Inés Horovitz* and The quaternary Cuban platyrrhine Ross D. E. Paralouatta varonai and the origin of MacPhee† Antillean monkeys *Department of Vertebrate Paleontology, American We describe recently recovered dental and mandibular remains of the Museum of Natural History, Cuban platyrrhine Paralouatta varonai, previously known from the New York, NY 10024-5192, holotype only (a nearly complete skull with very worn teeth). We also U.S.A. E-mail: expand on the original description of the type specimen. [email protected] and Paralouatta is one of three extinct taxa of Greater Antillean †Department of Mammalogy, Quaternary monkeys known from craniodental remains. The other American Museum of Natural two, Xenothrix mcgregori and Antillothrix bernensis, occurred in Jamaica History, New York, NY and Hispaniola, respectively. It has been common practice to assume 10024-5192, U.S.A. that Antillean monkeys were more closely related to individual mainland taxa than to each other. Thus, P. varonai was thought to be Received 23 February 1998 related to Alouatta; Antillothrix bernensis to Saimiri or Cebus; and X. revision received 20 June mcgregori to Callicebus, or to callitrichines, or even to be of unknown 1998 affinity. With the discovery of well-preserved dental remains of and accepted 6 July 1998 Paralouatta, it can now be ascertained that this species was in fact very different from Alouatta. Cladistic analysis reveals a sister-group Keywords: New World relationship between Antillothrix and Paralouatta, followed on the monkeys, platyrrhines, cladogram by Xenothrix and Callicebus (last taxon being the closest Antilles, Quaternary, mainlaind relative of the Antillean clade). This conclusion has an Antillothrix, Xenothrix, important biogeographic implication: recognition of an Antillean Paralouatta, Caribbean. clade, as advocated here, assumes only one primate colonization from the South American mainland, not several as previously believed. 1999 Academic Press Journal of Human Evolution (1999) 36, 33–68 Article No. jhev.1998.0259 Available online at http://www.idealibrary.com on Introduction 1984; Fleagle & Bown, 1983; Fleagle et al., 1987; Fleagle, 1990). By comparison, the Platyrrhine primates have been a notable part diversity of the endemic platyrrhines of the of the mammalian fauna of South America Greater Antilles (hereafter, the Antillean since at least the late Oligocene/early monkeys) is known only in barest outline. Miocene (Hoffstetter, 1969; MacFadden, The objective of this paper is to review 1990). The living species are found from recent progress in unravelling and interpret- southern Mexico to northern Argentina, but ing the phylogeny of these monkeys, with the fossil record establishes the New World particular emphasis on the late Quaternary monkeys once ranged more widely, from species from Cuba, Paralouatta varonai. Patagonia to the Greater Antilles (i.e., the Our understanding of the fossil history of island group consisting of Cuba, Hispaniola, Antillean monkeys may be getting broader Puerto Rico, and Jamaica). The past diver- in a geographical sense, but it is not very sity of platyrrhines at the southern end of deep temporally. Cuba, Jamaica, and their distribution is becoming increasingly Hispaniola each supported one or more well known, thanks to more than a century endemic platyrrhines during the late Quater- of dedicated work in this region (Ameghino, nary, but Cuba is the only island with a 1906; Rusconi, 1934, 1935; Kraglievich, Tertiary primate record—currently consist- 1951; Hershkovitz, 1970, 1974, 1981, ing of a single fossil, a talus of early Miocene 0047–2484/99/010033+36$30.00/0 1999 Academic Press 34 . . age from the important locality of Domo de rel monkey (Rímoli, 1977) or some kind of Zaza (MacPhee & Iturralde-Vinent, 1995; cebine related to both Saimiri and Cebus Iturralde-Vinent & MacPhee, in press). (MacPhee & Woods, 1982). Similarly, it Fossils of extinct Antillean monkeys were seemed unlikely that Jamaican Xenothrix, found (although not properly recognized as variously considerd to be a callicebin, such) as early as the 1920s in Hispaniola and cebine, or a taxon of unknown affinities Jamaica (MacPhee, 1996), but Williams & (Rosenberger, 1977; MacPhee & Fleagle, Koopman (1952) were responsible for shed- 1991), could be closely related to the mon- ding the first real light on the subject. Since keys from Cuba and Hispaniola. Adding to then, a number of new discoveries have been this already complex picture, Ford (1986a) made (Rímoli, 1977; Rosenberger, 1977; and Ford & Morgan (1988) argued that MacPhee & Woods, 1982; Ford & Morgan, several of the previously mentioned isolated 1988; Ford, 1990; Rivero & Arredondo, postcranials from Hispaniola and Jamaica 1991; MacPhee et al., 1995; MacPhee, are specifically marmoset like, which implied 1996), but the Antillean record is still that yet another major platyrrhine clade was clouded by a variety of uncertainties. From once represented in the Greater Antilles. a systematic standpoint, the most serious It is self-evident that, if the primate com- of these concerns how many species are plements of Cuba, Jamaica, and Hispaniola actually represented in existing fossil assem- had multiple origins, there had to have been blages, and how they are related to one multiple colonization events. However, another. There are three named taxa: thanks to a number of new discoveries, Paralouatta varonai Rivero & Arredondo, especially ones made in the present decade, 1991, from Cuba; Xenothrix mcgregori this picture of multiple separate coloniza- Williams & Koopman, 1952, from Jamaica; tions by unrelated propagules is no longer and Antillothrix bernensis MacPhee et al., so easily maintained. The multiple origins 1995 (formerly Saimiri bernensis Rímoli, viewpoint was explicitly challenged by 1977), from Hispaniola. There are, in MacPhee et al. (1995), who demonstrated addition, a number of primate-like post- on the basis of a preliminary cladistic study cranial remains from these islands that, for that Paralouatta and Antillothrix are related one reason or another, have not yet been as sister groups, and that Callicebus may be allocated; some may represent unknown the closest living relative of this dyad. The taxa (MacPhee & Fleagle, 1991; MacPhee, position of Xenothrix was not investigated. 1996). It is a safe bet that we are far However, it is of interest that Rosenberger from having a complete roster of Antillean (1977) independently posited that Xenothrix monkeys, even for the Quaternary (Ford, might be related to Callicebus on the basis 1990). of evidence then available. New fossils of The relationships of Antillean monkeys Xenothrix (description of which is in prep- are obscure. Indeed, until very recently, the aration) provide an opportunity to determine only conclusion one could draw from the whether all Antillean monkeys (or at least scanty literature on this subject was that those represented by adequate material) may these monkeys had little to do with one derive from a single colonizing ancestor. another. Thus, it might be concluded that One objective of this paper is to put on Cuban Paralouatta, supposedly a close rela- record a description and evaluation of all tive of Alouatta (Rivero & Arredondo, craniodental specimens of P. varonai so far 1991), could not also be a near affine of recovered. Another is to present a com- Hispaniolan Antillothrix bernensis because prehensive cladistic analysis of Antillean the latter was viewed as either a giant squir- monkeys. We address the latter goal within PARALOUATTA VARONAI 35 Figure 1. Map of Cuba showing principal localities: CA, Cueva Alta and Cueva del Mono Fósil (Quaternary); DZ, Domo de Zara (early Miocene). the larger issue of platyrrhine systematics. ated on the same south-facing slope of the To avoid ambiguity, in this paper we explic- Sierra de Galeras, one of several blocks of itly define pitheciins as including Pithecia, uplifted Jurassic limestones that make up the Chiropotes, and Cacajao, atelines as Ateles, Cordillera de Guaniguanico (Figure 1). Brachyteles, Lagothrix, and Alouatta, and cal- The first primate fossils from Cueva del litrichines as Callimico, Callithrix, Cebuella, Mono Fósil were found in 1988 and Leontopithecus, and Saguinus. included the type skull (Figure 2) and a distal humerus. Remains belonging to other Abbreviations vertebrates indicated that the temporal con- text of the site was Quaternary. The first AMNH, American Museum of Natural detailed account of this material, by Rivero History, New York & Arredondo (1991), included a formal CMC, Claremont-McKenna College, description of P. varonai and emphasized California the similarity of the type skull to that of the GPBSEC, Grupo Espeleológico ‘‘Pedro living howler monkey, Alouatta. Borrás’’ of the Sociedad Since the late 1980s, accessible portions Espeleológica de Cuba of Cueva Alta and Cueva Mono Fósil have MNHNH V, Museo Nacional de Historia been carefully combed for new fossils by Natural, La Habana, expeditions of the GPBSEC, MNHNH, and Vertebrate Collection AMNH. A number of additional platyrrhine M-D, mesiodistal remains have been recovered as a result, B-L, buccolingual including a mandible, numerous isolated teeth, and several postcranial pieces. Craniodental morphology and Although exploration continues in the relationships of Paralouatta varonai numerous caves that riddle the rest of the Fossils allocated to P. varonai have been south face of Sierra de Galeras, to date no recovered at only two localities in Cuba, new primate-bearing