New Evidence of Ara Autochthones from an Archeological Site in Puerto Rico: a Valid Species of West Indian Macaw of Unknown Geographical Origin (Aves: Psittacidae)
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3?? Caribbetiti iounml of Science. Vol. 44, No. 2, 215-222, 2008 Copyright 2008 College of Arts and Sciaices University of Puerto Rico, Mayagüez New evidence of Ara autochthones from an archeological site in Puerto Rico: a valid species of West Indian macaw of unknown geographical origin (Aves: Psittacidae) STORES L. OLSON^'' AND EDGAR J. MAíZ LóPEZ,^ 'Department of Vertebrate Zoologu, National Museum of Natural History, Smithsonian Institution, P. O. Box 37012, Washington, D.C. 20013-7012, U.S.A. ^Centro de Estudios Avanzados de Puerto Rico y el Caribe, San Juan, Puerto Rico (USA), 00902-3970. *Corresponding author: [email protected]. ABSTRACT.•The exinct macaw Ara autochthones, previously known only from a single bone from an archaeological site on St. Croix, Virgin Islands, is here identified from several associated bones from an archaeological site in south-central Puerto Rico. The species belongs to a distinctive intermediate size-class and was larger than the Cuban Macaw Ara tricolor. It is assumed to have been endemic to the West Indies, but prehistoric interisland transport of parrots by humans makes interpreting the natural distribution of the species impossible in the absence of fossils. Historical reports of macaws elsewhere in the West Indies are rendered dubious for the same reason. KEYWORDS.•Amazona, biogeography, extinction, human transport, parrots. INTRODUCTION described and named a new species of ma- caw as Ara autochthones, based on a single The history and natural distribution of tibiotarsus of an immature bird. Nothing macaws {Ara) in the West Indies are further regarding this species has turned clouded with uncertainties. The only speci- up in the 60 years since it was described men evidence apart from, archeological re- and there has been no further evaluation of mains is of the Cuban Macaw Ara tricolor, the species, which has been mentioned oc- known from about nineteen skins and ex- casionally in various checklists and compi- tinct since about 1864 (Greenway 1958, Ol- lations; e.g. the curious statement by son and Suárez, in press). Contrary to pre- Prestwich (1970: 199) that: "Nothing ap- vious belief, there is no historical evidence pears to have been recorded concerning for a macaw from Hispaniola (Olson 2005), this rather primitive macaw." the second largest of Üie AntUlean islands. flere we report on several associated From Jamaica, Guadeloupe, Martinique, skeletal elements from another archeologi- Dominica, and an unknown West Indian cal site in central Puerto Rico that we refer island, there are 18^*^ and 19*^^ century visi- to Ara autochthones. These confirm the va- tors' accounts of various macaws to which Kdity of the species and provide proof of no fewer than seven scientific names have the existence of a second species of macaw been applied {Clark 1905a,b; 1908; Roths- endemic to the West Indies. Information child 1905, 1907a,b), although these are all that we supplied concerning this material rightly to be considered entirely hypotheti- formed the basis for Wiley et al. (2004:96) cal (Prestwich 1970), In addition, Fisher and reporting "Ara unknown sp." from Puerto Warr (2003) discovered and reproduced a Rico. Unfortunately, as the with archeologi- previously unknown painting of a macaw cal specimens of parrots from elsewhere in supposedly from Jamaica dating from the West Indies reported by Williams and about 1765. Steadman (2001), and in the absence of a From an archeological deposit on St. fossil record, it is not possible to determine Croix in the Virgin Islands, Wetmore (1937) on which island this species of macaw 215 216 S. L. OLSON AND E. J. MAÍZ LÓPEZ originally evolved because of the potential rower shaft, ventral lip of glenoid facet for extensive trade in parrots among Am- more protrudent; humérus with ectepicon- erindians of the Antilles. dylar process and attachment of pronator brevis situated decidedly more proximad; MATERIALS AND METHODS carpometacarpus proportionately much longer, process of alular metacarpal not Comparative material examined.•Skel- curved proximad; femur with head propor- etons: Anodorhynchus hyacinthinus MHNT tionately larger; tibiotarsus very distinctive 1045, 1057, 1064, 1496, 1693, 1695, USNM in having the inner cnemial crest more 291249, 319969, 345230, 345854; A. leari pointed and extending farther proximad, FMNH 337716, 337860, 379161, MHNT internal condyle much narrower. Although 1540, 1547; Ara ambiguus LSUM2 90381; we have followed David and Gosselin USNM 224811; A. ararauna MHNT 242,983, (2002) in treating the generic name Ara as 1165, 1604, USNM 19355, 49891, 223952, masculine, we do not endorse splitting the 223993, 318791, 322286, 322337, 345207, genus into three by resurrecting the names 345848, 345849, 428243, 489411, 498698, PrimoUus and Orthopsittaca (Tavares et al. 502499, 502500; A. auricolUs USNM 345846, 2006). Recognition of monophyly of the 345847, 345851, 345852; A. chloropterus true macaws would be better served by in- MHNT 825, 1653, USNM 225132, 226876, cluding Cyanopsitta in Ara as it has long 345850,490125; A. couloni FMNH 291744; A. been delimited. glaucogularis FMNH 337727, LSUMZ 168622; A. rmcao MHNT 753, USNM 18508, Ara autochthones Wetmore, 1937 18988, 226164, 288772, 290508, 321173, 321981, 322058, 322212, 430513, 430516, Holotype.•USNM 483530, left tibiotar- 431614, 502497,502498; A. manilatus USNM sus; vertebrate paleontological collections 344700, 345853, 621711, 621949, 622388; A. (formerly USNM 343033 in the bird collec- maracaná FMNH 337756, 390830, 398918, tions). Collected in kitchen midden depos- USNM 320003, 344670; A. militaris USNM its from Concordia, southwestern St. Crobc, 288554, 288605, 344772, 344848; A. nobilis Virgin Islands, in 1934 by L. J. Kom (Wet- USNM 344080, 344081, 502284, 502503, more 1937). 622355; A. ruhrogenys FMNH 291402, Referred material.•USNM 448344 verte- 291404, 337744, MHNT 1812; A. severus brate paleontological collections: left cora- FMNH 104484, 290489, 337748, MHNT 388, coid lacking a portion of the head, proximal USNM 19115, 502504; Cyanopsitta spixii and distal ends of left humérus, proximal MHNT 820, USNM 346722. Measurements end of right radius, left carpometacarpus were also taken from X-radiographs of two lacking minor metacarpal, left femur lack- mounted specimens of Ara tricolor USNM ing distal end, right tibiotarsus lacking ex- 135137, 171767, Qualitative comparisons ternal part of proximal articular surface, were made with skeletons of Ara glaucogu- proximal fragment and worn distal portion laris i"Ara caninde" auct. FMNH 337727), of left tibiotarsus, fragment of shaft (hu- Anodorhynchus leari (FMNH 337716), and mérus?), unidentified fragm^ent (perhaps Amazona imperialis (USNM 318792, USNM not avian). These bones are evidently aä 321883). from a single individual. Locality and age.•Collected by Maiz dur- ing an excavation conducted in March and RESULTS April 1987 at the Hernández Colón (PO-13) Genus Ara Lacépède, 1799 archaeological site. The site, UTM E 755665/N1998980, represents an inland The new archaeological material is refer- Saladoid/Ostionoid pre-Columbian Indian able to Ara, rather than Amazona, the only village of approximately 15,000 m^. It is lo- other genus of large parrots in the West cated on the eastern bank of the CerriUos- Indies, by the following characters: cora- Bucaná River, south central Puerto Rico, coid more elongate with relatively nar- NE of the city of Ponce, Barrio Cerrillos (18° ARA AUTOCHTHONES IN PUERTO RICO 217 04' 05" N; 66° 35' 09 W). It lies at 76m amsl, 5.0. Carpometacarpus: length 55.8, proxi- 13,5 river km from the Caribbean Sea. mal depth 14.3, width of trochlea 5.7, width Physiographically, the Hernández Colón and depth of shaft at midpoint 4,5 x 5.0. site is situated in an alluvial terrace within Radius: greatest proximal diameter 6.0. Fe- the Semiarid Southern Foothills of Puerto mur: estimated length 51.5, proximal width Rico. 12.5, depth through trochanter 8.2, depth of Ten 2 X 1 m stratigraphie pits were ex- head 6.2, width and depth of shaft at mid- cavated after mapping the site. The pottery point 4.9 X 5.4. Tibiotarsus: length from seriation and two radiocarbon dates re- proximal articulating surface 74.5, length vealed a multi-component site, with a local from distal end of fibuiar crest to external sequence of three archaeological phases: condyle 47.2, depth through inner cnemial Pomarrosa Phase, Cerrillos Phase, and Ma- crest 11,4, width and depth of shaft at mid- ragiiez Phase. The Pomarrosa phase is sty- point 5.2 X 4.0, distal width 10.1. listically related to the Hacienda Grande ce- Comparisons.•The referred tibiotarsus is ramic style {ca. 200 B.C.-400 A.D.) as essentially identical in size with the holo- defined for Puerto Rico by Alegría (1965) type of Ara autochthones and the referred and Rouse & Alegría (1990). The Cerrillos material from Puerto Rico is therefore iden- and Maragüez phases are in turn related to tified as that species. As Wetmore (1937) the Cuevas (400-600 A.D.) and Early Osti- noted, the holotype is from a juvenile indi- ones (600-900 A.D.) styles as defined by vidual, so the new material is all the more Rouse (1952, 1992). The Hacienda Grande important for establishing the nature of the style is included within the Cedrosan Sala- species. In size, most living species of ma- doid subseries of the Saladoid series and caws fall into two separate clusters repre- corresponds with the first horticultural and senting large species and smaller species ceramics groups that migrated to Puerto (Table 1). Ara autochthones is distinct in be- Rico from northeastern South America ing intermediate between these two clus- {Rouse, 1992). All cultural and faunal re- ters. Only Ara glaucogularis and Anodorhyn- mains were collected using three gauges of chus leari (and presumably the very closely screens: 6 mm (1/4 inch), 3 mm (1/8 inch) related A. glaucus, which may be only sub- and 1.5 mm (1/16 inch).