Phorusrhacids the terror birds

7 Phorusrhacids: the Terror Birds
HERCULANO ALVARENGA, 1 LUIS CHIAPPE, 2
AND SARA BERTELLI 3
1
´ ´ ´
Museu de Historia Natural de Taubate, Taubate, Brazil
2
Los Angeles County Museum, Los Angeles, USA
3
Museum f€ r Naturkunde, Berlin, Germany
u

At the beginning of the Tertiary, while mammals Pleistocene (Alvarenga et al., 2010). Ratites and
were undergoing an evolutionary explosion, phorusrhacids apparently lived together, perhaps
several groups of birds developed a tendency to competing for similar terrestrial habitats, during
gigantism. These groups were scattered across the whole of the Tertiary. At that time South
almost the whole planet. The Gastornithidae America was an isolated island, and only the
(also known as Diatrymidae), with possible affin- ratites, perhaps the most vulnerable among
ity with the Anseriformes (Andors, 1992), have these birds, have survived to present times.
been recovered from Paleocene and Eocene depos- Scientific investigations on the Phorusrhacidae
its in North America, Europe and Asia (Matthew started at the end of the 19th century with the work
& Granger, 1917; Martin, 1992; Hou, 1980). In of Ameghino (1887), who described a mandible of a
Australia, another group of giant birds, the Dro- “probable toothless mammal”, which he named
mornithidae, with possible affinity to the Anser- Phorusrhacus longissimus. However, it was Mor-
iformes (Wroe, 1998; Murray & Vickers-Rich, eno (1889) who first called attention to the huge
2004) had a broad diversity in the mid-Tertiary. bones that he identified as giant birds from the
Among these, Dromornis stirtoni would have Tertiary period in Argentina. Later, Moreno &
been comparable in size to the largest birds ever Mercerat (1891) and two other publications of
found. The Ratitae, also present since the Paleo- Ameghino (1891a,b) recognized Phorusrhacus as
cene in South America and possibly in Europe a bird and named several genera and species for the
(Alvarenga, 1983; Martin, 1992), is another Phorusrhacidae. A number of bones and fragments
group remarkable for the large size reached in of bones were described as a new species or genera,
several species. Among the giant birds of the Ter- resulting in a complicated and extended synonymy
tiary, they are the only present-day survivors. The within the family; currently 14 genera and 18
elephant-bird, Aepyornis maximus, a ratite from species are recognized (Table 7.1). A more detailed
Madagascar that died out around 700 years ago, is history of the first investigations of this family, as
probably the biggest bird ever found (Amadon, well as of the first classifications proposed, is set
1947; Wetmore, 1967). out by Alvarenga & H€fling (2003).
o
The Phorusrhacidae, another group of giants In this chapter, our main objective is to present
was present in South America from the Paleocene a phylogenetic analysis of the Phorusrhacidae, and
(Alvarenga, 1985) and survived until the end of the at the same time call attention to anatomic details

Living Dinosaurs: The Evolutionary History of Modern Birds, First Edition. Edited by Gareth Dyke and Gary Kaiser.
Ó 2011 John Wiley & Sons, Ltd. Published 2011 by John Wiley & Sons, Ltd.

Table 7.1 List of the Phorusrhacidae species, including the junior synonymous, the geographical and stratigraphical distribution.
188

Taxon/reference Geographic locality Horizon/Age Additional references

Mesembriornis milneedwardsi Moreno, 1889 Argentina, Province, Buenos Aires: Upper Pliocene (Montehermosan) o
Alvarenga & H€fling, 2003
Paleociconia australis Moreno, 1889; Moreno & Monte Hermoso, Rio Loberia
Mercerat, 1891
Driornis pampeanus Moreno & Mercerat, 1891
(part: only femur)
Hermosiornis milneedwardsi Rovereto, 1914
Hermosiornis rapax Kraglievich, 1946
Prophororhacos australis Brodkorb, 1967
Mesembriornis incertus (Rovereto, 1914) Argentina, Province, Catamarca: Upper Miocene to Lower Pliocene Acosta Hospitaleche, 2002;
Phororhacos incertus Rovereto, 1914 Andalgal, Corral Quemado;
a (Huayquerian) o
Alvarenga H€fling, 2003
Province, Buenos Aires: P. de Villarino
Procariama simplex Rovereto,1914 Argentina, Province, Catamarca, Upper Miocene to Lower Pliocene Tonni, 1980; Alvarenga
Andalgal,Corral Quemado, Belém,
a (Huayquerian, Chasicoan) o
H€fling, 2003
Chiquimil, Rio Santa Maria
Paleopsilopterus itaboraiensis, Alvarenga,1985 Brazil, Est. Rio de Janeiro: S~o José de
a Upper Paleocene (Itaboraian) o
Itabora ı
Psilopterus bachmanni (Moreno Mercerat, 1891) Argentina, Province, Santa Cruz: Santa Middle Miocene (Santacrucian) o
Patagornis bachmanni Moreno Mercerat, 1891 Cruz, Lago Pueyrredon, Monte
Psilopterus communis Moreno Mercerat, 1891 Observación, La Cueva
Psilopteus intermedius Moreno Mercerat, 1891
Phororhacus delicatus Ameghino, 1891
Pelecyornis minutus Ameghino, 1891
Pelecyornis pueyrredonensis Sinclair Farr, 1932
Psilopterus affinis (Ameghino, 1899) Argentina, Province, Chubut: Cabe¸ a
c Middle Oligocene (Deseadan) o
Alvarenga H€fling, 2003;
Phororhacus affinis Ameghino, 1899 Blanca Agnolin, 2006
Psilopterus colzecus Tonni Tambussi, 1988 Argentina, Province, Buenos Aires: Upper Miocene (Chasicoan) o
Partido de Villarino
HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLI

Psilopterus lemoinei (Moreno Mercerat, 1891) Argentina, Province, Santa Cruz: Santa Middle Miocene (Santacrucian) o
Patagornis lemoinei Moreno Mercerat, 1891 Cruz, Killik Aike, Monte Observación, Degrange Tambussi, 2008
Psilopterus australis Moreno Mercerat, 1891; Take Harvey, La Cueva, Corriguen
Brodkorb, 1967 Kaik, Taguaquemada
Pelecyornis tubulatus Ameghino, 1895
Phororhacus modicus Ameghino, 1895
Staphylornis gallardoi Mercerat, 1897
Staphylornis erythacus Mercerat, 1897
Pelecyornis tenuirostris Sinclair Farr, 1932

Patagornis marshi Moreno Mercerat, 1891 Argentina, Province, Santa Cruz;Monte Lower to Middle Miocene. o
Tolmodus inflatus Ameghino, 1891 Observación, Tagua Quemada, La (Santacrucian)
Phororhacos inflatus Ameghino, 1891; Andrews, Cueva
1899; Brodkorb, 1967
Paleociconia cristata Brodkorb, 1967
Andrewsornis abbotti Patterson, 1941 Argentina, Province,Chubut (Cabe¸ a
c Middle to Upper Oligocene o
Blanca) and Santa Cruz (Pico Truncado) (Deseadan)
Andalgalornis steulleti (Kraglievich, 1931) Argentina, Province, Entre Rios and Upper Miocene to Lower Pliocene. o
Tonni, 1980; Alvarenga H€fling,
Phororhacos steulleti Kraglievich, 1931 Catamarca (Chiquimil) Fm Andalgal, (Huayquerian)
a 2003; Noriega Agnolin, 2008
Phororhacos deautieri Kraglievich, 1931
Andalgalornis ferox Paterson Kraglievich, 1960
Andalgalornis steulleti Brodkorb, 1967
Phorusrhacus longissimus Ameghino, 1887 Argentina, Province, Santa Cruz, La Lower and Middle Miocene o
Phororhacos longissimus, Ameghino, 1889 Cueva, Tagua/Quemada, Monte (Santacrucian)
Stereornis rollieri Moreno Mercerat, 1891 Observación, Rio Sehuén
Stereornis gaudryi Moreno Mercerat, 1891
Mesembriornis studeri Moreno Mercerat, 1891
Mesembriornis quatrefragesi Moreno Mercerat, 1891
Darwinornis copei Moreno Mercerat, 1891
Darwinornis zittelli Moreno Mercerat, 1891
Darwinornis socialis Moreno Mercerat, 1891
Owenornis affinis Moreno Mercerat, 1891
Owenornis lydekkeri Moreno Mercerat, 1891
Phororhacus sehuensis Ameghino, 1891
Phororhacus platygnathus Ameghino, 1891
Titanornis mirabilis, Moreno Mercerat, 1891
Phorusrhacids: the Terror Birds

Callornis giganteus, Ameghino, 1895
Eucallornis giganteus Ameghino, 1901
Liornis floweri Ameghino, 1895
Titanis walleri Brodkorb, 1963 USA, Florida (Inglish) and Texas (Baskin) Pliocene (Hemphillian to Late Gould Quitmer, 2005;
Blancan) MacFadden et al., 2006
(continued)
189

Table 7.1 (Continued)
190

Taxon/reference Geographic locality Horizon/Age Additional references

Devincenzia pozzi (Kraglievich, 1931) Argentina, Province, Buenos Aires and Upper Miocene to Upper Pliocene Tambussi et al., 1999; Alvarenga
Phororhacos pozzi Kraglievich, 1931 Entre Rios; Uruguay (Huayquerian and o
H€fling, 2003; Alfaro Perea,
Phororhacus longissimus mendocinus Kraglievich, Mesopotamien); Fm Raigon 2004; Agnolin, 2006; Noriega
1931 (Uruguay). Agnolin, 2008
Devincenzia gallinali Kraglievich, 1932
Onactornis depressus Cabrera, 1939
Onactornis pozzi Brodkorb, 1967
Onactornis mendocinus Brodkorb, 1967
Kelenken guillermoi Bertelli, Chiappe Argentina, Province,Rio Negro, Comallo. Midle Miocene (Fm Collón Cur)
a Chiappe Bertelli, 2007
Tambussi, 2007
Physornis fortis Ameghino, 1895 Argentina, Province, Santa Cruz, Puerto Middle Oligocene (Deseadan) Alvarenga, 1993; Alvarenga
Aucornis euryrhynchus Ameghino, 1898 Deseado, Punta Nova, La Flexa. o
H€fling, 2003
Paraphysornis brasiliensis (Alvarenga, 1982) Brazil, Est. S~o Paulo, Tremembé.
a Upper Oligocene or Lower Miocene Alvarenga, 1993
Physornis brasiliensis Alvarenga, 1982 (Upper Deseadan)
Brontornis burmeisteri Moreno and Mercerat, 1891 Argentina, Province,Santa Cruz (Lago Lower to Middle Miocene o
Rostrornis floweri Moreno Mercerat, 1891 Argentina, Monte León, Monte (Santacrucian) Agnolin, 2007
Brontornis platyonyx Ameghino, 1895 Observación, Karaiken, La Cueva,
Rio Gallegos.

Phorusrhacids: the Terror Birds 191

that need further clarification in these birds. The SYSTEMATIC PALEONTOLOGY
following institutional abbreviations are used in
this chapter: DGM, Divis~o de Geologia e Miner-
a Phylogenetic analysis
alogia do Departamento Nacional da Produ¸ ~o ca
Here we present a cladistic analysis based on 61
Mineral, Rio de Janeiro, Brazil; FMNH, Field
characters (see Appendix 7A); three are multistate,
Museum of Natural History, Chicago, USA;
two of them (i.e., 4 and 14) can be arranged as a
´
MHNT, Museu de Historia Natural de Taubate, ´
morphological series and were treated as ordered.
´
Taubate, Brazil; UF, University of Florida,
Damaged or absent structures are coded as miss-
Gainesville, USA.
ing-data, and all characters are weighted equally.
The character matrix (see Appendix 7B) was con-
structed using the software NDE 0.5.0 and sub-
GEOLOGICAL SETTING mitted to character optimization and parsimony
analysis in PAUPÃ 4.0b (Swofford, 2001). A heu-
Remains of Phorusrhacidae have been found in ristic search was conducted using the Tree Bisec-
continental beds of a great variety of Cenozoic tion Reconnection (TBR) algorithm with 1000
strata in South America, from all epochs of the replicates in the branch-swapping cycles with ran-
Tertiary. No species has been described for the dom addition of the taxa. The maximum saved
Eocene although a few fragmentary specimens trees (maxtress) were automatically increased by
for this epoch have been attributed to Psilopter- 100 when necessary, and branches were collapsed
inae (Acosta Hospitaleche Tambussi, 2005). if the minimum branch length was equal to zero.
Most of the fossil remains and described taxa of Bremer support was also calculated in PAUPÃ
phorusrhacids have been found in Argentina but based on creation of constrains for the clades result
they are also recorded in Uruguay and Brazil, in the previous analysis; this process was con-
and recently an ungual phalanx was recovered ducted with 10 replicates in order to optimize
from Oligo-Miocene deposits in Peru (Shockey computer time. The cladistic analysis was rooted
et al., 2006). There is also an important record of using the taxon Anseranas semipalmata. A total
a beak fragment from Seymour Island, Antarc- of 16 out of the18 species traditionally included in
tica (Case et al., 1987) that is certainly a man- Phorusrhacidae (Alvarenga H€fling, 2003; Ber-
o
dibular symphysis of a huge phorusrhacid, telli et al., 2007) were included in this analysis and
closely related to the Brontornis genus (Alvar- only the taxa Paleopsilopterus itaboaiensis Alvar-
enga H€fling, 2003) from the La Meseta For-
o enga, 1985 and Psilopterus affinis (Ameghino,
mation (Upper Eocene). In North America, 1899) were excluded due to the relatively meager
likely as a result of the Great American Biotic quality of the known material.
Interchange (GABI) that occurred during Paleopsilopterus itaboraiensis is similar in
the Tertiary, we have a single species, Titanis size to Procariama simplex and the morphology
walleri Brodkorb, 1963, from the Pliocene of of its tibiotarsus and tarsometatarsus is very sim-
Florida and Texas (Chandler, 1994; Baskin, ilar to that of other small Phorusrhacidae with
1995; Gould Quitmyer, 2005; MacFadden et long tarsometatarsi (Figure 7.2). In addition to
al., 2006). Claims supporting the existence of material previously described (Alvarenga, 1985),
European Phorusrhacidae have been discarded we can attribute five ungual phalanges to this
by Alvarenga H€fling (2003). In Table 7.1, a
o species (MHNT:5316–5320) (Figure 7.2J–N). R.
brief summary of the chronology of each site for Silva Santos collected this material during the
each species of Phorusrhacidae is given, as well ı
1970s, in the Itabora Basin, RJ, Brazil, which is
as the bibliographic references that may add the type locality of the species. The specimens of
details of the paleoenvironment and associated Psilopterus affinis suggest affinities with other
fauna for each species. species of Psilopterinae (Alvarenga H€fling, o

192 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLI

Fig. 7.1 Strict consensus cladogram resulting from the 48 most parsimonious trees from the present cladistic analysis
(length: 91; CI: 0.7; RI: 0.83). Numbers in nodes express the Bremer support.

2003; Agnolin, 2006). We have included both taxa (Appendix 7B) is shown in Figure 7.1. The group
tentatively in the subfamily Psilopterinae accord- traditionally named “Cariamae” is here phyloge-
ing to the features indicated by Alvarenga netically deﬁned as all clades descending from
H€ﬂing (2003).
o the common ancestor of Cariama and Phorusr-
The strict consensus of the 48 most parsimo- hacidae. A good number of other birds from the
nious trees resulting from analysis of the matrix Paleogene of North America, many of them


Fig. 7.2 Paleopsilopterus itaboraiensis: The
holotype right tarsometatarsus (MNRJ-
4040-V) in proximal (E), dorsal (F), lateral
(G), plantar (H), and medial (I) views. Referred
right tibiotasus (in the left side) and left
tibiotarsus (in the right side) in ventral
(A), lateral (B), medial (C), and distal (D)
views. Five ungual phalanges (MHNT-
5316-5320) from the same locality of the
holotype are tentatively attributed to the
digit III (J and K), digit II (L), digit IV (M) and
digit I (N) of the same species.

attributed to the “Bathornithidae” and “Idiornit- monophyletic and possibly broader Cariamidae
hidae” or “suborder Cariamae” (Wetmore, 1944; family.
Cracraft, 1968, 1971, 1973; Mourer-Chauvire, ´
´
1983; Mayr Mourer-Chauvire, 2006; Mayr,
The monophyly of the phorusrhacidae
2007), are evidently close to the extant Cariami-
dae (Olson, 1985). In fact, a revision of these birds Andrews (1896, 1899) was the ﬁrst to recognize
is necessary to deﬁne the basal cladistic position the close relationship between Phorusrhacidae
of the species of Cariamae, as well as establish a and the extant Cariamidae. There is a great


deal of confusion, however, in terms of which Miocene of South America. The phylogenetic
genera should be included in these families. analysis presented here recovers Brontornis in
Brodkorb (1967) included Psilopterus, Procar- the Phorusrhacidae, close to the other large
iama, and Mesembriornis in Cariamidae, caus- members of this clade and relatively distant
ing confusion for many subsequent authors. The from the Anseriformes. We believe that
extant Cariamidae are good fliers while the Phor- the particular diverted medial condyle of the
usrhacidae form a more derived group that is tibiotarsus may be an adaptation due to the
wholly flightless. The phylogenetic analysis pre- excessive weight of the bird, influencing its
sented here firmly establishes the systematic posture and walk. Significantly, this particular
position of these groups. character appears in Andalgalornis (Noriega
Another question regarding the monophyly Agnolin, 2008, figure 6-B) and also in Cariama.
of the Phorusrhacidae concerns Brontornis, the The quadrate bone mentioned by Angolin (2007)
largest member of the group. In Brontornis is fragmented to the point where meaningful
burmeisteri the internal condyle of the tibio- comparisons are impaired. An important char-
tarsus is medially diverted, in a way typical of acter for Brontornis can be seen in the thoracic
Anseriformes, and this is evident in at least two vertebra shown by Moreno Mercerat (1891,
specimens figured by Moreno Mercerat plate VII, figures 1 and 2). There is a large
(1891). Based on this feature and the morphol- recessus pneumaticus in the mid-centrum
ogy of an incomplete quadrate bone, Agnolin (character 22; Appendix 7A), which is a very
(2007) proposed recognizing Brontornis as a clearly defined character in the Phorusrhacidae
giant anseriform convergent on the Gastor- (Figure 7.3) and is well illustrated in the liter-
nithidae of the Northern Hemisphere and the ature for Psilopterus (Sinclair Farr, 1932,
Dromornithidae of Australia but coexisting figures 7 and 9, plate XXXI) and Titanis
with the Phorusrhacidae and ratites in the (Gould Quitmyer, 2005, figure 6-C).

Fig. 7.3 A thoracic vertebra of Paraphysornis brasiliensis (DGM-1418-R) in lateral (A), cranial (B), and dorsal (C) views.
Important anatomic similarities to Brontornis burmeisteri can be observed: the large recessus pneumaticus (A) in the
mid-centrum (character 22) is a diagnostic feature for phorusrhacids.


Taxonomic hierarchy flightless rails such as the Weka (Gallirallus aus-
tralis) the Takahe (Porphyrio mantelli) (Taylor,
Aves Linnaeus, 1758
1996) and among extant cariamas (Alvarenga, per-
Cariamae F€ rbringer
u
sonal observation), all taxa phylogentically close to
Phorusrhacidae Ameghino, 1889
Phorusrhacidae.
Diagnosis – The cladistic analysis offered here
provides a diagnosis of Phorusrhacidae based on 13
synapomorphies, of which nine are unambiguous
ANATOMY
and two are exclusive:
upper beak tip strongly curved (character 2);
There are many described and illustrated fossils of
palate desmognathous (character 3);
the Phorusrhacidae. The species of Psilopterus,
large temporal fossa almost meet at median line
the smallest phorusrhacid, are known from
(character 7);
complete or almost complete skeletons and are
foramen magnum oriented caudally (character 8);
particularly well described and illustrated
processus basipterygoid present (character 10);
(Sinclair Farr, 1932). Procariama simplex is
the pterygoid with articulation for the processus
also represented by a nearly complete skeleton
basipterygoid (character 11);
(FM–P14525), partially described by Alvarenga
processus zygomaticus present (character 13);
H€fling (2003). Unfortunately the larger species
o
acrocoracoidal process absent (character 29);
are known by much less complete specimens.
tuberculum ventrale of humerus projected proxi-
Among the larger phorusrhacids, Paraphysornis
mally (character 36);
brasiliensis (Alvarenga, 1982) is the best repre-
diaphysis of humerus is bowed and not in “sigma”
sented with about 70% of the skeleton available
(character 38);
for one specimen). As can be seen in Appendix 7A,
processus flexorius of humerus projected distally
a good number of anatomic characters can be
(character 39);
determined for the phorusrhacids but in spite
trochanter majus of the femur absent or not prom-
of the relative abundance of phorusrhacid fossils,
inent (character 49);
some important anatomic questions still persist.
trochlea metatarsi II (in dorsal view) not deflected
medially (character 59).
Sexual dimorphism – Alvarenga H€fling o
Cervical vertebrae
(2003) commented on the intraspecific differences
of size within the Phorusrhacidae and highlighted a Several specimens of Phorusrhacidae show an
variation of 33% in the size of the tarsometatarsus osseous bridge from the processus transversus
between two specimens of Brontornis burmeisteri to the middle of the corpus vertebrae, forming
as well as in the specimens attributed to Psilop- large dorsal fenestrae (Mayr Clark, 2003, char-
terus australis by Sinclair Farr (1932). The latter acter 52 and 53; Sinclair Farr, 1932, plate
taxon also suggests important intraspecific differ- XXXI; Patterson Kraglievich, 1960, figures 4
ences in the overall size and height of the maxilla. and 5). Noriega et al. (2009) illustrated a cervical
Gould Quitmyer (2005) summarized all the vertebra (certainly close to C-10) attributed to
material referred to Titanis walleri and highlighted Devincenzia pozzi, where this character is pres-
an important difference in the size of two quad- ent. In Paraphysornis brasiliensis, complete
ratojugals (UF 57580 and UF 57585) and two prox- vertebrae such as C3 and possibly C10 or C11
imal phalanges of the pedal digit III (UF 30001 and do not present this character (Figure 7.4). An
UF 171382). These differences may well be the examination of other existing vertebral frag-
expression of sexual dimorphism, which if con- ments also fail to confirm its presence. This
firmed would likely be female biased. Males may character appears to be absent in Paraphysornis
have been larger than females, as in the case of large and Brontornithinae.


Fig. 7.4 Some complete cervical vertebrae of Paraphysornis brasiliensis (DGM-1418-R) in dorsal views: 3rd cervical
(A), possibly the 10th cervical (B), and possibly the 11th cervical (C). The absence (at least in these vertebrae) of bridges
linking the processus transversus to the middle of corpus of the vertebrae, forming dorsal fenestras, may be an
important feature.

Thoracic vertebrae uncinate processes were present but not fused to
the ribs and were subsequently lost during fossil-
In addition to the recessus pneumaticus in
ization. We observe that while preparing the ske-
the middle centrum of some of the pre-synsacral
letons of some birds such as Psophiidae, Aramidae
thoracic vertebrae, there is another feature that
and some Rallidae (Rallus, Pardirallus) by macer-
relates to the processus dorsalis of the thoracic
ation, the uncinate processes are completely
vertebrae. This process is very tall in all complete
released and not fused to the ribs. If uncinate
specimens of thoracic vertebrae (Patterson
processes were present in the large phorusrhacids
Kraglievich, 1960, figure 6; Gould Quitmyer,
it seems unlikely that they were fused to the ribs
2005, figure 6). In Paraphysornis brasiliensis, a
(character 24; Appendix 7A).
fragment of processus dorsalis attributed to the
first pre-synsacral vertebra, as well as a crest that
Clavicles
we identified as a cranial extremity of the iliac
dorsalis crest (Figure 7.5), suggest a shorter pro- Within the Phorusrhacidae, the cranial tip
cessus dorsalis in the thoracic vertebrae, of Para- (extremitas omalis claviculae) of the coracoid
physornis. It may also be a feature of the is known to be fused to the clavicles only in
Brontornithinae. Mesembriornithinae (Rovereto, 1914). Neither
free clavicles nor a furcula are known for any
other representatives of the family. In Paraphy-
Uncinate process in ribs
sornis brasiliensis, a bone fragment not origi-
In the illustrations and descriptions of Sinclair nally described (Alvarenga, 1982) seems to
Farr (1932), the ribs of the Phorusrhacidae lack belong to the cranial extremity of the left clav-
uncinate processes. However, it is possible that icle (Figure 7.6).


Fig. 7.5 A reconstruction of the first pre-sinsacral thoracic vertebra in lateral view, close to the remains of the cranial
extremity of the dorsal iliac crest from Paraphysornis brasiliensis (DGM-1418R). This reconstruction suggests a short
processus dorsalis for Paraphysornis.

Pubis ending at around its medium portion. Sinclair
Farr (1932) similarly described the pubis in Psilop-
Andrews (1899) describes the pubis of Patagornis
terus, concluding that “the absence of the posterior
marshi as limited to its cranial portion, projecting
pubic projection, if subsequently confirmed, should
from the floor of the acetabulum as a bar closing the
prove to be a good diagnostic character, perhaps of
foramen obturatum, bordering the ischium and
ordinal value.” However, in a specimen of

Fig. 7.6 Possibly the left clavicle of Paraphysornis brasiliensis (DGM-1418R) from lateral (left) and medial (right)
views.


Fig. 7.7 Pelvis of Procariama simplex (FM- P14525) in lateral (A) and ventral (B) views. The most cranial portion of the
pubis (arrow) is closed and delimits the foramen obturatum; the most caudal portion of the pubis is articulated with the
ischiun, and is not continuous with the cranial portion.

Procariama simplex (FM-P14525), the nearly com-
plete pelvis preserves the pubis with both the prox-
imal and distal extremities (Figure 7.7), and a
delicate medium portion adhered to the ventral
surface of the ischium. Such a design is similar to
that of the Accipitridae and some Falconidae. The
distal extremity of the pubis may certainly have
been lost or not identified in Patagornis and Psilop-
terus in these cases. Pelvis fragments from Para-
physornis brasiliensis (DGM-1014), not previously
described by Alvarenga (1982), preserve the cranial
portions or the ischium (Figure 7.8) and the caudal
projections of the two pubes (Figure 7.9). Their
morphology is very similar to that observed in
Procariama. The pubis of Phorusrhacidae may be
defined as discontinuous, with the medial filamen-
tous portion adhering to the ventromedial surface of
the ischium. The caudal extremity of the pubis
articulates with the ischium (as in the Accipitridae).
In spite of these conclusions, we believe that some
variations, such as an open or closed obturator
foramen, may occur within the Phorusrhacidae.

TAXONOMY
Fig. 7.8 (A) Cranial fragment of the ischium of Para-
physornis brasiliensis (DGM-1418-R) in ventral view.
Alvarenga H€fling (2003) proposed the alloca-
o The detail of the right ischium (B) shows a pubis segment
tion of 17 species of Phorusrhacidae among five as a branch adhering to the ventral portion o the ischium
subfamilies. Recently, Bertelli et al. (2007) have (arrow).


Fig. 7.9 Caudal segments of the two pubes of Paraphysornis brasiliensis (DGM1418-R); the conformation is similar to
that seen in Procariama simplex.

described Kelenken guillermoi, an exceptionally 2 Subfamily Psilopterinae (Dolgopol de Saez,
large specimen from the Middle Miocene of Argen- 1927). (Diagnosed by the medial expansion of
tina as an additional genus and species. Its addition the articular surface of the trochea metatarsi II
to the family brings the total to 13 genera and 18 of the tarsometatarsus – character 59).
species (Table 7.1). The updated phylogenetic Genus Psilopterus (Moreno Mercerat, 1891)
analysis continues to support the five subfamilies P. bachmanni (Moreno Mercerat, 1891;
proposed by Alvarenga H€fling (2003) even
o Figure 7.10C)
though the cladogram sensu stricto does not sep- P. lemoinei (Moreno Mercerat, 1891)
arate the subfamilies Brontornithinae, Phorusrha- P. affinis (Ameghino, 1899)
cinae, and Patagornithinae. These taxa can be P. colzecus (Tonni Tambussi, 1988)
diagnosed by the characters described by the Genus Procariama (Rovereto, 1914)
authors cited above. Given this, we propose the P. simplex (Rovereto, 1914)
following family structure, from the more basal to Genus Paleopsilopterus (Alvarenga, 1985)
the more derived groups. P. itaboraiensis (Alvarenga, 1985)
3 Subfamily Patagornithinae (Mercerat, 1897).
(Diagnosed by Alvarenga H€fling (2003) as
o
medium-sized, smaller, and slimmer than the
Family phorusrhacidae
Phorusrhacinae, with a long and narrow man-
1 Subfamily Mesembriornithinae (Kraglievich, dibular symphysis, and a long and slender tar-
1932). (Diagnosed by fusion of the coracoid to sometatarsi that is more than 70% of the length
the clavicle – character 31.) of the tibiotarsus.)
Genus Mesembriornis (Moreno, 1889) Genus Patagornis (Moreno Mercerat, 1891)
M. milneedwardsi (Moreno, 1889; Figure 7.10B) P. marshi (Moreno Mercerat, 1891)
M. incertus (Rovereto, 1914) Genus Andrewsornis (Patterson, 1941)


Fig. 7.10 Reconstructions of some phorusrhacids compared to the extant Cariama. (A) Cariama cristata;
(B) Mesembriornis milneedwardsi; (C) Psilopterus bachmanni; (D) Andalgalornis steuletti; (E) Phorusrhacus long-
issimus; (F) Paraphysornis brasiliensis; and (G) Brontornis burmeiteri. A man’s silhouette (1.75 m) is used as scale.
(Drawing by Eduardo Brettas.)

A abbotti (Patterson, 1941) D. pozzi (Kraglievich, 1931)
Genus Andalgalornis (Patterson Kraglievich, Genus Kelenken (Bertelli et al., 2007)
1960) K. guilermoi (Bertelli et al., 2007)
A steulleti (Kraglievich, 1931; Figure 7.10D) Genus Titanis (Brodkorb, 1963)
4 Subfamily Phorusrhacinae (Ameghino, 1889). T. walleri (Brodkorb, 1963)
(Diagnosed by Alvarenga H€fling, (2003) as
o 5 Subfamily Brontornithinae (Moreno
gigantic; with a mandibular symphysis that is Mercerat, 1891). (Diagnosed by Alvarenga
relatively long and narrow but shallow, and H€fling (2003) as gigantic; the mandibular sym-
o
more than twice as long as the width of the physis is proportionally shorter, wider, and
base; tarsometatarsus is relatively long and higher than other Phorusrhacidae; the tarso-
slender, and is always longer than 60% of the metatarsus is proportionally short, widened,
tibiotarsus.) and flattened dorso-ventrally; also it is possible
Genus Phorusrhacos (Ameghino, 1889) that the condition of character 58 represents a
P. longissimus (Ameghino, 1899; Figure 7.10E) synapomorphy to this subfamily.)
Genus Devincenzia (Kraglievich, 1932) Genus Brontornis (Moreno Mercerat, 1891)


B. burmeisteri (Moreno Mercerat, 1891; ern continents were much closer together. In the
Figure 7.10G) Oligocene, extreme cooling of the planet and sub-
Genus Physornis (Ameghino, 1895) sequent lowering of sea levels might have facili-
P. fortis (Ameghino, 1895) tated further movements. It is also possible that
Genus Paraphysornis (Alvarenga, 1993; some birds made reverse movements but, later in
Figure 7.10F) the Tertiary, movement may have been restricted
P. brasiliensis (Alvarenga, 1982) by further vicarious geographic effects.

BIOGEOGRAPHY AND THE ORIGIN ACKNOWLEDGMENTS
OF THE PHORUSRHACIDAE
c
We thank Rafael Migotto, Ricardo Mendon¸ a, and
The biogeographical history of the Cariamae Graziella Couto-Ribeiro, from the Museu de His-
remains unclear but several genera and species ´ ´
toria Natural de Taubate, Brazil for important help
have been described from Eocene and Oligocene in the initial phase of this paper, including on
deposits in North America (Wetmore, 1944, 1967; phylogenetic analysis and editing of pictures. To
Cracraft, 1968, 1971, 1973; Olson, 1985) and Gary Kaiser and Gareth Dyke for the important
´
Europe (Mourer-Chauvire, 1983; Mayr, 2007, help in the revision on the final version of the
2009) although the interpretation of these birds manuscripts and also in the final treatment of the
as a monophyletic group needs to be re-examined. figures of this chapter. Finally, to Eduardo Brettas
They disappear from the fossil record of both areas for the excellent artistic reconstructions of some
in the Miocene. The Cariamidae, the sister group phorusrhacids in Figure 7.10.
to the Phorusrhacidae, appear to have been almost
absent from South America until the mid-
Tertiary. There are only two known representa-
APPENDIX 7A: CHARACTER LIST AND
tives, both from Argentina: Chunga incerta from
CHARACTER STATES USED FOR THE
the Late Miocene (Tonni, 1974) and Cariama san-
PRESENT CLADISTIC ANALYSIS
tacrucensis from the Early–Middle Miocene
(Noriega et al., 2009).
Skull and mandible
Members of the Phorusrhacidae and also an
unpublished Cariamae, closely related to the 1 Upper beak, maxilla and praemaxilla: wider
European Idiornithidae (Alvarenga, personal than tall (0); taller than wide (1).
observation), were present in South America dur- 2 Upper beak, praemaxilla tip: straight or slight
ing the Paleocene. They must represent the South curved (0); strongly curved (1).
American portion of a stock of Cariamae that 3 Palate: squizognathous (0); desmognathous (1).
enjoyed an early (even Cretaceous) diversity in Eur- 4 Rostral border of antorbital fenestrae: strongly
ope, North America and certainly Africa. Claims obliquous (0); obliquous (1); almost vertical (2).
supporting the presence of phorusrhacids in Europe 5 Neurocranium: wider than tall (0); taller than
´,
(Mourer-Chauvire 1983; Peters, 1987) were dis- wider (1).
cussed and discarded by Alvarenga H€fling, 2003.
o 6 Os lacrimale (in adult): not ankilosed to frontal
It is possible that Phorusrhacidae and also Car- (0); ankilosed to frontal (1).
iamidae arose in South America, but unfortunately 7 Temporal fossa: small (0); large – almost meet
it will be necessary to find new fossil evidence to up in median line (1).
provide direct support for reconstruction of the 8 Foramen magnum oriented: ventrally or ven-
biogeographic history of these birds. They might trally-caudal (0); caudally (1).
have arrived in South America from Europe by 9 Processus supraorbitales of lacrimale: short (0);
traveling through Africa at a time when the south- caudally long (1).


10 Processus basipterigoid: absent or very small 27 Coracoid – pneumatic foramina in dorsal surface of
(0); present (1). extremitas sternalis: absent (0); present (1).
11 Os pterigoid with articulation for the proces- 28 Procoracoidal process: present (0); absent (1).
sus basipterigoid: absent (0); present (1). 29 Acrocoracoidal process; present (0); absent (1).
12 Premaxillare nasal process (in adult): not com- 30 An osseous bridge linking the acrocoracoidal
pletelly fused (0); completelly fused (1). to the procoracoidal process: absent (0);
13 Processus zigomaticus: present (0); absent (1). present (1).
14 Jugal bar very tall (the hight is two times or 31 Coracoid fused to claviculae: absent (0); present (1).
more than the wide): absent (0); tall two times 32 Coracoid articular facet for the scapula: an
(1); tall more than two times (2). excavated cotila (0); not a cotila (1).
15 Mandibulae – pars symphysialis: lenght equal 33 Scapula, acromion cranially projected; absent (0);
or bigger than one quarter of the mandibulae: present (1).
absent (0); present (1). 34 Scapula corpus: curved (0); straight (1).
16 Mandibulae – pars symphysialis: longer than 35 Scapula pneumatic foramen: present (0);
than wide, strong and massive: absent(0); pres- absent (1).
ent (1).
17 Mandibulae – pars symphysialis: straight or
Thoracic limb
ventrally curved (0); dorsally curved (1).
18 Fenestra caudalis mandibulae: absent (0); 36 Humerus – tuberculum ventrale projected prox-
present (1). imally (more than caput humeri); absent (0);
19 Fenestra rostralis mandibulae:absent (0); present (1).
present (1). 37 Humerus – pneumatic foramen: absent or very
20 Mandibulae – processus retroarticularis: small (0); large (1).
absent or small (0); present and large (1). 38 Humerus – diaphysis in anconal view: double
curve in “sigma” (0); one curve with concavity
anconal and medial (1).
Vertebral column and ribs
39 Humerus – processus flexorius projected distally:
21 Third cervical vertebrae – an osseous bridge absent (0); present (1).
linking the processus transversus to processus 40 Ulna length: equal or longer than the humerus
articularis (pos-zygapophysis) making a dorsal (0); shorter than the humerus (1).
fenestrae (see Mayr Clarke, 2003 – char. 52): 41 Carpometacarpus – distal end of metacarpale
absent (0); present (1). minus (see Alvarenga H€fling, 2003, Fig. 6):
o
22 Thoracicae vertebrae – a large recessus pneuma- same level of metacarpale majus (0); shorter than
ticus in the mid centrum (see Livezey Zusi, 2006 metacarpale majus (1).
– char. 0850): absent (0); present (1). 42 Carpometacarpus – os metacarpale minus
23 Presinsacral vertebrae form a notarium: absent (shaft) (see Mayr Clarke, 2003, character 85):
(0); present (1). almost parallel to metacarpale majus (0);
24 Ribs – processus uncinati: absent or unfused to bowed (1).
ribs (0); fused to ribs (1).
Pelvic girdle
Thoracic girdle
43 Pelvis elongated and compressed laterally:
25 Coracoid – ?pneumatic foramen directly absent (0); present (1).
below facies articularis scapularis (see Mayr 44 A strong transversal crest supracetabularis ilii:
Clarke, 2003 –char. 66): absent (0); present (1). absent (0); present (1).
26 Coracoid – foramen nervi supracoracoidei: 45 Pars preacetabularis ilii: fused only in the top of
present (0); absent (1). spinous process of synsacral vertebrae (0); fused


in the top and lateral face of spinous process of 55 Tarsometatarsus long and slender: the ratio of
synsacral vertebrae (1). total length/width of middle of diaphysis is
46 Pubis incomplete: absent (0); present (1). smaller than 12 (0); the ratio is bigger than 12 (1).
56 Tarsometatarsus – facies dorsalis excavated (an
evident longitudinal sulcus): absent (0); present
Pelvic limb
(1).
47 Femur length: shorter than tarsometatarsus 57 Tarsometatarsus – hypotarsus with well-devel-
(0); equal or longer (1). oped crista/sulci (see Mayr Clarke, 2003,
48 Femur length: shorter than humerus (0); equal or character 103): absent (0); present (1).
longer (1). 58 Tarsometatarsus (dorsal view) – articular surface
49 Femur – trochanter majus: prominent proximally of middle trochlea – a dorsomedial expansion (see
(0); absent or not prominent (1). Alvarenga H€fling, 2003, figure 8): absent (0);
o
50 Femur – fossa poplitea: shallow (90); deep (1). present (1).
51 Tibiotarsus – distal rim of condylus medialis 59 Tarsometatarsus (dorsal view), trochlea metatarsi
distinctly notched (see Mayr Clarke, 2003, II: deflected medially (0); almost parallel to the
char. 102): absent (0); present (1). trochlea III (1); articular surface extended medi-
52 Tibiotarsus – condylus medialis medially de- ally (2).
flected: absent (0); present (1). 60 Tarsometatarsus (distal view), trochlea meta-
53 Tarsometatarsus proportions: larger than 60% tarsi II: not deflec ted plantarly (0); deflected
the length of tibiotarsus (0); smaller (1). plantarly (1).
54 Tarsometatarsus strong and short: ratio of 61 Tarsometatarsus (dorsal view), trochlea
total length/width middle of diaphysis is bigger metatarsi IV, a longitudinal sulcus: present (0);
than 6 (0); the ratio is smaller than 6 (1). absent (1).

47
46
45
44
43
42
41
40
39
38
37
36
35
34
33
32
31
30
29
28
27
26
25
24
23
22
21
20
19
18
17
16
15
14
13
12
11
10
9
8
7
6
5
4
3
2
1
APPENDIX 7B: CHARACTER MATRIX
204

Character
Taxon
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
Anseranasse- 0 0 1 1 1 0 0 1 0 1 1 0 1 1 0 0 0 0 1 1 1 0 1 1 1 0 1 0 0 0 0
mipalmata
Ardeotis kori 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 1 0 1 0 0 0 0 0
Opisthocomus 0 0 0 1 1 ? 0 0 0 0 0 1 0 0 0 0 0 1 1 1 1 0 1 1 1 ? 1 0 0 0 1
hoazin
Cariama cristata 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 1 0 0 0 1 0
Elaphrocnemus 0 0 0 1 0 0 0 0 1 0 0 1 1 ? ? ? ? ? ? ? ? ? ? ? 0 1 0 1 0 0 0
phasianus
Bathornis grallator 0 0 0 0 0 1 0 0 0 ? ? 0 1 0 0 0 0 0 0 0 ? ? ? ? ? ? 0 ? ? ? ?
Brontornis ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 ? 1 ? ? 1 ? ? ? ? ? ? ? ? ?
burmeisteri
Physornis fortis ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 ? 1 ? ? ? ? ? ? ? ? ? ? ? ?
Paraphysornis 1 1 ? ? ? 0 ? ? 1 ? 1 ? ? 1 1 1 1 1 1 0 0 1 0 0 0 1 0 1 1 0 0
brasiliensis
Phorusrhacos 1 1 ? 1 ? ? ? ? 1 ? ? ? ? 1 1 1 1 1 1 0 ? 1 ? ? ? ? ? ? ? ? ?
longissimus
Devincenzia pozzii 1 1 1 1 0 0 1 1 1 1 1 ? 0 2 1 1 1 ? ? ? ? 1 ? ? ? ? ? ? ? ? ?
Kelenken guilhermoi 1 1 1 1 0 ? 1 1 ? ? ? ? 0 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?
Titanis walleri ? ? ? ? ? ? ? ? ? ? 1 ? ? 1 ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ?
Patagornis marshi 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 0 1 0 ? 1 ? ? 0 1 0 1 1 0 0
Andrewsornis abbotti 1 1 1 1 ? ? ? ? ? ? ? ? ? 1 1 1 1 1 1 0 ? ? ? ? ? ? ? ? ? ? ?
Andalgalornis steulleti 1 1 1 2 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 0 1 1 0 0 ? ? ? ? ? ? ?
Psilopterus bachmanni 1 1 1 0 0 0 1 1 1 1 1 1 0 0 0 1 1 1 1 0 1 1 0 0 0 1 0 1 1 0 0

Psilopterus lemoinei 1 1 1 1 0 0 1 1 1 1 1 ? 0 0 0 1 1 1 1 0 1 1 0 0 0 1 0 1 1 0 0
Psilopterus colzecus ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 ? ? ? ? 1 ? ? ? ? ? ? ? ? ?
Procariama simplex 1 1 1 1 0 0 1 1 1 1 1 1 0 0 0 1 1 1 1 0 ? 1 ? ? 0 1 0 1 1 0 0
Mesembriornis 0 ? 1 1 0 0 1 1 ? ? ? 1 0 0 0 0 1 1 1 ? 1 1 0 0 0 1 ? 1 1 0 1
milneedwardsi
Mesembriornis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 0 1 ? 1 1 0 1
incertus

(Continued)

Character
Taxon
32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61
Anseranas semipalmata 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 1 0
Ardeotis kori 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 1
Opisthocomus hoazin 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 1
Cariama cristata 1 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 0 0 1
Elaphrocnemus phasianus 1 ? ? ? 0 0 0 0 1 0 1 ? ? ? ? 0 1 0 1 1 0 0 0 1 1 0 0 0 0 0
Bathornis grallator ? ? 0 1 ? ? 0 ? 1 0 1 ? ? ? ? 0 1 ? ? 0 0 0 0 1 1 ? 0 0 0 ?
Brontornis burmeisteri ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? ? 1 ? 1 1 1 0 1 0 1 1 0 1
Physornis fortis ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?
Paraphysornis brasiliensis 1 1 1 1 1 1 1 1 1 1 1 ? ? ? 1 1 1 1 1 1 0 1 1 0 1 0 1 1 0 1
Phorusrhacos longissimus ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? 0 1 1 1 1 0 0 0 0 1 0 0 1 0 1
Devincenzia pozzii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 ? 1 0 0 0 0 1 ? 0 1 0 1
Kelenken guilhermoi ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 1 0 0 1 0 1
Titanis walleri ? ? ? ? ? ? ? ? ? 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 1
Patagornis marshi 1 1 1 1 ? ? 1 1 1 0 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 0 1 0 1
Andrewsornis abbotti ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ? ? ?
Andalgalornis steulleti ? ? ? ? ? ? ? ? ? ? ? 1 1 1 1 ? ? ? 1 ? 1 ? ? ? ? ? ? ? ? ?
Psilopterus bachmanni 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 0 2 0 1
Psilopterus lemoinei 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 0 2 0 1
Phorusrhacids: the Terror Birds

Psilopterus colzecus ? ? ? ? ? ? 1 1 1 ? ? ? ? ? ? ? 1 ? ? ? ? 0 ? ? ? ? 0 2 ? 1
Procariama simplex 1 1 1 1 1 1 1 1 1 ? ? 1 1 1 1 0 1 1 1 0 0 0 0 1 1 0 0 2 0 1
Mesembriornis milneedwardsi 1 ? ? ? ? ? 1 1 1 0 1 1 1 1 ? 0 1 1 1 0 0 0 0 1 1 0 0 1 0 1
Mesembriornis incertus 1 ? ? ? 1 1 1 1 1 ? ? ? ? ? ? 0 1 ? ? 0 0 0 0 1 1 0 0 1 0 1
205


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