Phorusrhacidae

Reconstruction of Psilopterus lemoinei, one of the smaller phorusrhacids, by FunkMonk.

Belongs within: Cariamae.

The Phorusrhacidae include the 'terror birds' of South America, flightless carnivorous birds that lived from the mid-Palaeocene to the Upper Pliocene/Lower Pleistocene. The latest surviving species, Titanis walleri, is also notable for being found in southern North America. Some phorusrhacids reached significant sizes, with heights comparable to a human; other species were smaller, possibly a little over two feet in height (Alvarenga & Höfling 2003).

Alvarenga & Höfling (2003) recognised five subfamilies within the Phorusrhacidae (excluding Cunampaia, which they regarded as undiagnostic). The smallest phorusrhacids belonged to the Psilopterinae, and the largest to the Phorusrhacinae and Brontornithinae. The Phorusrhacinae were of a more slender and cursorial build than the heavy, robust Brontornithinae.

Characters (from Alvarenga & Höfling 2003): Large or even gigantic build; body slim in frontal view, especially the premaxila, pelvis and thorax being laterally flattened; premaxilla bulky, especially high, with a pointed strong apex in the form of a hook; jaw with very solid symphysis; ample and pervious nostrils (septum absent); palate desmognate; basipterigoid processes well-developed; pterigoids with evident articulation for basipterigoid process in mid-portion; ribs lacking uncinate processes; pubis atrophied in cranial half; wings reduced; coracoid with extreme reduction of procoracoidal and acrocoracoidal processes, possessing an ample scapular facet, in the form of a groove in the apex; humerus with internal tuberosity bulging proximally, proximal half of diaphysis being strongly bent and processus flexorius distally prominent; tarsometatarsus with triangular-shaped hypotarsus when viewed from below and without tendon grooves; strongly curved ungual phalanx.

<==Phorusrhacidae (see below for synonymy)
    |  i. s.: Patagornithinae [Tolmodinae] AH03
    |           |--Patagornis Moreno & Mercerat 1891 (see below for synonymy) AH03
    |           |    `--*P. marshi Moreno & Mercerat 1891 (see below for synonymy) AH03
    |           |--Andalgalornis Patterson & Kraglievich 1960 AH03
    |           |    `--A. steulletti (Kraglievich 1931) (see below for synonymy) AH03
    |           `--Andrewsornis Patterson 1941 AH03
    |                |--*A. abbotti Patterson 1941 AH03
    |                `--‘Aucornis’ solidus Ameghino 1898 (n. d.) AH03
    |--+--Phorusrhacinae [Phororhacinae] FP64
    |  |    |--Phorusrhacos Ameghino 1887 AH03 (see below for synonymy)
    |  |    |    `--*P. longissimus Ameghino 1887 AH03 (see below for synonymy)
    |  |    |--Titanis Brodkorb 1963 AH03
    |  |    |    `--*T. walleri Brodkorb 1963 AH03
    |  |    `--Devincenzia Kraglievich 1932 [incl. Onactornis Cabrera 1939] AH03
    |  |         `--D. pozzi (Kraglievich 1931) (see below for synonymy) AH03
    |  `--+--Cunampaia Rusconi 1946 FP64, AH03 [Cunampaiidae]
    |     |    `--C. simplex FP64
    |     `--Brontornithinae [Brontorniinae] FP64
    |          |--Brontornis Moreno & Mercerat 1891 [incl. Rostrornis Moreno & Mercerat 1891] AH03
    |          |    `--*B. burmeisteri Moreno & Mercerat 1891 (see below for synonymy) AH03
    |          |--Physornis Ameghino 1895 [incl. Aucornis Ameghino 1898] AH03
    |          |    `--*P. fortis Ameghino 1895 [incl. Aucornis euryrhynchus Ameghino 1899] AH03
    |          `--Paraphysornis Alvarenga 1993 AH03
    |               `--*P. brasiliensis (Alvarenga 1982) [=Physornis brasiliensis] AH03
    `--Psilopteridae [Pelecyornidae, Pelecyornithidae, Psilopteriidae] FP64
         |--Mesembriornis Moreno 1889 (see below for synonymy) AH03
         |    |--*M. milneedwardsi Moreno 1889 (see below for synonymy) AH03
         |    `--M. incertus (Rovereto 1914) [=Prophororhacos incertus] AH03
         `--Psilopterinae AH03
              |  i. s.: ‘Paleociconia’ cristata Moreno & Mercerat 1891 (n. d.) AH03
              |--Smiliornis Ameghino 1891 AH03
              |    `--S. penetrans Ameghino 1899 AH03
              |--Procariama Rovereto 1914 AH03
              |    `--*P. simplex Rovereto 1914 AH03
              |--Riacama Ameghino 1899 AH03
              |    `--R. caliginea Ameghino 1899 AH03
              |--Lophiornis Ameghino 1891 AH03
              |    `--L. obliquus Ameghino 1891 AH03
              |--Paleopsilopterus Alvarenga 1985 AH03
              |    `--*P. itaboraiensis Alvarenga 1985 AH03
              |--Pseudolarus Ameghino 1891 AH03
              |    |--P. eocaenus Ameghino 1891 A94
              |    `--P. guaraniticus Ameghino 1899 AH03
              `--Psilopterus Moreno & Mercerat 1891 [incl. Pelecyornis Ameghino 1891, Staphylornis Mercerat 1897] AH03
                   |--P. bachmanni (Moreno & Mercerat 1891) AH03 (see below for synonymy)
                   |--P. affinis (Ameghino 1899) [=Phororhacos affinis] AH03
                   |--P. colzecus Tonni & Tambussi 1988 AH03
                   `--P. lemoinei (Moreno & Mercerat 1891) AH03 (see below for synonymy)

Nomen nudum: Paleociconia australis Moreno 1889 AH03

Andalgalornis steulletti (Kraglievich 1931) [=Phororhacos steulletti; incl. P. deautieri Kraglievich 1931, *Andalgalornis ferox Patterson & Kraglievich 1960] AH03

*Brontornis burmeisteri Moreno & Mercerat 1891 [incl. Rostrornis floweri Moreno & Mercerat 1891, B. platyonyx Ameghino 1895] AH03

Devincenzia pozzi (Kraglievich 1931) [=Phororhacos pozzi, Onactornis pozzi; incl. O. depressus Cabrera 1939, *Devincenzia gallinali Kraglievich 1932, Phororhacos longissimus mendocinus Kraglievich 1931] AH03

Mesembriornis Moreno 1889 [incl. Hermosiornis Moreno 1889, Paleociconia Moreno 1889, Prophororhacos Rovereto 1914; Hermosiornidae, Hermosiorniidae, Hermosiornithidae, Hermosiornithinae, Mesembriornithinae, Prophororhacinae] AH03

*Mesembriornis milneedwardsi Moreno 1889 [incl. *Paleociconia australis Moreno 1889, Prophororhacos australis, Hermosiornis milneedwardsi Rovereto 1914, Driornis pampeanus Moreno & Mercerat 1891, Hermosiornis rapax Kraglievich 1946] AH03

Patagornis Moreno & Mercerat 1891 [incl. Morenomerceraria Lambrecht 1933, Tolmodus Ameghino 1891] AH03

*Patagornis marshi Moreno & Mercerat 1891 [incl. Tolmodus inflatus Ameghino 1891, Phororhacos inflatus] AH03

Phorusrhacidae [Brontornithes, Brontornithidae, Darwinornithidae, Devincenziidae, Mesembriornidae, Mesembriornithidae, Mesembriornithinae, Paleociconiinae, Patagornithidae, Phororhacidae, Phororhacoidea, Phororhacosidae, Phorusrhacoidae, Phorusrhacoidea, Phorusrhacosidae, Stereornithidae]

Phorusrhacos Ameghino 1887 AH03 [=Phororhacos Ameghino 1889 AH03; incl. Callornis Ameghino 1895 AH03, Darwinornis Moreno & Mercerat 1891 AH03, Dryornis Moreno & Mercerat 1891 A94, Eucallornis Ameghino 1901 AH03, Liornis Ameghino 1895 AH03, Owenornis Moreno & Mercerat 1891 AH03, Stephanornis Mercerat 1893 A94, Stereornis Moreno & Mercerat 1891 AH03, Titanornis Mercerat 1893 AH03; Liorninae, Liornithidae]

Phorusrhacos longissimus Ameghino 1887 AH03[=Phororhacos longissimus AH03; incl. Owenornis affinis Moreno & Mercerat 1891 AH03, Darwinornis copei Moreno & Mercerat 1891 AH03, *Liornis floweri Ameghino 1895 A94, AH03, Stereornis gaundryi Moreno & Mercerat 1891 AH03, *Callornis giganteus Ameghino 1895 A94, AH03, Eucallornis giganteus AH03, Owenornis lydekkeri Moreno & Mercerat 1891 AH03, Liornis minor Dolgopol de Saez 1927 AH03, Titanornis mirabilis Mercerat 1893 AH03, Phororhacos platygnathus Ameghino 1891 AH03, Mesembriornis quatrefragesi Moreno & Mercerat 1891, Phororhacos sehuensis Ameghino 1891 AH03, Stereornis rollieri Moreno & Mercerat 1891 AH03, Darwinornis socialis Moreno & Mercerat 1891 AH03, Mesembriornis studeri Moreno & Mercerat 1891 AH03, D. zitteli Moreno & Mercerat 1891 AH03]

Psilopterus bachmanni (Moreno & Mercerat 1891) AH03 [=Patagornis bachmanni AH03; incl. *Ps. communis Moreno & Mercerat 1891 AH03, Pelecyornis communis A94, Phororhacos delicatus Ameghino 1891 AH03, Psilopterus intermedius Moreno & Mercerat 1891 AH03, Ps. minutus Ameghino 1891 AH03, Pelecyornis minutus A94, Pe. pueyrredonensis Sinclair & Farr 1932 AH03]

Psilopterus lemoinei (Moreno & Mercerat 1891) AH03 [=Patagornis lemoinei AH03; incl. Ps. australis Moreno & Mercerat 1891 AH03, Pelecyornis australis A94, Staphylornis erythacus Mercerat 1897 AH03, S. gallardoi Mercerat 1897 AH03, Phororhacos modicus Ameghino 1895 AH03, Pelecyornis tenuirostris Sinclair & Farr 1932 AH03, Pe. tubulatus Ameghino 1895 AH03]

*Type species of generic name indicated

REFERENCES

[AH03] Alvarenga, H. M. F., & E. Höfling. 2003. Systematic revision of the Phorusrhacidae (Aves: Ralliformes). Papéis Avulsos de Zoologia 43 (4): 55-91.

[A94] Ameghino, F. 1894. Sur les oiseaux fossiles de Patagonie et la aune mammalogique des couches a Pyrotherium. Boletín del Instituto Geográfico Argentino 15 (11-12): 501-602.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.