3235:
3118:
3065:
2867:
2608:
1641:), the hind part of which appears to overhang downward as seen from the side. In
1615:(that formed the lower edge of the eye socket) and the outer side margin of each
4078:
3954:
3931:
3843:
3530:
3433:
2810:
2245:
2109:
1915:
1879:
1847:
1718:
1705:, and low, which gives it a rectangular appearance when viewed from behind. The
1678:
1458:
1394:
1374:
761:
752:
713:
643:
613:
462:
417:
372:
121:
47:
34:
3160:
2100:
had not been adequately determined, but compared with other fossil beds of the
1784:
3924:
3901:
3795:
3701:
3554:
3499:
2559:
2206:
2163:
2136:
2008:
1905:
1888:
1801:
1793:
1760:
1592:
1555:
1535:
1464:
474:
465:
age of South America, when open environments predominated, which allowed more
440:
405:
92:
57:
4069:
2779:
2670:
1603:
as in other phorusrhacids) is not discernible. The quadrangular shape of the
553:
540:
3984:
3577:
2296:
2250:
2190:
2155:
2143:
2128:
2124:
2120:
that were disconnected from each other. The formation is composed mainly of
1927:
1884:
1867:
1612:
1401:
1370:
1343:
756:
739:
The Brazilian paleontologist Herculano Alvarenga and colleagues published a
617:
532:
490:
466:
436:
424:
367:
325:
154:
97:
41:
3084:
2987:
2938:
2920:
2886:
2485:
581:, the specimen was reported and discussed by the Argentine paleontologists
439:
mammalian predators, though they did co-exist with some large, carnivorous
3357:
3282:
2011:
and stress distribution in its skull. They found its bite force to be 133
1587:
exaggerated by crushing, and their hind margin is formed by the maxillary
4063:
3646:
3592:
3421:
2167:
2151:
2147:
2132:
2097:
1995:
A 2010 study by Degrange and colleagues of the medium-sized phorusrhacid
1935:
1755:
1709:(the rounded prominence at the back of the head which contacted with the
1545:
1378:
1363:
1327:
586:
510:
432:
392:'s skull. The tarsometatarsus leg bone is 437 mm (17 in) long.
359:
329:
174:
87:
82:
67:
62:
52:
3330:
2701:"A revision of skull morphology in Phorusrhacidae (Aves, Cariamiformes)"
2678:
2662:
1909:. In a 2006 news article about the discovery, Chiappe stated that while
4111:
3864:
3710:
2978:
2953:
2787:
2747:
2617:
2259:
2004:
1863:
1859:
1835:
1768:
1596:
1335:
769:(an unnatural grouping). The following cladogram shows the position of
678:
524:
321:
309:
102:
77:
3275:
10.1002/(SICI)1098-2345(1998)45:4<317::AID-AJP1>3.0.CO;2-Z
2839:"Mechanical analysis of feeding behavior in the extinct "terror bird"
2805:. Springer Briefs in Earth System Sciences. Springer. pp. 59–86.
3415:
3257:
Kay, Richard Frederick; Johnson, Derek; Meldrum, Don Jeffrey (1998).
3147:(Xenarthra, Megatheriinae) from the Miocene of Patagonia Argentina".
2268:
1813:
1600:
1575:
1342:, and some bird groups around the world developed a tendency towards
701:
470:
164:
4040:
2476:
2096:
had only been preliminarily studied at the time, and the age of the
1626:
1358:, and the Phorusrhacidae. Phorusrhacids are an extinct group within
1373:(when the continent was an isolated island) and survived until the
3427:
2068:
1871:
1828:
1818:
1783:
1737:
1625:
1437:
1411:
684:
668:
594:
389:
301:
4098:
2647:
Ksepka, Daniel T. (2014). "Flights of fancy in avian evolution".
2171:
2089:
654:
566:). The outcrops where the specimen was discovered belong to the
313:
184:
4044:
3452:
3374:
2514:"Systematic revision of the Phorusrhacidae (Aves: Ralliformes)"
2131:
that were accumulated in continental environments ranging from
1436:
at the upper back of the head (a size likened to the size of a
3259:"A new pitheciin primate from the middle Miocene of Argentina"
1717:(the large opening at the base of the skull through which the
1404:
predators that entered from North America in the Pleistocene.
696:
In their 2007 description, Bertelli and colleagues classified
416:(the large opening at the base of the skull through which the
3101:
Kramarz, Alejandro; Garrido, Alberto; Bond, Mariano (2019). "
2410:
Bertelli, Sara; Chiappe, Luis M.; Tambussi, Claudia (2007).
1894:
According to Chiappe and Bertelli in 2006, the discovery of
2952:
Hudson, Penny E.; Corr, Sandra A.; Wilson, Alan M. (2012).
1792:
Phorusrhacids are thought to have been ground predators or
632:(lower leg bone of birds), a small upper portion of a foot
423:
Phorusrhacids are thought to have been ground predators or
2015:
at the bill tip, and showed it had lost a large degree of
1677:). The postorbital process contains scars left by massive
1362:, the only living members of which are the two species of
2746:
Tambussi, Claudia; Ubilla, Martín; Perea, Daniel (1999).
2428:
10.1671/0272-4634(2007)27[409:ANPACF]2.0.CO;2
708:, the part of the jaws that formed the beak), and convex
1377:. They also appeared in North America at the end of the
3037:"Flexibility along the neck of the Neogene terror bird
2909:
Proceedings of the Royal Society B: Biological Sciences
2803:
South American and Antarctic Continental Cenozoic Birds
2903:
Blanco, Rudemar Ernesto; Jones, Washington W. (2005).
2512:
Alvarenga, Herculano M.F.; Höfling, Elizabeth (2003).
4163:
2507:
2505:
2503:
1870:
of medium-sized mammals, the size, for example, of a
1369:
Phorusrhacids were present in South America from the
3972:
3942:
3912:
3875:
3831:
3783:
3771:
3755:
3744:
3733:
3723:
3678:
3627:
3604:
3565:
3542:
3526:
3516:
3506:
3495:
3484:
3474:
1495:. The ratio between the upper beak and the skull of
4053:
3971:
3941:
3911:
3874:
3830:
3782:
3770:
3700:
3677:
3626:
3603:
3564:
3541:
3465:
3187:
The Late Cenozoic of Patagonia and Tierra del Fuego
3011:"Huge "terror bird" fossil discovered in Patagonia"
2174:landscapes would have happened simultaneously with
732:, and a tarsometatarsus that is similar to that of
2694:
2692:
2690:
2688:
2545:
2543:
2541:
3216:Palaeogeography, Palaeoclimatology, Palaeoecology
3138:
3136:
3096:
3094:
2405:
2403:
2401:
2399:
2397:
2395:
2393:
2326:"Fossil found of a Big Bird Kermit wouldn't like"
2073:The holotype was found near the Comallo railroad.
443:mammals. The long and slender tarsometatarsus of
2391:
2389:
2387:
2385:
2383:
2381:
2379:
2377:
2375:
2373:
1878:. This strength could be used for accessing the
597:name refers to a spirit in the mythology of the
2962:): spatio-temporal and kinetic characteristics"
2352:"Teen finds fossil skull of biggest bird known"
2186:event which had a drying effect on continents.
712:(the top of the upper beak). Five phorusrhacid
338:in 2007. The genus name references a spirit in
1746:lower leg bone; the upper side is on the left.
316:, which lived in what is now Argentina in the
3386:
2832:
2830:
1669:, and these two projections enclose a narrow
601:of Patagonia which is represented as a giant
8:
1812:narrowing of the pelvis, upper maxilla, and
1701:. The occipital table is very wide, like in
1665:process is separated narrowly from a robust
1548:skull from the right (top) and back (bottom)
657:of different sizes, until the discovery of
527:, a small village in the north-west of the
4041:
3827:
3779:
3462:
3449:
3393:
3379:
3371:
3208:
3206:
3180:
3178:
1595:(a projection from the nasals towards the
120:
20:
3329:
3074:
3064:
2977:
2928:
2876:
2866:
2642:
2640:
2638:
2636:
2616:
2529:
2475:
2458:Chiappe, Luis M.; Bertelli, Sara (2006).
1981:(left, A-C, the other skulls belong to a
473:, with a few fossils of birds, reptiles,
2898:
2896:
2460:"Skull morphology of giant terror birds"
2453:
2451:
2449:
2447:
2445:
4170:
2308:
1722:condyle is not visible, differing from
370:Phorusrhacinae, along with for example
3189:. Elsevier Science. pp. 269–278.
3009:Joyce, Christopher (25 October 2006).
2954:"High speed galloping in the cheetah (
2316:
2314:
2312:
743:of Phorusrhacidae in 2011 that found
412:, and in having an almost triangular
7:
2350:Hotz, Robert Lee (26 October 2006).
2154:. The open environment allowed more
1472:, which was itself based on that of
1332:extinction of the non-bird dinosaurs
677:outcrops; the holotype was found in
2170:stage. The transition towards more
1158:
1151:
1126:
1101:
1076:
997:
945:
938:
931:
856:
831:
806:
794:
787:
776:
689:Comparison between skulls of large
4248:Prehistoric birds of South America
2752:Journal of Vertebrate Paleontology
2705:Journal of Vertebrate Paleontology
2589:Journal of Vertebrate Paleontology
2416:Journal of Vertebrate Paleontology
535:region of Argentina (coordinates:
14:
2180:Middle Miocene Climate Transition
2043:, based on the morphology of its
1841:Florida Museum of Natural History
1796:, and have often been considered
1383:Great American Biotic Interchange
427:, and have often been considered
408:) that fits into a socket of the
131:Fukui Prefectural Dinosaur Museum
4185:
4173:
4025:
4020:
4014:
3356:
2135:(deposited by running water) to
2116:system characterized by several
1965:
1956:
1534:
1523:
500:
489:
435:South America in the absence of
141:
3263:American Journal of Primatology
2966:Journal of Experimental Biology
2531:10.1590/S0031-10492003000400001
4223:Miocene birds of South America
3149:Journal of Mammalian Evolution
2772:10.1080/02724634.1999.10011154
2699:Degrange, Federico J. (2020).
2102:South American land mammal age
1926:would have been as quick as a
1788:Restoration of a feeding group
1:
4243:Fossil taxa described in 2007
2843:(Gruiformes: Phorusrhacidae)"
2725:10.1080/02724634.2020.1848855
1918:. The article suggested that
1883:well-armored mammals such as
1730:, whose fossae are distinct.
1491:, and is longer than that of
773:following the 2015 analysis:
624:preserved, while most of the
332:of the new genus and species
3322:10.5710/AMGH.06.08.2021.3439
3236:10.1016/j.palaeo.2022.110932
3119:10.5710/AMGH.15.07.2019.3258
3066:10.1371/journal.pone.0037701
2958:) and the racing greyhound (
2868:10.1371/journal.pone.0011856
2609:10.1080/02724634.2014.912656
1742:Sketch of the holotype left
1340:evolutionary diversification
2811:10.1007/978-94-007-5467-6_7
2146:and temperate to warm, dry
1839:mounted in a kicking pose,
841:Mesembriornis milneedwardsi
720:, and the describers found
366:was found to belong in the
4269:
3161:10.1007/s10914-019-09469-6
3145:Megathericulus patagonicus
2518:Papéis Avulsos de Zoologia
1833:Reconstructed skeleton of
1452:Prior to the discovery of
1420:Phorusrhacids were large,
1267:Paraphysornis brasiliensis
519:Around 2004, fossils of a
404:(a rounded edge above the
379:Phorusrhacids were large,
312:group of large, predatory
129:Reconstructed skeleton in
4011:
3461:
3448:
3410:
2560:10.1002/9781119990475.ch7
2178:that corresponded to the
1262:
1235:
1217:
1199:
1181:
1163:
1156:
1149:
1131:
1124:
1106:
1099:
1081:
1074:
1020:
1002:
995:
968:
950:
943:
936:
929:
879:
861:
854:
836:
829:
811:
804:
792:
785:
271:
264:
138:Scientific classification
136:
128:
119:
23:
4213:Extinct flightless birds
3405:and their extinct allies
1862:, approaching that of a
1416:Size compared to a human
1389:itself lived during the
1168:Phorusrhacos longissimus
4218:Prehistoric bird genera
3039:Andalgalornis steulleti
2841:Andalgalornis steulleti
2356:Pittsburgh Post-Gazette
2017:intracranial immobility
2001:Finite Element Analysis
1948:Skull and neck function
1186:Andalgalornis steulleti
609:honors its discoverer.
554:41.033167°S 70.258250°W
513:skull in left side view
461:, and lived during the
3306:Cladosictis patagonica
3041:(Aves Phorusrhacidae)"
2921:10.1098/rspb.2005.3133
2139:(deposited by lakes).
2074:
1843:
1789:
1767:. The distal vascular
1747:
1633:
1417:
816:Mesembriornis incertus
693:
682:
559:-41.033167; -70.258250
16:Extinct genus of birds
4146:Paleobiology Database
2072:
1832:
1787:
1741:
1629:
1554:The front end of the
1415:
973:Psilopterus bachmanni
741:phylogenetic analysis
688:
675:Collón Curá Formation
672:
572:prepared the specimen
568:Collón Curá Formation
459:Collón Curá Formation
350:lower leg bone and a
4238:Fossils of Argentina
3367:at Wikimedia Commons
2554:. pp. 187–208.
2106:radioisotopic dating
1699:temporal musculature
1346:; this included the
1204:Andrewsornis abbotti
1025:Psilopterus lemoinei
1007:Psilopterus colzecus
866:Llallawavis scagliai
308:("terror bird"), an
3228:2022PPP...59410932G
3057:2012PLoSO...737701T
2915:(1574): 1769–1773.
2859:2010PLoSO...511856D
2764:1999JVPal..19..404T
2717:2020JVPal..40E8855D
2663:10.1511/2014.106.36
2601:2015JVPal..35E2656D
2324:(31 October 2006).
2322:Wilford, John Noble
1711:first neck vertebra
1605:antorbital fenestra
1086:Kelenken guillermoi
955:Psilopterus affinis
730:postorbital process
700:as a member of the
591:Kelenken guillermoi
550: /
410:postorbital process
354:. The discovery of
342:mythology, and the
335:Kelenken guillermoi
328:, and was made the
324:, in the region of
278:Kelenken guillermoi
3946:Mesembriornithinae
3608:Eleutherornithidae
2979:10.1242/jeb.066720
2650:American Scientist
2330:The New York Times
2086:volcanic eruptions
2084:(rocks ejected by
2080:was discovered in
2075:
2029:white-tailed eagle
2025:red-legged seriema
1987:white-tailed eagle
1983:red-legged seriema
1844:
1790:
1748:
1634:
1418:
884:Procariama simplex
800:Mesembriornithinae
718:cladistic analysis
694:
683:
529:Río Negro Province
457:is known from the
257:K. guillermoi
4233:Neogene Argentina
4161:
4160:
4133:Open Tree of Life
4047:Taxon identifiers
4038:
4037:
4034:
4033:
4009:
4008:
4005:
4004:
4001:
4000:
3546:Ameghinornithidae
3361:Media related to
3196:978-0-08-055889-9
2972:(14): 2425–2434.
2820:978-94-007-5466-9
2569:978-1-119-99047-5
1876:Thomson's gazelle
1707:occipital condyle
1667:zygomatic process
1481:skull, unlike in
1326:During the early
1323:
1322:
1314:
1313:
1305:
1304:
1296:
1295:
1287:
1286:
1278:
1277:
1251:
1250:
1222:Patagornis marshi
1111:Devincenzia pozzi
1063:
1062:
1054:
1053:
1045:
1044:
1036:
1035:
984:
983:
913:
912:
904:
903:
895:
894:
570:. Aguirre-Zabala
293:
292:
245:
4260:
4190:
4189:
4188:
4178:
4177:
4176:
4169:
4154:
4153:
4141:
4140:
4128:
4127:
4115:
4114:
4102:
4101:
4089:
4088:
4087:
4074:
4073:
4072:
4042:
4029:
4024:
4018:
3974:
3944:
3914:
3877:
3833:
3828:
3812:Paleopsilopterus
3785:
3780:
3773:
3757:
3746:
3735:
3725:
3680:
3629:
3606:
3567:
3544:
3528:
3518:
3508:
3497:
3486:
3476:
3463:
3450:
3395:
3388:
3381:
3372:
3360:
3344:
3343:
3333:
3301:
3295:
3294:
3254:
3248:
3247:
3210:
3201:
3200:
3182:
3173:
3172:
3140:
3131:
3130:
3098:
3089:
3088:
3078:
3068:
3032:
3026:
3025:
3023:
3021:
3006:
3000:
2999:
2981:
2960:Canis familiaris
2956:Acinonyx jubatus
2949:
2943:
2942:
2932:
2900:
2891:
2890:
2880:
2870:
2834:
2825:
2824:
2798:
2792:
2791:
2743:
2737:
2736:
2696:
2683:
2682:
2644:
2631:
2630:
2620:
2580:
2574:
2573:
2552:Living Dinosaurs
2547:
2536:
2535:
2533:
2509:
2498:
2497:
2479:
2455:
2440:
2439:
2407:
2368:
2367:
2365:
2363:
2347:
2341:
2340:
2338:
2336:
2318:
2213:Paraeucinepeltus
2065:Paleoenvironment
2007:, estimated its
1969:
1960:
1780:Feeding and diet
1673:(opening at the
1631:Life restoration
1544:Sketches of the
1538:
1527:
1422:flightless birds
1240:Physornis fortis
1159:
1152:
1127:
1102:
1077:
998:
946:
939:
932:
857:
832:
807:
795:
788:
777:
599:Tehuelche people
565:
564:
562:
561:
560:
555:
551:
548:
547:
546:
543:
504:
493:
381:flightless birds
352:phalanx toe bone
280:
276:
239:
232:
219:
206:
146:
145:
124:
114:
44:
29:Temporal range:
21:
4268:
4267:
4263:
4262:
4261:
4259:
4258:
4257:
4198:
4197:
4196:
4186:
4184:
4174:
4172:
4164:
4162:
4157:
4149:
4144:
4136:
4131:
4123:
4118:
4110:
4105:
4097:
4092:
4083:
4082:
4077:
4068:
4067:
4062:
4049:
4039:
4030:
4019:
3997:
3967:
3937:
3907:
3879:Patagornithinae
3870:
3826:
3775:Phorusrhacoidea
3766:
3696:
3673:
3622:
3599:
3560:
3537:
3457:
3444:
3406:
3399:
3353:
3348:
3347:
3303:
3302:
3298:
3256:
3255:
3251:
3212:
3211:
3204:
3197:
3184:
3183:
3176:
3142:
3141:
3134:
3100:
3099:
3092:
3034:
3033:
3029:
3019:
3017:
3008:
3007:
3003:
2951:
2950:
2946:
2902:
2901:
2894:
2836:
2835:
2828:
2821:
2800:
2799:
2795:
2745:
2744:
2740:
2711:(6): e1848855.
2698:
2697:
2686:
2646:
2645:
2634:
2582:
2581:
2577:
2570:
2549:
2548:
2539:
2511:
2510:
2501:
2477:10.1038/443929a
2457:
2456:
2443:
2409:
2408:
2371:
2361:
2359:
2349:
2348:
2344:
2334:
2332:
2320:
2319:
2310:
2305:
2264:Proteropithecia
2176:climate changes
2067:
1993:
1992:
1991:
1990:
1972:
1971:
1970:
1962:
1961:
1950:
1827:
1782:
1777:
1744:tarsometatarsus
1736:
1552:
1551:
1550:
1549:
1541:
1540:
1539:
1530:
1529:
1528:
1450:
1410:
1324:
1315:
1306:
1297:
1288:
1279:
1252:
1136:Titanis walleri
1064:
1055:
1046:
1037:
985:
914:
905:
896:
726:occipital table
667:
638:morphologically
630:tarsometatarsus
583:Luis M. Chiappe
579:scientific name
558:
556:
552:
549:
544:
541:
539:
537:
536:
517:
516:
515:
514:
507:
506:
505:
496:
495:
494:
483:
431:that dominated
348:tarsometatarsus
289:
282:
274:
273:
260:
238:
230:
217:
204:
140:
115:
113:
112:
111:
110:
105:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
39:
38:
27:
17:
12:
11:
5:
4266:
4264:
4256:
4255:
4253:Phorusrhacinae
4250:
4245:
4240:
4235:
4230:
4225:
4220:
4215:
4210:
4208:Phorusrhacidae
4200:
4199:
4195:
4194:
4182:
4159:
4158:
4156:
4155:
4142:
4129:
4116:
4103:
4090:
4075:
4059:
4057:
4051:
4050:
4045:
4036:
4035:
4032:
4031:
4013:
4012:
4010:
4007:
4006:
4003:
4002:
3999:
3998:
3996:
3995:
3988:
3980:
3978:
3976:Physornithinae
3969:
3968:
3966:
3965:
3958:
3950:
3948:
3939:
3938:
3936:
3935:
3928:
3920:
3918:
3909:
3908:
3906:
3905:
3898:
3891:
3883:
3881:
3872:
3871:
3869:
3868:
3861:
3854:
3847:
3839:
3837:
3835:Phorusrhacinae
3825:
3824:
3816:
3808:
3800:
3791:
3789:
3787:Phorusrhacidae
3777:
3768:
3767:
3765:
3764:
3753:
3742:
3731:
3721:
3714:
3706:
3704:
3698:
3697:
3695:
3694:
3686:
3684:
3675:
3674:
3672:
3671:
3664:
3657:
3650:
3643:
3635:
3633:
3624:
3623:
3621:
3620:
3612:
3610:
3601:
3600:
3598:
3597:
3589:
3581:
3573:
3571:
3569:Bathornithidae
3562:
3561:
3559:
3558:
3550:
3548:
3539:
3538:
3536:
3535:
3524:
3514:
3504:
3493:
3482:
3471:
3469:
3467:incertae sedis
3459:
3458:
3453:
3446:
3445:
3443:
3442:
3436:
3430:
3424:
3418:
3411:
3408:
3407:
3400:
3398:
3397:
3390:
3383:
3375:
3369:
3368:
3352:
3351:External links
3349:
3346:
3345:
3296:
3269:(4): 317–336.
3249:
3202:
3195:
3174:
3155:(3): 445–460.
3132:
3103:Astrapotherium
3090:
3027:
3001:
2944:
2892:
2826:
2819:
2793:
2758:(2): 404–406.
2738:
2684:
2632:
2595:(2): e912656.
2575:
2568:
2537:
2499:
2441:
2422:(2): 409–419.
2369:
2342:
2307:
2306:
2304:
2301:
2231:Astrapotherium
2222:Protypotherium
2195:Megathericulus
2184:global cooling
2122:volcaniclastic
2066:
2063:
2045:neck vertebrae
1974:
1973:
1964:
1963:
1955:
1954:
1953:
1952:
1951:
1949:
1946:
1826:
1823:
1798:apex predators
1781:
1778:
1776:
1773:
1735:
1732:
1715:foramen magnum
1683:squamosal bone
1671:temporal fossa
1609:lacrimal bones
1543:
1542:
1533:
1532:
1531:
1522:
1521:
1520:
1519:
1518:
1449:
1446:
1409:
1406:
1391:middle Miocene
1352:Dromornithidae
1348:Gastornithidae
1321:
1320:
1317:
1316:
1312:
1311:
1308:
1307:
1303:
1302:
1299:
1298:
1294:
1293:
1290:
1289:
1285:
1284:
1281:
1280:
1276:
1275:
1272:
1271:
1261:
1258:
1257:
1254:
1253:
1249:
1248:
1245:
1244:
1234:
1231:
1230:
1227:
1226:
1216:
1213:
1212:
1209:
1208:
1198:
1195:
1194:
1191:
1190:
1180:
1177:
1176:
1173:
1172:
1162:
1157:
1155:
1150:
1148:
1145:
1144:
1141:
1140:
1130:
1125:
1123:
1120:
1119:
1116:
1115:
1105:
1100:
1098:
1095:
1094:
1091:
1090:
1080:
1075:
1073:
1070:
1069:
1066:
1065:
1061:
1060:
1057:
1056:
1052:
1051:
1048:
1047:
1043:
1042:
1039:
1038:
1034:
1033:
1030:
1029:
1019:
1016:
1015:
1012:
1011:
1001:
996:
994:
991:
990:
987:
986:
982:
981:
978:
977:
967:
964:
963:
960:
959:
949:
944:
942:
937:
935:
930:
928:
920:
919:
916:
915:
911:
910:
907:
906:
902:
901:
898:
897:
893:
892:
889:
888:
878:
875:
874:
871:
870:
860:
855:
853:
850:
849:
846:
845:
835:
830:
828:
825:
824:
821:
820:
810:
805:
803:
793:
791:
786:
784:
781:Phorusrhacidae
775:
666:
663:
616:, skull roof,
509:
508:
499:
498:
497:
488:
487:
486:
485:
484:
482:
479:
429:apex predators
414:foramen magnum
318:middle Miocene
291:
290:
283:
269:
268:
262:
261:
253:
251:
247:
246:
228:
224:
223:
221:Phorusrhacinae
215:
211:
210:
208:Phorusrhacidae
202:
198:
197:
192:
188:
187:
182:
178:
177:
172:
168:
167:
162:
158:
157:
152:
148:
147:
134:
133:
126:
125:
117:
116:
108:
107:
106:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
45:
31:Middle Miocene
28:
15:
13:
10:
9:
6:
4:
3:
2:
4265:
4254:
4251:
4249:
4246:
4244:
4241:
4239:
4236:
4234:
4231:
4229:
4226:
4224:
4221:
4219:
4216:
4214:
4211:
4209:
4206:
4205:
4203:
4193:
4183:
4181:
4171:
4167:
4152:
4147:
4143:
4139:
4134:
4130:
4126:
4121:
4117:
4113:
4108:
4104:
4100:
4095:
4091:
4086:
4080:
4076:
4071:
4065:
4061:
4060:
4058:
4056:
4052:
4048:
4043:
4028:
4023:
4017:
3994:
3993:
3992:Paraphysornis
3989:
3987:
3986:
3982:
3981:
3979:
3977:
3970:
3964:
3963:
3962:Mesembriornis
3959:
3957:
3956:
3952:
3951:
3949:
3947:
3940:
3934:
3933:
3929:
3927:
3926:
3922:
3921:
3919:
3917:
3916:Psilopterinae
3910:
3904:
3903:
3899:
3897:
3896:
3892:
3890:
3889:
3888:Andalgalornis
3885:
3884:
3882:
3880:
3873:
3867:
3866:
3862:
3860:
3859:
3855:
3853:
3852:
3848:
3846:
3845:
3841:
3840:
3838:
3836:
3829:
3822:
3821:
3817:
3814:
3813:
3809:
3806:
3805:
3801:
3798:
3797:
3793:
3792:
3790:
3788:
3781:
3778:
3776:
3769:
3762:
3761:
3754:
3751:
3750:
3743:
3740:
3739:
3732:
3730:
3729:
3722:
3720:
3719:
3715:
3713:
3712:
3708:
3707:
3705:
3703:
3699:
3693:
3692:
3688:
3687:
3685:
3683:
3676:
3670:
3669:
3665:
3663:
3662:
3658:
3656:
3655:
3651:
3649:
3648:
3644:
3642:
3641:
3637:
3636:
3634:
3632:
3631:Idiornithidae
3625:
3619:
3618:
3617:Eleutherornis
3614:
3613:
3611:
3609:
3602:
3595:
3594:
3590:
3587:
3586:
3582:
3580:
3579:
3575:
3574:
3572:
3570:
3563:
3557:
3556:
3552:
3551:
3549:
3547:
3540:
3533:
3532:
3525:
3523:
3522:
3515:
3513:
3512:
3505:
3502:
3501:
3494:
3491:
3490:
3483:
3481:
3480:
3479:Elaphrocnemus
3473:
3472:
3470:
3468:
3464:
3460:
3456:
3455:Cariamiformes
3451:
3447:
3441:
3440:Cariamiformes
3437:
3435:
3431:
3429:
3425:
3423:
3419:
3417:
3413:
3412:
3409:
3404:
3396:
3391:
3389:
3384:
3382:
3377:
3376:
3373:
3366:
3365:
3359:
3355:
3354:
3350:
3341:
3337:
3332:
3327:
3323:
3319:
3315:
3311:
3307:
3300:
3297:
3292:
3288:
3284:
3280:
3276:
3272:
3268:
3264:
3260:
3253:
3250:
3245:
3241:
3237:
3233:
3229:
3225:
3221:
3217:
3209:
3207:
3203:
3198:
3192:
3188:
3181:
3179:
3175:
3170:
3166:
3162:
3158:
3154:
3150:
3146:
3139:
3137:
3133:
3128:
3124:
3120:
3116:
3112:
3108:
3104:
3097:
3095:
3091:
3086:
3082:
3077:
3072:
3067:
3062:
3058:
3054:
3051:(5): e37701.
3050:
3046:
3042:
3040:
3031:
3028:
3016:
3012:
3005:
3002:
2997:
2993:
2989:
2985:
2980:
2975:
2971:
2967:
2963:
2961:
2957:
2948:
2945:
2940:
2936:
2931:
2926:
2922:
2918:
2914:
2910:
2906:
2899:
2897:
2893:
2888:
2884:
2879:
2874:
2869:
2864:
2860:
2856:
2853:(8): e11856.
2852:
2848:
2844:
2842:
2833:
2831:
2827:
2822:
2816:
2812:
2808:
2804:
2797:
2794:
2789:
2785:
2781:
2777:
2773:
2769:
2765:
2761:
2757:
2753:
2749:
2742:
2739:
2734:
2730:
2726:
2722:
2718:
2714:
2710:
2706:
2702:
2695:
2693:
2691:
2689:
2685:
2680:
2676:
2672:
2668:
2664:
2660:
2656:
2652:
2651:
2643:
2641:
2639:
2637:
2633:
2628:
2624:
2619:
2614:
2610:
2606:
2602:
2598:
2594:
2590:
2586:
2579:
2576:
2571:
2565:
2561:
2557:
2553:
2546:
2544:
2542:
2538:
2532:
2527:
2523:
2519:
2515:
2508:
2506:
2504:
2500:
2495:
2491:
2487:
2483:
2478:
2473:
2470:(7114): 929.
2469:
2465:
2461:
2454:
2452:
2450:
2448:
2446:
2442:
2437:
2433:
2429:
2425:
2421:
2417:
2413:
2406:
2404:
2402:
2400:
2398:
2396:
2394:
2392:
2390:
2388:
2386:
2384:
2382:
2380:
2378:
2376:
2374:
2370:
2357:
2353:
2346:
2343:
2331:
2327:
2323:
2317:
2315:
2313:
2309:
2302:
2300:
2298:
2294:
2293:
2292:Prolagostomus
2288:
2287:
2282:
2281:
2276:
2275:
2270:
2266:
2265:
2261:
2257:
2256:
2252:
2248:
2247:
2242:
2241:
2237:
2236:sparassodonts
2233:
2232:
2228:
2224:
2223:
2219:
2215:
2214:
2209:
2208:
2203:
2202:
2197:
2196:
2192:
2187:
2185:
2181:
2177:
2173:
2169:
2165:
2161:
2160:Early Miocene
2157:
2153:
2149:
2145:
2140:
2138:
2134:
2130:
2126:
2123:
2119:
2115:
2111:
2107:
2103:
2099:
2095:
2092:. The area's
2091:
2087:
2083:
2079:
2071:
2064:
2062:
2061:larger prey.
2059:
2055:
2054:plesiomorphic
2050:
2049:Andalgalornis
2046:
2042:
2041:Andalgalornis
2037:
2034:
2033:Andalgalornis
2030:
2026:
2022:
2018:
2014:
2010:
2006:
2002:
1998:
1997:Andalgalornis
1988:
1984:
1980:
1979:
1978:Andalgalornis
1968:
1959:
1947:
1945:
1943:
1942:
1941:Tyrannosaurus
1937:
1933:
1929:
1925:
1921:
1917:
1912:
1908:
1907:
1902:
1897:
1892:
1890:
1886:
1881:
1877:
1873:
1869:
1865:
1861:
1857:
1856:Mesembriornis
1853:
1849:
1842:
1838:
1837:
1831:
1824:
1822:
1820:
1815:
1809:
1807:
1806:biomechanical
1803:
1799:
1795:
1786:
1779:
1774:
1772:
1770:
1766:
1762:
1757:
1753:
1745:
1740:
1733:
1731:
1729:
1725:
1720:
1716:
1712:
1708:
1704:
1700:
1696:
1692:
1686:
1684:
1680:
1676:
1672:
1668:
1663:
1659:
1654:
1652:
1648:
1644:
1640:
1637:supraorbital
1632:
1628:
1624:
1622:
1618:
1614:
1610:
1606:
1602:
1598:
1594:
1590:
1584:
1581:
1577:
1573:
1572:Andalgalornis
1569:
1565:
1561:
1557:
1547:
1537:
1526:
1517:
1515:
1514:
1513:Andalgalornis
1509:
1508:
1503:
1498:
1494:
1490:
1486:
1485:
1484:Mesembriornis
1479:
1475:
1471:
1467:
1466:
1461:
1460:
1455:
1447:
1445:
1443:
1439:
1435:
1431:
1427:
1423:
1414:
1407:
1405:
1403:
1399:
1398:thylacosmilid
1396:
1392:
1388:
1384:
1381:, during the
1380:
1376:
1372:
1367:
1365:
1361:
1360:Cariamiformes
1357:
1356:Palaeognathae
1353:
1349:
1345:
1341:
1338:underwent an
1337:
1333:
1329:
1319:
1318:
1310:
1309:
1301:
1300:
1292:
1291:
1283:
1282:
1274:
1273:
1270:
1269:
1268:
1260:
1259:
1256:
1255:
1247:
1246:
1243:
1242:
1241:
1233:
1232:
1229:
1228:
1225:
1224:
1223:
1215:
1214:
1211:
1210:
1207:
1206:
1205:
1197:
1196:
1193:
1192:
1189:
1188:
1187:
1179:
1178:
1175:
1174:
1171:
1170:
1169:
1161:
1160:
1154:
1153:
1147:
1146:
1143:
1142:
1139:
1138:
1137:
1129:
1128:
1122:
1121:
1118:
1117:
1114:
1113:
1112:
1104:
1103:
1097:
1096:
1093:
1092:
1089:
1088:
1087:
1079:
1078:
1072:
1071:
1068:
1067:
1059:
1058:
1050:
1049:
1041:
1040:
1032:
1031:
1028:
1027:
1026:
1018:
1017:
1014:
1013:
1010:
1009:
1008:
1000:
999:
993:
992:
989:
988:
980:
979:
976:
975:
974:
966:
965:
962:
961:
958:
957:
956:
948:
947:
941:
940:
934:
933:
926:
925:Psilopterinae
922:
921:
918:
917:
909:
908:
900:
899:
891:
890:
887:
886:
885:
877:
876:
873:
872:
869:
868:
867:
859:
858:
852:
851:
848:
847:
844:
843:
842:
834:
833:
827:
826:
823:
822:
819:
818:
817:
809:
808:
801:
797:
796:
790:
789:
782:
779:
778:
774:
772:
768:
764:
763:
758:
754:
750:
746:
742:
737:
735:
731:
727:
723:
719:
715:
711:
707:
703:
699:
692:
691:phorusrhacids
687:
680:
676:
671:
664:
662:
660:
656:
652:
651:
646:
645:
639:
635:
631:
627:
626:palatal bones
623:
622:quadrate bone
619:
615:
610:
608:
607:specific name
604:
600:
596:
592:
588:
584:
580:
575:
573:
569:
563:
534:
530:
526:
522:
512:
503:
492:
480:
478:
476:
472:
468:
464:
460:
456:
452:
451:
450:Andalgalornis
446:
442:
438:
434:
430:
426:
421:
419:
415:
411:
407:
403:
399:
395:
391:
386:
382:
377:
375:
374:
369:
365:
361:
357:
353:
349:
345:
344:specific name
341:
337:
336:
331:
327:
323:
319:
315:
311:
307:
303:
299:
298:
287:
281:
279:
270:
267:
266:Binomial name
263:
259:
258:
252:
249:
248:
243:
237:
236:
229:
226:
225:
222:
216:
213:
212:
209:
203:
200:
199:
196:
195:Cariamiformes
193:
190:
189:
186:
183:
180:
179:
176:
173:
170:
169:
166:
163:
160:
159:
156:
153:
150:
149:
144:
139:
135:
132:
127:
123:
118:
104:
99:
94:
89:
84:
79:
74:
69:
64:
59:
54:
49:
43:
36:
32:
26:
22:
19:
4192:Paleontology
4054:
3990:
3983:
3960:
3953:
3930:
3923:
3900:
3895:Andrewsornis
3893:
3886:
3863:
3858:Phorusrhacos
3856:
3850:
3849:
3842:
3820:Patagorhacos
3818:
3810:
3802:
3794:
3758:
3747:
3736:
3726:
3716:
3709:
3689:
3666:
3661:Occitaniavis
3659:
3652:
3645:
3638:
3615:
3591:
3585:Eutreptornis
3583:
3576:
3553:
3529:
3519:
3509:
3498:
3487:
3477:
3466:
3432:Superorder:
3363:
3331:11336/171288
3313:
3309:
3305:
3299:
3266:
3262:
3252:
3219:
3215:
3186:
3152:
3148:
3144:
3110:
3106:
3102:
3048:
3044:
3038:
3030:
3018:. Retrieved
3014:
3004:
2969:
2965:
2959:
2955:
2947:
2912:
2908:
2850:
2846:
2840:
2802:
2796:
2755:
2751:
2741:
2708:
2704:
2654:
2648:
2592:
2588:
2578:
2551:
2521:
2517:
2467:
2463:
2419:
2415:
2360:. Retrieved
2358:. p. 13
2355:
2345:
2333:. Retrieved
2329:
2299:, and fish.
2290:
2284:
2280:Protacaremys
2278:
2272:
2262:
2253:
2244:
2240:Patagosmilus
2238:
2229:
2227:astrapothere
2220:
2218:notoungulate
2211:
2205:
2201:Prepotherium
2199:
2193:
2188:
2141:
2094:stratigraphy
2077:
2076:
2057:
2048:
2040:
2038:
2032:
2020:
1996:
1994:
1976:
1939:
1923:
1919:
1910:
1904:
1900:
1895:
1893:
1855:
1851:
1845:
1834:
1825:Leg function
1810:
1791:
1775:Paleobiology
1764:
1752:Phorusrhacos
1751:
1749:
1727:
1723:
1702:
1694:
1690:
1687:
1655:
1650:
1646:
1642:
1639:ossification
1635:
1620:
1617:frontal bone
1585:
1579:
1571:
1568:Andrewsornis
1567:
1563:
1560:Phorusrhacos
1559:
1553:
1511:
1507:Andrewsornis
1505:
1501:
1496:
1493:Phorusrhacos
1492:
1488:
1482:
1477:
1473:
1470:Phorusrhacos
1469:
1463:
1457:
1453:
1451:
1441:
1434:nuchal crest
1430:Phorusrhacos
1429:
1425:
1419:
1386:
1368:
1330:, after the
1325:
1265:
1263:
1238:
1236:
1220:
1218:
1202:
1200:
1184:
1182:
1166:
1164:
1134:
1132:
1109:
1107:
1085:
1084:
1082:
1023:
1021:
1005:
1003:
971:
969:
953:
951:
882:
880:
864:
862:
839:
837:
814:
812:
770:
767:polyphyletic
760:
748:
744:
738:
733:
721:
697:
695:
658:
650:Phorusrhacos
648:
642:
634:phalanx bone
611:
603:bird of prey
590:
576:
545:70°15′29.7″W
542:41°01′59.4″S
521:phorusrhacid
518:
477:, and fish.
454:
448:
444:
422:
402:ossification
397:
393:
384:
378:
371:
363:
355:
334:
333:
306:phorusrhacid
296:
295:
294:
285:
277:
272:
256:
255:
241:
234:
233:
24:
18:
4228:Colloncuran
4079:Wikispecies
3955:Llallawavis
3932:Psilopterus
3844:Devincenzia
3738:Pseudolarus
3668:Propelargus
3531:Lavocatavis
3511:Similiornis
3434:Australaves
3310:Ameghiniana
3107:Ameghiniana
2618:11336/38650
2246:Cladosictis
2191:xenarthrans
2110:Colloncuran
2082:pyroclastic
1999:, based on
1889:glyptodonts
1848:tibiotarsus
1719:spinal cord
1679:jaw muscles
1651:Devincenzia
1459:Psilopterus
1408:Description
1395:metatherian
1375:Pleistocene
762:Llallawavis
753:sister taxa
749:Devincenzia
714:subfamilies
681:(8, middle)
644:Devincenzia
620:, and left
614:eye sockets
557: /
463:Colloncuran
418:spinal cord
373:Devincenzia
214:Subfamily:
35:Colloncuran
4202:Categories
3925:Procariama
3902:Patagornis
3796:Brontornis
3760:Smiliornis
3728:Miocariama
3702:Cariamidae
3682:Salmilidae
3555:Strigogyps
3521:Talantatos
3500:Itaboravis
3489:Gradiornis
3401:Genera of
3222:: 110932.
3113:(4): 290.
2303:References
2297:amphibians
2207:Prozaedyus
2164:cordillera
2137:lacustrine
2129:sandstones
2125:limestones
2009:bite force
1906:Brontornis
1885:armadillos
1868:long bones
1852:Patagornis
1802:borhyaenid
1794:scavengers
1765:Devicenzia
1761:metatarsal
1728:Devicenzia
1724:Patagornis
1703:Devicenzia
1695:Patagornis
1691:Patagornis
1643:Patagornis
1621:Patagornis
1593:nasal bone
1580:Patagornis
1564:Patagornis
1556:premaxilla
1502:Patagornis
1489:Patagornis
1474:Patagornis
1465:Patagornis
728:, a blunt
605:, and the
475:amphibians
441:borhyaenid
425:scavengers
406:eye socket
3985:Physornis
3804:Macrornis
3654:Oblitavis
3640:Gypsornis
3578:Bathornis
3414:Kingdom:
3340:240529252
3244:247423841
3169:163164163
3127:199099778
2780:0272-4634
2733:234119602
2671:0003-0996
2657:(1): 39.
2286:Neoreomys
2274:Maruchito
2255:Abderites
2251:marsupial
2156:cursorial
2152:bushlands
2148:woodlands
2144:semi-arid
2098:sediments
1936:dinosaurs
1928:greyhound
1662:parietals
1613:jugal bar
1402:placental
1371:Paleocene
1344:gigantism
757:cladogram
665:Evolution
618:braincase
533:Patagonia
467:cursorial
437:placental
420:enters).
368:subfamily
340:Tehuelche
326:Patagonia
284:Bertelli
250:Species:
240:Bertelli
161:Kingdom:
155:Eukaryota
4099:47444327
4085:Kelenken
4064:Wikidata
4055:Kelenken
3851:Kelenken
3647:Idiornis
3593:Paracrax
3422:Chordata
3420:Phylum:
3416:Animalia
3403:Seriemas
3364:Kelenken
3291:22214720
3085:22662194
3045:PLOS ONE
2996:13543638
2988:22723482
2939:16096087
2887:20805872
2847:PLOS ONE
2679:43707746
2627:85212917
2486:17066027
2436:85693135
2271:such as
2168:Friasian
2133:alluvial
2114:foreland
2078:Kelenken
2058:Kelenken
2021:Kelenken
2005:CT scans
1924:Kelenken
1920:Kelenken
1911:Kelenken
1901:Kelenken
1896:Kelenken
1756:tubercle
1734:Leg bone
1647:Kelenken
1546:holotype
1497:Kelenken
1478:Kelenken
1454:Kelenken
1442:Kelenken
1426:Kelenken
1387:Kelenken
1379:Pliocene
1364:seriemas
1328:Cenozoic
771:Kelenken
745:Kelenken
722:Kelenken
698:Kelenken
659:Kelenken
587:holotype
511:Holotype
481:Taxonomy
455:Kelenken
445:Kelenken
433:Cenozoic
398:Kelenken
394:Kelenken
385:Kelenken
364:Kelenken
360:seriemas
356:Kelenken
330:holotype
297:Kelenken
235:Kelenken
201:Family:
175:Chordata
171:Phylum:
165:Animalia
151:Domain:
40:15
25:Kelenken
4166:Portals
4138:3600058
4125:1474098
4112:4966577
4070:Q133129
3865:Titanis
3749:Riacama
3711:Cariama
3691:Salmila
3438:Clade:
3426:Class:
3283:9702279
3224:Bibcode
3076:3360764
3053:Bibcode
3015:NPR.org
2930:1559870
2878:2923598
2855:Bibcode
2788:4524003
2760:Bibcode
2713:Bibcode
2597:Bibcode
2494:4381103
2269:rodents
2260:primate
2013:newtons
1932:grazing
1864:cheetah
1860:ostrich
1836:Titanis
1769:foramen
1658:sutures
1597:maxilla
1591:of the
1589:process
1336:mammals
927:
802:
783:
734:Titanis
706:rostrum
679:Comallo
673:Map of
531:in the
525:Comallo
471:mammals
322:Comallo
310:extinct
275:†
254:†
227:Genus:
191:Order:
181:Class:
109:↓
4151:105905
3718:Chunga
3338:
3289:
3281:
3242:
3193:
3167:
3125:
3083:
3073:
3020:26 May
2994:
2986:
2937:
2927:
2885:
2875:
2817:
2786:
2778:
2731:
2677:
2669:
2625:
2566:
2492:
2484:
2464:Nature
2434:
2362:26 May
2335:26 May
2289:, and
2267:, and
2258:, the
2249:, the
2234:, the
2225:, the
2216:, the
2210:, and
2118:basins
2003:using
1985:and a
1880:marrow
1814:thorax
1675:temple
1601:septum
1576:tomium
1570:, and
1510:, and
1354:, the
1350:, the
1264:
1237:
1219:
1201:
1183:
1165:
1133:
1108:
1083:
1022:
1004:
970:
952:
923:
881:
863:
838:
813:
798:
751:to be
710:culmen
702:family
593:; the
286:et al.
242:et al.
4180:Birds
4120:IRMNG
3336:S2CID
3316:(6).
3287:S2CID
3240:S2CID
3165:S2CID
3123:S2CID
2992:S2CID
2784:JSTOR
2729:S2CID
2675:JSTOR
2623:S2CID
2524:(4).
2490:S2CID
2432:S2CID
2090:tuffs
1938:like
1872:saiga
1819:hawks
1448:Skull
1438:horse
595:genus
390:horse
314:birds
302:genus
300:is a
4107:GBIF
3428:Aves
3279:PMID
3191:ISBN
3081:PMID
3022:2022
2984:PMID
2935:PMID
2883:PMID
2815:ISBN
2776:ISSN
2667:ISSN
2564:ISBN
2482:PMID
2364:2022
2337:2022
2243:and
2182:, a
2172:arid
2127:and
2104:and
2027:and
1916:rhea
1887:and
1726:and
1487:and
1462:and
747:and
655:taxa
647:and
288:2007
244:2007
185:Aves
48:PreꞒ
4094:EoL
3326:hdl
3318:doi
3271:doi
3232:doi
3220:594
3157:doi
3115:doi
3071:PMC
3061:doi
2974:doi
2970:215
2925:PMC
2917:doi
2913:272
2873:PMC
2863:doi
2807:doi
2768:doi
2721:doi
2659:doi
2655:102
2613:hdl
2605:doi
2556:doi
2526:doi
2472:doi
2468:443
2424:doi
2150:or
1874:or
589:of
304:of
4204::
4148::
4135::
4122::
4109::
4096::
4081::
4066::
3334:.
3324:.
3314:58
3312:.
3285:.
3277:.
3267:45
3265:.
3261:.
3238:.
3230:.
3218:.
3205:^
3177:^
3163:.
3153:27
3151:.
3135:^
3121:.
3111:56
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3093:^
3079:.
3069:.
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2990:.
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2968:.
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2933:.
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2907:.
2895:^
2881:.
2871:.
2861:.
2849:.
2845:.
2829:^
2813:.
2782:.
2774:.
2766:.
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2754:.
2750:.
2727:.
2719:.
2709:40
2707:.
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2687:^
2673:.
2665:.
2653:.
2635:^
2621:.
2611:.
2603:.
2593:35
2591:.
2587:.
2562:.
2540:^
2522:43
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2516:.
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2430:.
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2418:.
2414:.
2372:^
2354:.
2328:.
2311:^
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2277:,
2204:,
2198:,
1891:.
1623:.
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1562:,
1516:.
1504:,
1334:,
661:.
376:.
362:.
98:Pg
42:Ma
37:)~
4168::
3973:†
3943:†
3913:†
3876:†
3832:†
3823:?
3815:?
3807:?
3799:?
3784:†
3772:†
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3527:†
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3380:v
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3293:.
3273::
3246:.
3234::
3226::
3199:.
3171:.
3159::
3129:.
3117::
3087:.
3063::
3055::
3049:7
3024:.
2998:.
2976::
2941:.
2919::
2889:.
2865::
2857::
2851:5
2823:.
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2790:.
2770::
2762::
2735:.
2723::
2715::
2681:.
2661::
2629:.
2615::
2607::
2599::
2572:.
2558::
2534:.
2528::
2496:.
2474::
2438:.
2426::
2366:.
2339:.
231:†
218:†
205:†
103:N
93:K
88:J
83:T
78:P
73:C
68:D
63:S
58:O
53:Ꞓ
33:(
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