(Aves: Tinamiformes, Tinamidae) from the Early-Middle Miocene of Argentina

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Chandler, New Species of Tinamou PalArch’s Journal of Vertebrate Palaeontology, 9(2) (2012)

A NEW SPECIES OF TINAMOU (AVES: TINAMIFORMES, TINAMIDAE) FROM THE EARLY-MIDDLE MIOCENE OF ARGENTINA

Robert M. Chandler*

* Department of Biological and Environmental Sciences, Georgia College and State Uni- versity, Milledgeville, GA 31061, [email protected]

Robert M. Chandler. 2012. A New Species of Tinamou (Aves: Tinamiformes, Tinami- dae) from the Early-Middle Miocene of Argentina. – Palarch’s Journal of Vertebrate Palaeontology 9(2) (2012), 1-8. ISSN 1567-2158. 8 pages + 2 ﬁ gures, 1 table.

Keywords: fossil, Miocene, tinamou, Crypturellus, Argentina

ABSTRACT

A new species of tinamou from the early-middle Miocene (Santacrusian), Santa Cruz Forma- tion of Argentina is named. The new species is approximately 16 million year old and has an afﬁ nity with the modern genus Crypturellus based on the unique characteristics of the humerus, hence, the designation aff. Crypturellus. Fossil species and the zooarchaeological record of modern tinamous are given.

Introduction Among the fossils collected by Brown is an almost complete left humerus of a tinamou. In In 1898-1899, Barnum Brown, assistant curator recent years, tinamou fossils have been collect- in the Department of Vertebrate Paleontology at ed in other Santacrusian (South American Land the American Museum of Natural History, repre- Mammal Age, SALMA) localities from both the sented the AMNH as a member of the third Pa- coastal and northwestern areas (Bertelli & Chi- tagonian expedition led by John Bell Hatcher of appe, 2005) of Santa Cruz Province (ﬁ gure 1). Princeton. As told by Simpson (1984: 118-121), Additional records of this Neotropical family are this was not a happy partnership and eventu- abundant in the Pliocene and Pleistocene depos- ally Brown was left behind in Patagonia. How- its of the Pampean region (Tambussi, 1987) and ever, Brown preferred to be in the ﬁ eld alone and other South American localites (Cuello, 1988), returned to New York with a good collection of and also are known from late Miocene depos- Santacrusian vertebrate fossils. its (Tonni & Tambussi, 1986; Tambussi, 1987).

Map of the general region of the Santa Cruz Province, showing localities (solid circles) where fossil tinamous have been collected. Modiﬁ ed from Marshall (1976) by Linda D. Chandler.

All assignable fossils have been referred to ex- tion (Bertelli & Chiappe, 2005; Chiappe, 1991) tant genera with the exception of one species in northwestern Santa Cruz Province. The Pin- in the poorly represented extinct genus Tina- turas Formation is slightly older in age than the misornis (Brodkorb, 1961; Tonni, 1977; Tam- typical Santacrusian beds of the Santa Cruz For- bussi, 1987). mation (Marshall, et al. 1983; Bown & Fleagle, The middle Miocene (Santacrucian) tinamou 1993). reported here is the earliest record for a tinamou with an afﬁ nity to the genus Crypturellus. Institutional Abbreviations Additional described tinamou fossils have been collected in the middle Miocene formations of AMNH, FAM, American Museum of Natural Santa Cruz Province (Bertelli & Chiappe, 2005; History, Frick American Museum; Chiappe, 1991). Three different localities in the FMNH, Field Museum of Natural History; Santa Cruz Formation have yielded tinamou KU, Charles Bunker Memorial Osteological Col- fossils (ﬁ gure 1). In addition to the humerus of lection, Museum of Natural History, Univer- the new species (named herein) from Cañadón sity of Kansas; de las Vacas, another humerus and several cora- LSU, Museum of Natural Science, Louisiana coids have been found at Monte Observación, State University; and a humerus and a tibiotarsus at Monte León. MACN-SC, Museo Argentino de Ciencias Natu- Furthermore, a tinamid tibiotarsus was found rales, Colección Santa Cruz; in continental deposits of the Pinturas Forma- SDSNH, San Diego Society of Natural History;

UF, Florida Museum of Natural History, Univer- 1988; Houde, 1988), but there is a broad consen- sity of Florida; sus that both Lithornis and galliforms are basal UMMZ, University of Michigan, Museum of Zo- neornithine birds (Olson, 1985; Cracraft, 1988; ology. Houde, 1988; Sibley & Alquist, 1995). The anatomical terminology used in this Geological Ocurrence study is that of Baumel et al. (1979) with the exception that the Latin terms were replaced with The holotype was collected by Barnum Brown the English equivalents. from beds of the Santa Cruz Formation in a Comparative materials used in this study wash at Cañadón de las Vacas, southeastern are as follows: Lithornis plebius (AMNH:FAM Santa Cruz Province, Argentina (fi gure 1). In 21902), L. promiscuus (AMNH:FAM 21903), Crax Brown’s personal Patagonian fi eld notebook globulosa (UF 24052, 24053), Mitu mitu (UF (Brown, 1899, AMNH Archives) he states that 24051), Ortalis garrula (UF 33148), O. polioceph- Cañadón de las Vacas is “40 miles south of San- ala (UF 33147), O. vetula (UF 33145, 33146), Pip- ta Cruz [Puerto Santa Cruz], ... North and south ile cumanensis (UF 24048), Crypturellus bartlet- Vacas the Santa Cruzian [sic] has been eroded ti (LSU 109286; KU 84731), C. boucard (LSU back from the sea a distance of three miles ....” 23333), C. cinereus (KU 85127), C. cinnamomens This geographic reference is in agreement with (KU 41861; SDSNH 41693), C. obsoletus (LSU the locality given for Cañadón de las Vacas clari- 83940), C. parvirostris (LSU 125624), C. soui (KU fi ed by Marshall (1976). 34657), C. tataupa (LSU 125626), C. transfaciatus The Santa Cruz Formation is a rich fossilifer- (LSU 93849), C. undulatus (FMNH 320358; KU ous continental deposit, largely exposed in the 84742), Eudromia elegans (KU 78065), Nothocer- Argentinean Province of Santa Cruz (Ameghino, cus bonapartei (UMMZ 155485), N. julius (LSU 1889; Simpson, 1940; Marshall, 1976; Marshall 120893), N. nigrocapillus (LSU 99310), Notho- et al., 1986). Traditionally, the age of the Santa procta cinerascens (FMNH 314655), N. ornata Cruz Formation has been considered as early (UMMZ 209964), N. pentlandii (LSU 79742), Miocene (Marshall, 1976; Marshall et al., 1977; Nothura darwini (KU 77981), N. maculosa (KU 1983). However, recent radiometric dates sug- 77978; SDSNH 38630), Rhynchotus rufescens gest a younger age for these deposits (Marshall (FMNH 105649; KU 84358), Tinamotis ingoufi et al., 1986; Bown & Larriestra, 1990; MacFad- (SDSNH 38632), T. pentlandi (FMNH 105919), den, 1990). Marshall et al. (1986) considered the Tinamus guttatus (LSU 121484), T. solitarius age of the Santa Cruz Formation to be between (UMMZ 203318), T. tao (FMNH 330220). 18 and 15 Ma. New radiometric dates confi rm this younger age, with dates ranging from 16.6 Systematic Paleontology to 16.4 Ma for the lower part of the formation (Bown & Fleagle, 1993; Flynn & Swisher, 1995). Order Tinamiformes (Huxley, 1872) Family Tinamidae Gray, 1840 Methods aff. Crypturellus Brabourne & Chubb, 1914 Crypturellus reai, new species The monophyletic nature of the genera of the Tinamiformes, Tinamidae used in this study is Figure 2 based on the work of other Bertelli & Chiappe (2005). For purposes of identifi cation and look- Holotype – AMNH:FAM 9151, left humerus. ing for informative phylogenetic characteris- Type locality – Wash at Cañadón de las Va- tics, I have compared the fossil with represen- cas, southeastern Santa Cruz Province, Argen- tatives of all nonpasserine bird orders and in tina (fi gure 1). particular with the extinct paleognathous bird Type horizon – Santacrucian SALMA, early- Lithornis (L. plebius and L. promiscuus) and middle Miocene, 16.3-17.5 Ma, Santa Cruz Fm. with several species of cracids (Crax globulosa, (see Flynn & Swisher, 1995). Mitu mitu, Ortalis garrula, O. poliocephala, O. Measurements holotype – Total length: vetula, Penelope jacquagu; Pipile cumanensis). 50.2 mm; proximal breadth: 13.0 mm; depth The relationships between Lithornis, cracids, and width of shaft at mid-point: 8.5 x 9.8 mm; and the Tinamidae are controversial (Cracraft, distal breadth: 10.2 mm.

Figure 2. Crypturellus reai, AMNH:FAM 9151. A) Anconal; B) Palmar; C) Internal; D) External; E) Anconal; F) Palmar; G) Internal; H) External views. Abbreviations are: M. brachialis fossa (bra), bicipital crest (bic), capital incision (cap), M. infraspinatus attachment (inf), M. pectoralis attachment (pec), transverse ligamental groove (tlg), M. triceps brachii (tri), and ventral supracondylar tubercle (vst). Scale equals 1cm. Photography by the author. Drawing by Linda D. Chandler. Courtesy of AMNH: FAM.

Etymology – This new species is named in of the humerus extends distad only far enough honor of Amadeo M. Rea, a consummate educa- to occlude the capital incision medially (fi g- tor, mentor, and friend. Amadeo’s passion for ures 2A, E), unlike other species of Crypturellus birds, living and fossil, botany, and ethnobiol- where it extends farther distad and is broader. ogy fi rst sparked a desire in me to study fossil In addition, the attachment of the M. infraspi- birds. natus is concave (fi gures 2C, G), which differs Diagnosis – The holotype humerus (AMNH: from the condition of a fl at or slightly convex FAM 9151), Crypturellus, Tinamus, and Rhyn- attachment as is exhibited in the other species chotus have the distal border of the head of the of Crypturellus. Crypturellus reai also differs humerus extending distad, medially occluding from all other species examined in having the the capital incision; Nothocercus, Nothura, Eu- proximal end of the attachment for the ven- dromia, Nothoprocta, and Tinamotis have a dis- tral collateral ligament (ventral supracondylar tinctly rounded distal border with a medially tubercle) slanted more distally (fi gures 2D, H). open incision. The holotype, Crypturellus, and The attachment is more cranial in position in Tinamotis share the attachment for the M. in- the other genera. fraspinatus oriented ventrally, not caudally as In Crypturellus reai the transverse ligamen- in Tinamus, Rhynchotus, and the other tinamid tal groove is deep and extends ventrally almost genera examined. to the bicipital crest (fi gures 2B, C, F, G), a con- Within Crypturellus, C. reai differs from all dition approached only by C. bartletti and C. other species examined in having the head of transfaciatus. In Tinamotis and Nothocercus, the humerus more deeply undercut medially. the transverse ligamental groove forms a shal- Crypturellus reai also differs from all other spe- low pit. Both C. reai and C. tataupa have a dis- cies of Crypturellus examined in that the fossa tinct groove ascending proximad from the at- of the M. brachialis is nearly straight and ex- tachment of the M. pectoralis medially across tends diagonally across the shaft. Crypturellus the head of the humerus (fi gures 2A, E). reai has the attachment for the ventral collat- A nearly straight diagonal fossa of the eral ligament (ventral supracondylar tubercle) M. brachialis is present in C. reai (fi gures 2B, F) slanted more caudally, and not located in the and the other genera of tinamous. The other plane of the long axis of the humerus as in all species of Crypturellus, however, have a cresent- the examined species of Crypturellus and other shaped impression. tinamid genera. Comparisons with Crypturellus reai can be Description and comparison – Crypturellus made with humeri from extinct species from reai is a tinamou of medium size similar to the two other localities, Monte Observación and pale-browed, C. transfaciatus, and slaty-breast- Monte Léon. The two fossils (MACN-SC 1449 ed, C. boucardi, tinamous. The nearly complete and MACN-SC 3608) both are the distal ends left humerus is missing the lateral condyle and of right humeri. They virtually are identical to the dorsal epicondyle. It is from an adult indi- each other in size and appearance with the ex- vidual and is heavily mineralized. ception that one has an additional bony spur The new species has an affi nity with species above the dorsal epicondyle, which is a patho- in the genus Crypturellus based on two char- logical artifact. Both humeri have a shorter and acters: 1) the direct ventral orientation of the broader crescent-shaped fossa of the M. bra- M. infraspinatus attachment; and 2) the head chialis, whereas C. reai has a longer and nar- of the humerus extends distad occluding the rower impression. Also, the attachment for the capital incision medially (fi gure 2). Individually, ventral collateral ligament is fl at in cranial view, these characters are found in three other gen- which differs from the unique condition of the era (see Diagnosis), but not together in any ge- holotype. The late Pliocene extinct genus and nus other than Crypturellus. A caudally oriented species Tinamisornis parvulus does not have M. infraspinatus attachment and a non-obstruct- a humerus referred to it. Therefore, it cannot ed capital incision is present in Nothocercus. be directly compared to C. reai. (See Tambussi, Crypturellus reai has several unique features. 1987 for a review of this genus). The head of the humerus is more deeply under- Pleistocene records of modern species of tin- cut by the head of the M. triceps brachii than amous have been reported by Brodkorb (1963), in any other species of Crypturellus. The head Tambussi (1987; see table 1), and Cuello (1988).

Genus Miocene Pliocene Pleistocene Reference Tnamus x Brodkorb (1963) Brodkorb (1963); this Crypturellus xx paper Rhynchotus x Brodkorb (1963) Nothoprocta x Brodkorb (1963) Brodkorb (1963); Nothura xx Tambussi (1987) Tambussi (1987); Cuello Eudromia xxx (1988) Taoniscus x Brodkorb (1963) Tinamisornis x Tambussi (1987)

Table 1. Chronology of the fossil tinamid genera.

Discussion of Natural History, San Diego, California; Mu- seum of Zoology, University of Michigan, Ann Crypturellus reai is an early tinamou with an Arbor, Michigan. I received a Collection Study afﬁ nity to the genus Crypturellus. It is reported Grant from the AMNH to conduct this study. I from the early-middle Miocene, Santacrusian, thank Linda D. Chandler for skillfully prepar- continental deposits of Cañadón de las Vacas, ing the illustrations and for proof-reading the Santa Cruz Province, Argentina. In addition, manuscript. I especially would like to thank there is at least one more species of tinamou Luis Chiappe for his early participation, help- from the same deposit from two localities, Mon- ful comments, and use of tinamou fossils in his te Observacion and Monte Leon, which are in care. Also, I thank James L. Knight, Bradley C. close proximity. Also, there is an undescribed Livezey, Alfred J. Mead, and S. David Webb for tinamou from the slightly older Pinturas For- critically reviewing this paper. mation (Bertelli & Chiappe, 2005). Living congeners of Crypturellus reai are all Cited Literature found in tropical or subtropical environments (Blake, 1986). Previous analyses of the verte- Ameghino, F. 1889. Contribución al conocimien- brate fauna indicate tropical-subtropical climat- to de los mamíferos fósiles de la República ic conditions during the deposition of the Santa Argentina. – Actas Academia de las Ciencias Cruz Formation (Pascual, 1984; 1986; Pascual & de Córdoba 6: 1-1027. Ortiz Jaureguizar, 1990). The occurrence of C. Baumel, J.J., A.S. King, A.M. Lucas, J.E. Breazile & reai supports such an inference. H.E. Evans. 1979. Nomina Anatomica Avi- um. – New York, Academic Press. Acknowledgements Bertelli, S. & L.M. Chiappe. 2005. Earliest tinamous (Aves: Palaeognathae) from the Mio- I would like to thank the curators and staff of cene of Argentina and their phylogenetic po- the Department of Vertebrate Paleontology, sition. – Contributions in Science 502: 1-20. AMNH, for making the holotype and other fos- Bown, T. & J.G. Fleagle. 1993. Systematics, bio- sils available to me for study. Also, I thank the stratigraphy, and dental evolution of the curators of the following museums for mak- palaeothentidae, later Oligocene to early- ing modern comparative specimens or fossils middle Miocene (Deseadean-Santacrucian) available to me: Museo Argentino de Ciencias caenolestoid marsupials of South America. – Naturales, Buenos Aires, Argentina; Field Mu- Journal of Paleontology 67, 2 (Supplement): seum of Natural History, Chicago, Illinois; Flor- 1-76. ida Museum of Natural History, Gainesville, Bown, T. & C. Larriestra. 1990. Sedimentary Florida; Museum of Natural History, The Uni- paleoenvironments of fossil platyrrhine lo- versity of Kansas, Lawrence, Kansas; Museum calities, Miocene Pinturas Formation, Santa of Natural Science, Louisiana State University, Cruz Province, Argentina. – Journal of Hu- Baton Rouge, Louisiana; San Diego Museum man Evolution 19: 87-119.

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