The last terror birds

Skeleton of the terror bird Titanis walleri at the Florida Museum of Natural History.

In 1887, Florentino Ameghino, the “father of Argentinian Palaeontology”, described a large, toothless jaw from the Miocene of the Province of Santa Cruz, naming it Phorusrhacos longissimus and assigning it to a new family of edentulous mammal. He used this finding as a critical evidence for his contention that modern mammalian lineages originated in Argentina and later spread across the globe. Four years later, Moreno and Mercerat recognized for the first time that the mandible described by Ameghino was really that of a bird.

The Phorusrhacidae, the so-called “terror birds”, were a group of medium-to large sized extinct ground-dwelling birds, which lived during the Cenozoic, and became the dominant carnivores of South America while it was an isolated continent. They are characterized by their elongated hindlimbs, narrow pelvis, reduced forelimbs, and their huge skull with their tall, long, narrow, and hollow beaks ended in a hook. Kelenken guillermoi, is the largest known phorusrhacid and lived in the Miocene of Argentina. The skull reaches a length of 71.6 cm and the whole animal would reach 3 m high. Kelenken is also represented by a tarsometatarsus and a broken phalanx and proceeds from the locality of Comallo (Río Negro Province, Argentina). Titanis walleri, lived during the Pliocene and Pleistocene of North America. It was 2.5 metres tall and weighed approximately 150 kilograms. This giant bird is interpreted as an early immigrant during the Great American Interchange.

Proximal portion of the left humerus of Psilopterus sp. Caudal, b ventral, c cranial and d dorsal views (From Jones et. al., 2017)

At the end of the Pliocene, Phorusrhacids decline in diversity. Two new specimens support the hypothesis that the latest geologic occurrence of the Phorusrhacidae comes from late Pleistocene sediments of Uruguay. The remains comprise the distal portion of right tarsometatarsus and a left humerus; the latter is assigned to the genus Psilopterus. The first material (MPAB-520) comes from Soriano, Uruguay, and the sediments belong to the Dolores Formation (Lujanian Stage-Age, late Pleistocene/early Holocene). The following features identify the specimen as a phorusrhacid bird. (1) a large and distally expanded trochlea metatarsi III; (2) a very narrow trochlea metatarsi II with the articular surface transversally convex and without any longitudinal sulcus (in dorsal and distal views); (3) in dorsal view the trochlea metatarsi II is almost parallel and much shorter than the middle trochlea, and forming a narrow notch between trochleae II and III. The second material consists of a left humerus without distal epiphysis belonged to Museo Paleontológico Alejandro Berro (MPAB-2024).

There are two explanatory hypotheses proposed for the decline of the terror birds: environmental reasons or direct competition (at least for the larger specimens) with placental carnivore’s immigrants to South America after the setting of the Panamanian bridge. 



Jones, W., Rinderknecht, A., Alvarenga, H. et al. PalZ (2017), The last terror birds (Aves, Phorusrhacidae): new evidence from the late Pleistocene of Uruguay,

ALVARENGA, Herculano M.F.  and  HOFLING, Elizabeth. Systematic revision of the Phorusrhacidae (Aves: Ralliformes). Pap. Avulsos Zool. (São Paulo) [online]. 2003, vol.43, n.4 [cited  2015-03-24], pp. 55-91 .


Solving a Darwinian mystery

Macrauchenia patachonica by Robert Bruce Horsfall.

During the first two years of his voyage aboard HMS Beagle, Charles Darwin collected a considerable number of fossil mammals from various South American localities. Darwin sent all the specimens to the Reverend Professor John Stevens Henslow, his mentor and a close friend. The samples were deposited in the Royal College of Surgeons where Richard Owen began its study. Between 1837 and 1845, Owen described eleven taxa, including: Toxodon platensis, Macrauchenia patachonica, Equus curvidens, Scelidotherium leptocephalum, Mylodon darwinii, and Glossotherium sp.

Macrauchenia, meaning “big neck,” was named by Richard Owen based on limb bones and vertebrae collected by Charles Darwin on January 1834 at Puerto San Julian, in Santa Cruz Province, Argentina. The bizarre animal had a camel-like body, with sturdy legs, a long neck and a relatively small head. Owen described as “A large extinct Mammiferous Animal, referrible to the Order Pachydermata; but with affinities to the Ruminantia, and especially to the Camelidae”Macrauchenia is now considered among the more derived native South American litopterns, an endemic order whose fossil record extends from the Paleocene to the end of the Pleistocene and includes some 50 described genera. Darwin also made inferences about the environment which Macrauchenia lived: “Mr. Owen… considers that they form part of an animal allied to the guanaco or llama, but fully as large as the true camel. As all the existing members of the family of Camelidae are inhabitants of the most sterile countries, so we may suppose was this extinct kind… It is impossible to reflect without the deepest astonishment, on the changed state of this continent. Formerly it must have swarmed with great monsters, like the southern parts of Africa, but now we find only the tapir, guanaco, armadillo, capybara; mere pigmies compared to antecedents races… Since their loss, no very great physical changes can have taken place in the nature of the Country. What then has exterminated so many living creatures?…We are so profoundly ignorant concerning the physiological relations, on which the life, and even health (as shown by epidemics) of any existing species depends, that we argue with still less safety about either the life or death of any extinct kind” (Voyage of the Beagle, Chapter IX, Jan. 1834).

Dated mitogenomic phylogenetic tree. (From Westbury, M. et al)

The unusual morphological traits displayed by extinct South American native ungulates defied both Charles Darwin and Richard Owen, who tried to resolve their relationships. Two recently published molecular studies, using protein (collagen) sequence information, found that litopterns as well as notoungulates formed a monophyletic unit that shared more recent common ancestry with Perissodactyla than with any other extant placental group.

A valuable tool for uncovering phylogenetic relationships of extinct animals is ancient DNA (aDNA), although, attempts to use standard aDNA methodologies to collect genetic material from specimens from low-latitude localities have been largely unsuccessful. However, a new study recovered a nearly complete mitochondrial genome for Macrauchenia from a cave in southern Chile. The small size of the mitochondrial genome simplifies the assembly of fossil sequences using de novo methods.

In theory, reconstructing an ancient genome de novo can be undertaken without relying on a close relative’s DNA for guidance, but due to contaminant DNA and low average fragment lengths, de novo assembly is generally considered not computationally feasible. A promising new approach is using  the genetic codes of numerous living species as reference points, allowing them to reliably predict the fossil’s likeliest genetic sequences. Using the new approach, the phylogenetic analyses place Macrauchenia as a sister taxon to all living Perissodactyla, with the origin of Panperissodactlya at 66 Ma.



Westbury, M. et al. A mitogenomic timetree for Darwin’s enigmatic South American mammal Macrauchenia patachonicaNat. Commun. 8, 15951 doi: 10.1038/ncomms15951 (2017).

Welker, F. et al. Ancient proteins resolve the evolutionary history of Darwin’s South American ungulates. Nature 522, 81–84 (2015). doi:10.1038/nature14249

Buckley, M. Ancient collagen reveals evolutionary history of the endemic South American ‘ungulates’. Proc. Biol. Sci. 282, 20142671 (2015). DOI: 10.1098/rspb.2014.2671

The Megafauna extinction in South América.


Megatherium americanum Cuvier, 1796. Museo Argentino de La Plata.

Megatherium americanum Cuvier, 1796. Museo Argentino de La Plata.

During the Pleistocene and the early Holocene,  most of the terrestrial megafauna became extinct. It was a deep global-scale event. The extinction was notably more selective for large-bodied animals than any other extinction interval in the last 65 million years. Multiple explanatory hypotheses have been proposed for this event: climatic change, over hunting, habitat alteration, and the introduction of a new disease. Traditionally, the focus of research and debate has been on the Eurasian and North American extinctions. In North America, two mammalian orders (Perissodactyla, Proboscidea) were eliminated completely. At the species level, the extinction was total for mammals larger than 1000 kg and greater than 50% for size classes between 1000 and 32 kg. Early observations confirm that extinctions could be severe even in relatively climatically stable regions where the vegetation changed little. In South America the event was  more severe, with the loss of 50 megafaunal genera. Three orders of mammals disappeared (Notoungulata, Proboscidea, Litopterna), as did all megafaunal xenarthrans and at the species level, the extinction was total for mammals larger than 320 kg (Koch and Barnosky, 2006).

“Descuartizando un gliptodonte. Escenas de la vida del hombre primitivo” (Quartering a glyptodont. Scenes from the life of primitive man). Painting by Luis de Servi. Museo de la Plata).

“Descuartizando un gliptodonte. Escenas de
la vida del hombre primitivo” (Quartering a
glyptodont. Scenes from the life of primitive man). Painting by Luis de Servi, Museo de la Plata.

Before the Great American Biotic Interchange, about 3 million years ago, the largest mammals in South America were mainly endemic notoungulates, litopterns and xenarthrans. But, during the interchange, many other megamammals and large mammals arrived to South America. The late Pleistocene in this region is first characterized by a rapid cooling. During the Pleistocene-Holocene transition pollen sequences suggest a change to sub-humid climatic conditions. In addition to rapid climate change, the extinctions are seen as the result of habitat loss, reduced carrying capacity for herbivores, resource fragmentation or disturbances in the co-evolutionary equilibrium between plants, herbivores, and carnivores. The death event of the gomphothere population in Águas de Araxá (Brazil)  about 55,000 years ago, is probably an example of individuals that were suffering with the climate changes during the Late Pleistocene.

Paleontological and archaeological data indicate that extinctions seem more common after the human arrival and during the rapid climate change between 11.200 and 13.500 years. This pattern suggests that a synergy of human impacts and rapid climate change—analogous to what is happening today — may enhance extinction probability (Prado et al., 2015).



J.L. Prado et al. (2015). “Megafauna extinction in South America: A new chronology for the Argentine Pampas.” Palaeogeography, Palaeoclimatology, Palaeoecology 425: 41–49

Alroy, John. (2001). “A Multispecies Overkill Simulation of the End-Pleistocene Megafaunal Mass Extinction.” Science 292:1893-1896

Koch PL, Barnosky AD (2006) Late Quaternary extinctions: State of the debate. Annu Rev Ecol Evol Syst 37:215–250.

Prescott GW, Williams DR, Balmford A, Green RE, Manica A. (2012) Quantitative global analysis of the role of climate and people in explaining late Quaternary megafaunal extinctions. Proc. Natl Acad. Sci. USA 109,45274531

Barnosky AD, Lindsey EL.(2010) Timing of Quaternary megafaunal extinction in South America in relation to human arrival and climate change