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 uAt the beginning of the Tertiary, while mammals Pleistocene (Alvarenga et al., 2010). Ratites andwere undergoing an evolutionary explosion, phorusrhacids apparently lived together, perhapsseveral groups of birds developed a tendency to competing for similar terrestrial habitats, duringgigantism. These groups were scattered across the whole of the Tertiary. At that time Southalmost 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 amongity with the Anseriformes (Andors, 1992), have these birds, have survived to present times.been recovered from Paleocene and Eocene depos- Scientiﬁc investigations on the Phorusrhacidaeits 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 aAustralia, another group of giant birds, the Dro- “probable toothless mammal”, which he namedmornithidae, 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 huge2004) had a broad diversity in the mid-Tertiary. bones that he identiﬁed as giant birds from theAmong 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 offound. The Ratitae, also present since the Paleo- Ameghino (1891a,b) recognized Phorusrhacus ascene 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 fragmentsgroup 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 synonymytiary, they are the only present-day survivors. The within the family; currently 14 genera and 18elephant-bird, Aepyornis maximus, a ratite from species are recognized (Table 7.1). A more detailedMadagascar that died out around 700 years ago, is history of the ﬁrst investigations of this family, asprobably the biggest bird ever found (Amadon, well as of the ﬁrst classiﬁcations proposed, is set1947; Wetmore, 1967). out by Alvarenga & H€ﬂing (2003). o The Phorusrhacidae, another group of giants In this chapter, our main objective is to presentwas 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 detailsLiving 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. 188Taxon/reference Geographic locality Horizon/Age Additional referencesMesembriornis milneedwardsi Moreno, 1889 Argentina, Province, Buenos Aires: Upper Pliocene (Montehermosan) o Alvarenga & H€ﬂing, 2003Paleociconia australis Moreno, 1889; Moreno & Monte Hermoso, Rio Loberia Mercerat, 1891Driornis pampeanus Moreno & Mercerat, 1891 (part: only femur)Hermosiornis milneedwardsi Rovereto, 1914Hermosiornis rapax Kraglievich, 1946Prophororhacos australis Brodkorb, 1967Mesembriornis 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 VillarinoProcariama 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 MariaPaleopsilopterus itaboraiensis, Alvarenga,1985 Brazil, Est. Rio de Janeiro: S~o José de a Upper Paleocene (Itaboraian) o Alvarenga H€ﬂing, 2003 Itabora ıPsilopterus bachmanni (Moreno Mercerat, 1891) Argentina, Province, Santa Cruz: Santa Middle Miocene (Santacrucian) o Alvarenga H€ﬂing, 2003Patagornis bachmanni Moreno Mercerat, 1891 Cruz, Lago Pueyrredon, MontePsilopterus communis Moreno Mercerat, 1891 Observación, La CuevaPsilopteus intermedius Moreno Mercerat, 1891Phororhacus delicatus Ameghino, 1891Pelecyornis minutus Ameghino, 1891Pelecyornis pueyrredonensis Sinclair Farr, 1932Psilopterus afﬁnis (Ameghino, 1899) Argentina, Province, Chubut: Cabe¸ a c Middle Oligocene (Deseadan) o Alvarenga H€ﬂing, 2003;Phororhacus afﬁnis Ameghino, 1899 Blanca Agnolin, 2006Psilopterus colzecus Tonni Tambussi, 1988 Argentina, Province, Buenos Aires: Upper Miocene (Chasicoan) o Alvarenga H€ﬂing, 2003 Partido de Villarino HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIPsilopterus lemoinei (Moreno Mercerat, 1891) Argentina, Province, Santa Cruz: Santa Middle Miocene (Santacrucian) o Alvarenga H€ﬂing, 2003;Patagornis lemoinei Moreno Mercerat, 1891 Cruz, Killik Aike, Monte Observación, Degrange Tambussi, 2008Psilopterus australis Moreno Mercerat, 1891; Take Harvey, La Cueva, Corriguen Brodkorb, 1967 Kaik, TaguaquemadaPelecyornis tubulatus Ameghino, 1895Phororhacus modicus Ameghino, 1895Staphylornis gallardoi Mercerat, 1897Staphylornis erythacus Mercerat, 1897Pelecyornis tenuirostris Sinclair Farr, 1932
Patagornis marshi Moreno Mercerat, 1891 Argentina, Province, Santa Cruz;Monte Lower to Middle Miocene. o Alvarenga H€ﬂing, 2003Tolmodus inﬂatus Ameghino, 1891 Observación, Tagua Quemada, La (Santacrucian)Phororhacos inﬂatus Ameghino, 1891; Andrews, Cueva 1899; Brodkorb, 1967Paleociconia cristata Brodkorb, 1967Andrewsornis abbotti Patterson, 1941 Argentina, Province,Chubut (Cabe¸ a c Middle to Upper Oligocene o Alvarenga H€ﬂing, 2003 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, 2008Phororhacos deautieri Kraglievich, 1931Andalgalornis ferox Paterson Kraglievich, 1960Andalgalornis steulleti Brodkorb, 1967Phorusrhacus longissimus Ameghino, 1887 Argentina, Province, Santa Cruz, La Lower and Middle Miocene o Alvarenga H€ﬂing, 2003Phororhacos longissimus, Ameghino, 1889 Cueva, Tagua/Quemada, Monte (Santacrucian)Stereornis rollieri Moreno Mercerat, 1891 Observación, Rio SehuénStereornis gaudryi Moreno Mercerat, 1891Mesembriornis studeri Moreno Mercerat, 1891Mesembriornis quatrefragesi Moreno Mercerat, 1891Darwinornis copei Moreno Mercerat, 1891Darwinornis zittelli Moreno Mercerat, 1891Darwinornis socialis Moreno Mercerat, 1891Owenornis afﬁnis Moreno Mercerat, 1891Owenornis lydekkeri Moreno Mercerat, 1891Phororhacus sehuensis Ameghino, 1891Phororhacus platygnathus Ameghino, 1891Titanornis mirabilis, Moreno Mercerat, 1891 Phorusrhacids: the Terror BirdsCallornis giganteus, Ameghino, 1895Eucallornis giganteus Ameghino, 1901Liornis ﬂoweri Ameghino, 1895Titanis 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) 190Taxon/reference Geographic locality Horizon/Age Additional referencesDevincenzia 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, 2008Devincenzia gallinali Kraglievich, 1932Onactornis depressus Cabrera, 1939Onactornis pozzi Brodkorb, 1967Onactornis mendocinus Brodkorb, 1967Kelenken guillermoi Bertelli, Chiappe Argentina, Province,Rio Negro, Comallo. Midle Miocene (Fm Collón Cur) a Chiappe Bertelli, 2007 Tambussi, 2007Physornis 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, 2003Paraphysornis brasiliensis (Alvarenga, 1982) Brazil, Est. S~o Paulo, Tremembé. a Upper Oligocene or Lower Miocene Alvarenga, 1993Physornis brasiliensis Alvarenga, 1982 (Upper Deseadan)Brontornis burmeisteri Moreno and Mercerat, 1891 Argentina, Province,Santa Cruz (Lago Lower to Middle Miocene o Alvarenga H€ﬂing, 2003;Rostrornis ﬂoweri Moreno Mercerat, 1891 Argentina, Monte León, Monte (Santacrucian) Agnolin, 2007Brontornis platyonyx Ameghino, 1895 Observación, Karaiken, La Cueva, Rio Gallegos. HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLI
Phorusrhacids: the Terror Birds 191that need further clariﬁcation in these birds. The SYSTEMATIC PALEONTOLOGYfollowing institutional abbreviations are used inthis chapter: DGM, Divis~o de Geologia e Miner- a Phylogenetic analysisalogia do Departamento Nacional da Produ¸ ~o ca Here we present a cladistic analysis based on 61Mineral, 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 1000strata 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 savedEocene although a few fragmentary specimens trees (maxtress) were automatically increased byfor this epoch have been attributed to Psilopter- 100 when necessary, and branches were collapsedinae (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 PAUPÃphorusrhacids have been found in Argentina but based on creation of constrains for the clades resultthey 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 optimizefrom Oligo-Miocene deposits in Peru (Shockey computer time. The cladistic analysis was rootedet al., 2006). There is also an important record of using the taxon Anseranas semipalmata. A totala beak fragment from Seymour Island, Antarc- of 16 out of the18 species traditionally included intica (Case et al., 1987) that is certainly a man- Phorusrhacidae (Alvarenga H€ﬂing, 2003; Ber- odibular symphysis of a huge phorusrhacid, telli et al., 2007) were included in this analysis andclosely 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 meagerlikely as a result of the Great American Biotic quality of the known material.Interchange (GABI) that occurred during Paleopsilopterus itaboraiensis is similar inthe Tertiary, we have a single species, Titanis size to Procariama simplex and the morphologywalleri 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 with1995; Gould Quitmyer, 2005; MacFadden et long tarsometatarsi (Figure 7.2). In addition toal., 2006). Claims supporting the existence of material previously described (Alvarenga, 1985),European Phorusrhacidae have been discarded we can attribute ﬁve ungual phalanges to thisby 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 theeach species of Phorusrhacidae is given, as well ı 1970s, in the Itabora Basin, RJ, Brazil, which isas the bibliographic references that may add the type locality of the species. The specimens ofdetails of the paleoenvironment and associated Psilopterus afﬁnis suggest afﬁnities with otherfauna for each species. species of Psilopterinae (Alvarenga H€ﬂing, o
192 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIFig. 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 grouptentatively in the subfamily Psilopterinae accord- traditionally named “Cariamae” is here phyloge-ing to the features indicated by Alvarenga netically deﬁned as all clades descending fromH€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 thenious trees resulting from analysis of the matrix Paleogene of North America, many of them
Phorusrhacids: the Terror Birds 193Fig. 7.2 Paleopsilopterus itaboraiensis: Theholotype right tarsometatarsus (MNRJ-4040-V) in proximal (E), dorsal (F), lateral(G), plantar (H), and medial (I) views. Referredright tibiotasus (in the left side) and lefttibiotarsus (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 theholotype are tentatively attributed to thedigit III (J and K), digit II (L), digit IV (M) anddigit I (N) of the same species.attributed to the “Bathornithidae” and “Idiornit- monophyletic and possibly broader Cariamidaehidae” or “suborder Cariamae” (Wetmore, 1944; family.Cracraft, 1968, 1971, 1973; Mourer-Chauvire, ´ ´1983; Mayr Mourer-Chauvire, 2006; Mayr, The monophyly of the phorusrhacidae2007), are evidently close to the extant Cariami-dae (Olson, 1985). In fact, a revision of these birds Andrews (1896, 1899) was the ﬁrst to recognizeis necessary to deﬁne the basal cladistic position the close relationship between Phorusrhacidaeof the species of Cariamae, as well as establish a and the extant Cariamidae. There is a great
194 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIdeal of confusion, however, in terms of which Miocene of South America. The phylogeneticgenera should be included in these families. analysis presented here recovers Brontornis inBrodkorb (1967) included Psilopterus, Procar- the Phorusrhacidae, close to the other largeiama, and Mesembriornis in Cariamidae, caus- members of this clade and relatively distanting confusion for many subsequent authors. The from the Anseriformes. We believe thatextant Cariamidae are good ﬂiers while the Phor- the particular diverted medial condyle of theusrhacidae form a more derived group that is tibiotarsus may be an adaptation due to thewholly ﬂightless. The phylogenetic analysis pre- excessive weight of the bird, inﬂuencing itssented here ﬁrmly establishes the systematic posture and walk. Signiﬁcantly, this particularposition 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 meaningfulburmeisteri 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 thoracicAnseriformes, 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-centrumogy 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 Phorusrhacidaegiant 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 Titaniswith 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 themid-centrum (character 22) is a diagnostic feature for phorusrhacids.
Phorusrhacids: the Terror Birds 195 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 toPhorusrhacidae Ameghino, 1889 Phorusrhacidae. Diagnosis – The cladistic analysis offered hereprovides a diagnosis of Phorusrhacidae based on 13synapomorphies, of which nine are unambiguous ANATOMYand two are exclusive:upper beak tip strongly curved (character 2); There are many described and illustrated fossils ofpalate 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 areforamen magnum oriented caudally (character 8); particularly well described and illustratedprocessus basipterygoid present (character 10); (Sinclair Farr, 1932). Procariama simplex isthe 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 oacrocoracoidal 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 spitetrochanter 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 differencesof size within the Phorusrhacidae and highlighted a Several specimens of Phorusrhacidae show anvariation of 33% in the size of the tarsometatarsus osseous bridge from the processus transversusbetween two specimens of Brontornis burmeisteri to the middle of the corpus vertebrae, formingas 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, platetaxon also suggests important intraspeciﬁc differ- XXXI; Patterson Kraglievich, 1960, ﬁgures 4ences in the overall size and height of the maxilla. and 5). Noriega et al. (2009) illustrated a cervicalGould Quitmyer (2005) summarized all the vertebra (certainly close to C-10) attributed tomaterial 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, completeratojugals (UF 57580 and UF 57585) and two prox- vertebrae such as C3 and possibly C10 or C11imal phalanges of the pedal digit III (UF 30001 and do not present this character (Figure 7.4). AnUF 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. Thisﬁrmed would likely be female biased. Males may character appears to be absent in Paraphysornishave been larger than females, as in the case of large and Brontornithinae.
196 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIFig. 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 bridgeslinking the processus transversus to the middle of corpus of the vertebrae, forming dorsal fenestras, may be animportant 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, Aramidaethoracic 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 completelyvertebrae. This process is very tall in all complete released and not fused to the ribs. If uncinatespecimens of thoracic vertebrae (Patterson processes were present in the large phorusrhacidsKraglievich, 1960, ﬁgure 6; Gould Quitmyer, it seems unlikely that they were fused to the ribs2005, ﬁgure 6). In Paraphysornis brasiliensis, a (character 24; Appendix 7A).fragment of processus dorsalis attributed to theﬁrst pre-synsacral vertebra, as well as a crest that Clavicleswe identiﬁed as a cranial extremity of the iliacdorsalis crest (Figure 7.5), suggest a shorter pro- Within the Phorusrhacidae, the cranial tipcessus dorsalis in the thoracic vertebrae, of Para- (extremitas omalis claviculae) of the coracoidphysornis. It may also be a feature of the is known to be fused to the clavicles only inBrontornithinae. 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 toFarr (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).
Phorusrhacids: the Terror Birds 197Fig. 7.5 A reconstruction of the ﬁrst pre-sinsacral thoracic vertebra in lateral view, close to the remains of the cranialextremity of the dorsal iliac crest from Paraphysornis brasiliensis (DGM-1418R). This reconstruction suggests a shortprocessus 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 posteriormarshi as limited to its cranial portion, projecting pubic projection, if subsequently conﬁrmed, shouldfrom the ﬂoor of the acetabulum as a bar closing the prove to be a good diagnostic character, perhaps offoramen obturatum, bordering the ischium and ordinal value.” However, in a specimen ofFig. 7.6 Possibly the left clavicle of Paraphysornis brasiliensis (DGM-1418R) from lateral (left) and medial (right)views.
198 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIFig. 7.7 Pelvis of Procariama simplex (FM- P14525) in lateral (A) and ventral (B) views. The most cranial portion of thepubis (arrow) is closed and delimits the foramen obturatum; the most caudal portion of the pubis is articulated with theischiun, 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 adelicate medium portion adhered to the ventralsurface of the ischium. Such a design is similar tothat of the Accipitridae and some Falconidae. Thedistal extremity of the pubis may certainly havebeen lost or not identiﬁed in Patagornis and Psilop-terus in these cases. Pelvis fragments from Para-physornis brasiliensis (DGM-1014), not previouslydescribed by Alvarenga (1982), preserve the cranialportions or the ischium (Figure 7.8) and the caudalprojections of the two pubes (Figure 7.9). Theirmorphology is very similar to that observed inProcariama. The pubis of Phorusrhacidae may bedeﬁned as discontinuous, with the medial ﬁlamen-tous portion adhering to the ventromedial surface ofthe ischium. The caudal extremity of the pubisarticulates with the ischium (as in the Accipitridae).In spite of these conclusions, we believe that somevariations, such as an open or closed obturatorforamen, 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 segmenttion of 17 species of Phorusrhacidae among ﬁve as a branch adhering to the ventral portion o the ischiumsubfamilies. Recently, Bertelli et al. (2007) have (arrow).
Phorusrhacids: the Terror Birds 199Fig. 7.9 Caudal segments of the two pubes of Paraphysornis brasiliensis (DGM1418-R); the conformation is similar tothat 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 oftina as an additional genus and species. Its addition the articular surface of the trochea metatarsi IIto 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)
200 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLIFig. 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)
Phorusrhacids: the Terror Birds 201 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, andThe 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 helphave been described from Eocene and Oligocene in the initial phase of this paper, including ondeposits in North America (Wetmore, 1944, 1967; phylogenetic analysis and editing of pictures. ToCracraft, 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 the2009) although the interpretation of these birds manuscripts and also in the ﬁnal treatment of theas a monophyletic group needs to be re-examined. ﬁgures of this chapter. Finally, to Eduardo BrettasThey disappear from the fossil record of both areas for the excellent artistic reconstructions of somein the Miocene. The Cariamidae, the sister group phorusrhacids in Figure 7.10.to the Phorusrhacidae, appear to have been almostabsent from South America until the mid-Tertiary. There are only two known representa- APPENDIX 7A: CHARACTER LIST ANDtives, both from Argentina: Chunga incerta from CHARACTER STATES USED FOR THEthe Late Miocene (Tonni, 1974) and Cariama san- PRESENT CLADISTIC ANALYSIStacrucensis from the Early–Middle Miocene(Noriega et al., 2009). Skull and mandible Members of the Phorusrhacidae and also anunpublished Cariamae, closely related to the 1 Upper beak, maxilla and praemaxilla: widerEuropean Idiornithidae (Alvarenga, personal than tall (0); taller than wide (1).observation), were present in South America dur- 2 Upper beak, praemaxilla tip: straight or slighting 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: stronglyope, 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 meetit 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).
202 HERCULANO ALVARENGA, LUIS CHIAPPE, AND SARA BERTELLI10 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 very20 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): o22 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 minus23 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
Phorusrhacids: the Terror Birds 203 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 is46 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 surface49 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); o50 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).
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