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Bahia, Brazil Proc. Natl. Acad. Sci. USA Vol. 93, pp. 6405-6409, June 1996 Evolution A new extinct primate among the Pleistocene megafauna of Bahia, Brazil (New World monkeys/paleontology/Atelinae) CASTOR CARTELLE* AND WALTER CARL HARTWIGt *Instituto de Geociencias, Universidade Federal de Minas Gerais, Belo Horizonte, MG, Brazil; and tDepartment of Anthropology, George Washington University, Washington, DC 20052 Communicated by F. Clark Howell, University of California, Berkeley, CA, February 12 1996 (received for review November 1, 1995) ABSTRACT A nearly complete skeleton of a robust- Table 1. Mammals discovered in Toca da Boa Vista bodied New World monkey that resembles living spider mon- Order Species keys was recovered from undisturbed Pleistocene deposits in the Brazilian state of Bahia. The skeleton displays the highly Chiroptera Mormoops megalophylla specialized postcranial pattern typical of spider and woolly Pteronotus parnelli spider monkeys and shares cranial similarities to the spider Chrotopterus auritus monkey exclusively. It is generically distinct on the basis of its Loncophylla mordax robustness (>20 kg) and on the shape of its braincase. This Desmodus rotundus new genus indicates that New World monkeys nearly twice the +Desmodus draculae size of those living today were part of the mammalian biomass Eptesicus brasiliensis of southern Amazonia in the late Pleistocene. The discovery of Tadarida brasiliensis this specimen expands the known adaptive diversity of New Edentata + Nothrotherium maquinense World monkeys and demonstrates that they underwent body + Scelidodon cuvieri size expansion in the terminal Pleistocene, as did many other Myrmecophaga tridactyla types of mammals. Euphractus sexcistus Rodentia Coendou prehensilis Artiodactyla Tayassu tajacu Anthropoid primates colonized South America in the early Lama guanicoe part of the Tertiary (>30 million years ago) and radiated Mazama gouazoubira widely throughout the neotropics. Sixteen New World mon- Carnivora Cerdocyon thous key genera survive today, all of which are arboreal and + Protocyon troglodytes relatively small-bodied compared with most Old World +Arctotherium brasiliense anthropoids. Four living genera (Alouatta, Ateles, Brachy- Procyon cancrivorus teles, and Lagothrix) are closely related to one another and Conepatus semistriatus are distinct among New World monkeys for their relatively Felis pardalis large body size (6-12 kg), suspensory postures, and prehen- Felis tigrina sile tails (1-3). They are usually classified in a separate Felis yagouaroundi subfamily, Atelinae, within which the howler monkey (Al- Felis concolor ouatta) is considered less closely related to the other three + Smilodon populator than they are to each other (2). Ateles and Brachyteles are Primates + Protopithecus brasiliensis further distinguished by a specialized postcranial adaptation + Caipora bambuiorum to a brachiating mode of locomotion, in which the forelimbs are relatively elongated (3). The Pleistocene skeleton de- The plus sign indicates extinct species. scribed here resembles the spider monkey (Ateles) more than relative completeness of several fossil skeletons in this chamber it does any other living genus. However, this skeleton is suggests that the sediments were in primary depositional context unusually robust, and its neurocranium is more rounded at the time of discovery. compared with any living or extinct ateline. Systematics. The order is Primates (Linnaeus, 1758); the suborder is Haplorhini (Pocock, 1918); the infraorder is Platyrrhini (E. Geoffroy, 1812); the subfamily is Atelinae DEPOSITIONAL CONTEXT (Gray, 1825); the tribe is Atelini (Gray, 1825); the genus is The fossil skeleton was discovered in 1992 in Toca da Boa Caipora; and the species is Caipora bambuiorum. Vista, a cave in the Brazilian state of Bahia (Fig. 1). The Type Specimen. IGC-UFMG 05 (Instituto de Geociencias - skeleton was found in an undisturbed deposit in a chamber that Universidade Federal de Minas Gerais) is a nearly complete also contained other Pleistocene fossils and a nearly complete skeleton of a late-stage subadult, including cranium, mandible, skeleton of a second large platyrrhine referred to Protopithecus axial skeleton including caudal vertebrae, incomplete scapulae Lund, 1838 (4-5). Toca da Boa Vista is an extensive limestone and pelvis, upper and lower limb long bones, carpals, tarsals, travertine with over 100 km of interfingering galleries. The fossils metacarpals, metatarsals, and numerous phalanges. were recovered in a chamber along the northeastern wall of the Age and Locality. Toca da Boa Vista is located at 40o51'39R W cave several hundred meters from the principal entrance. The longitude and 10°09'36" S latitude at an altitude of 600 m above mean sea level. No radiometric dates are available from the The publication costs of this article were defrayed in part by page charge deposits, but other mammalian taxa found in the same sedimen- payment. This article must therefore be hereby marked "advertisement" in tary context represent a mix of living and extinct Pleistocene accordance with 18 U.S.C. §1734 solely to indicate this fact. species (Table 1). Pending a more precise radiometric assessment, 6405 Downloaded by guest on September 29, 2021 6406 Evolution: Cartelle and Hartwig Proc. Natl. Acad. Sci. USA 93 (1996) + 8° + 8° 420 38° State of Bahia FIG. 1. The location of Toca da Boa Vista in the northern part of the state of Bahia, Brazil, is shown. A V e 1cm 1cm FIG. 2. Anterior (A), lateral (B), and posterior (C) views of the cranium of IGC-UFMG 05, type specimen of Caipora bambuiorum. Table 2. Cranial measurements for Caipora and the four genera of living ateline New World monkeys Caipora (1) Ateles (92) Brachyteles (11) Lagothrix (73) Alouatta (25) NCL 94.1 77.9 86.6 73.7 61.2 68.5-84.4 79.8-91.7 67.0-81.2 54.9-68.8 NCB 75.4 60.8 61.9 58.6 51.0 54.9-65.7 57.7-65.1 - 53.8-63.1 47.3-56.2 TSL 131.5 114.1 114.8 104.9 107.6 104.0-122.0 100.0-122.0 97.2-114.6 96.3-121.4 BAS-NAS 77.2 63.3 68.2 63.1 64.9 55.4-71.9 58.2-74.4 57.9-70.0 57.6-78.2 PL 40.6 34.4 38.7 31.6 39.9 29.9-40.7 34.2-44.1 26.2-37.4 34.7-59.9 BOB 63.3 55.6 57.3 54.0 52.3 49.9-64.2 52.0-61.2 47.9-59.0 47.2-60.7 NCL, neurocranial length; NCB, neurocranial breadth; TSL, total skull length; BAS-NAS, basion-nasion; PL, palate length; BOB, biorbital breadth. Sample sizes are in parentheses. All measurements are in millimeters. Downloaded by guest on September 29, 2021 Evolution: Cartelle and Hartwig Proc. Natl. Acad. Sci. USA 93 (1996) 6407 4.60 4.4 0.5 'Z )J'I IV B _ * Aor 7 _~ 4.48 k A A teles, W 4.3 - 0.4 - yy BrachivteheS EI - L-agothliv. X c) 42k c. 4.36 F r Al99ittilit/ ZA I It 0.3 L TP: C E;4.1h --7' .4- 4.24 H ; . 4.0 C.) z 4.12 K Z 0.1 L 3.9 V Q 4.00; 3.8 4.20 4.32 4.44 4.56 4.68 4.80 4.20 4.32 4.44 4.56 4.68 4.80 4.20 4.32 4.44 4.56 4.68 4.80 Cranial Size Composite Cranial Size Composite Cranial Size Composite FIG. 3. Log-log plots of cranial dimensions on overall cranial size [(total skull length + basicranial length + facial length + basion-nasion)/4] for living atelines and Caipora. The ellipse represents 90% of the variation in the Ateles, Brachyteles and Lagothrix sample. Neurocranial length (A) and breadth (B) of Caipora have opposite relative relationships to the distribution of living species, such that a measure of their differential (C) demonstrates a more spherical neurocranium than is typical in living atelines. A B FIG. 4. Drawings of the palate (A) and mandible (B) of Caipora 1 cm bambuiorum, in occlusal view. Table 3. Postcranial measurements for Caipora and the four genera of living ateline New World monkeys Caipora (1) Ateles (31) Brachyteles (3) Lagothrix (17) Alouatta (25) FHD 22.9 17.9 18.2 15.0 13.4 15.8-20.2 16.9-19.8 14.0-15.7 11.8-15.9 FL 216* 205.6 202.0 166.4 154.2 190.5-226.0 186.5-212.0 157.5-176.5 139.0-171.0 BCB 38.5 31.8 29.0 27.1 23.9 29.1-34.9 28.0-30.9 24.2-29.3 21.4-26.7 HHD 25.1 20.5 19.8 20.1 19.8 17.8-24.1 18.2-21.6 18.6-22.2 16.9-23.1 HL 222.0 207.3 208.7 - 166.2 145.9 184.5-225.5 192.0-223.0 154.5-177.0 129.0-165.0 BIEPI 37.7 30.9 30.0 28.0 26.6 28.5-33.2 26.7-33.0 25.6-30.0 22.5-30.8 I-I 1.06* 1.05 1.07 0.98 0.95 1.01-1.07 1.05-1.08 0.96-1.00 0.92-0.98 FV 70 ml 40 ml 20 ml 20 ml FHD, femoral head diameter; FL, femoral length; BCB, femoral bicondylar breadth; HHD, humeral head diameter; HL, humeral length; BIEPI, humeral biepicondylar breadth; I-I, intermembral index (forelimb length/hindlimb length); FV, femoral volume (measured by water displacement in a graduated cylinder). Sample sizes are in parentheses. All measurements are in millimeters except where indicated. *Measurements that reflect incomplete growth of the fossil. Downloaded by guest on September 29, 2021 6408 Evolution: Cartelle and Hartwig Proc.
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