american museum

AMERICAN MUSEUMNovitatesPUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORYCENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024

Number 3141, 31 pp., 11 figures, 3 tables June 30, 1995

Orgin of the Greater Antillean Land MammalFauna, 1: New Tertiary Fossils from

Cuba and Puerto Rico

R. D. E. MAcPHEE' AND MANUEL A. ITURRALDE-VINENT2

ABSTRACT

A new isolobodontine capromyid rodent (Za-zamys veronicae, new genus and species) and a newplatyrrhine primate (unnamed genus and speciesA) are described from material recovered at Domode Zaza, an important late Early Miocene localityin south-central Cuba near the town ofSancti Spir-itus. Also described is a femur believed to repre-sent a small megalonychid sloth (unnamed genusand species B), from a well-dated Early Oligocenesite near the town ofYauco in southwestern PuertoRico.These and other discoveries in the Greater An-

tilles establish that these islands supported a di-verse land mammal fauna from the Oligocene on-ward. Pertinent geological evidence is consistentwith the proposition that the initiators of all orpractically all founder clades of Antillean landmammals were emplaced there near the Eocene-

Oligocene boundary. Under the terms ofthis mod-el it is not expected that any land mammal faunalelements of greater than Late Eocene age will befound in these islands. It is reasonable to infer thatTertiary Antillean mammal groups were widelydistributed throughout this terrain, and the dis-covery of a biogeographical "ghost" (Zazamys)supports this.Emergent parts of the present-day Greater An-

tilles were in a close-packed array until the com-mencement of the Neogene. They formed part ofa long-lasting positive topographic feature of theCaribbean Basin, here dubbed GAARlandia. Dur-ing the Late Eocene-Oligocene, this feature in-cluded Cuba, northern Hispaniola, Puerto Rico,Virgin Islands, Aves Rise, and the Venezuelan An-tillas Menores. Since latest Oligocene, GAARian-dia has undergone disruption and fragmentation

' Chairman and Curator, Department of Mammalogy, American Museum of Natural History.2 Curator, Geology and Paleontology Group, Museo Nacional de Historia Natural, Capitolio Nacional, CH- 10200,

La Habana, Cuba.

Copyright © American Museum of Natural History 1995 ISSN 0003-0082 / Price $4.90

AMERICAN MUSEUM NOVITATES

into its modem insular components, a process thatforced vicariance ofthe original fauna. Extinctionhas also shaped the distribution of Antillean land

mammals throughout the later Cenozoic, and thishas important implications for interpreting theirhistorical biogeography.

RESUMEN

Un nuevo roedor capromydo isolobodontido(Zazamys veronicae, nuevo genero y especie) y unnuevo primate platyrrhinido (genero y especie A,sin denominar) se describen a partir del materialrecobrado en Domo de Zaza, una importante lo-calidad fosilifera del Mioceno Inferior tardio si-tuada en la region sur-central de Cuba, cercana ala ciudad de Sancti Spiritus. Tambien se describeun femur que se cree representa un pequeiio pe-rezoso megalonychido (genero y especie B, sin de-nominar), de un sitio paleontologico bien datadode edad Oligoceno Inferior, localizado cerca de laciudad de Yauco, en el suroeste de Puerto Rico.

Estos descubrimientos, unido a otros recien rea-lizados en las Antillas Mayores, permiten estable-cer que estas islas ya soportaban una variada faunade mamiferos terrestres a partir del Oligocene. Lasevidencias geologicas son consistentes con la pro-posicion de que los iniciadores de todos, o prac-ticamente de todos los clados fundadores de lafauna de mamiferos terrestres antillanos, se em-plazaron en estas tierras cerca del limite Eoceno-Oligoceno. Bajo los terminos de este modelo, esde esperar que ningun mamifero terrestre masantiguo que Eoceno Superior sea encontrado en

estas islas. Es razonable, asimismo, inferir que losgrupos de mamiferos terrestres estaban amplia-mente distribuidos en las tierras antillanas, y eldescubrimiento de un "fantasma" biogeografico(Zazamys) en el Mioceno Inferior de Cuba, sus-tenta esta afirmacion.

Las partes emergidas de las actuales AntillasMayores formaban entonces un conjunto mascompacto hasta el comienzo del Neogeno. Ellasformaban parte de un elemento topografico po-sitivo muy duradero en la cuenca del Caribe, elcual aqui se denomina GAARlandia. Durante elEoceno Superior-Oligoceno, este elemento incluiaa Cuba, La Espaniola septentrional, Puerto Rico,Islas Virgenes, Cresta de Aves y las Antillas Me-nores Venezolanas. A partir del Oligoceno tardio,comenzo la fracturacion y dispersion de GAAR-landia hasta formar sus componentes insulares ac-tuales, un proceso que forz6o la vicariancia de lafauna original. Las extinciones que han ocurridoa traves del Cenozoico tardio, tambien es probableque hayan contribuido a conformar la distribucionactual de los mamiferos terrestres Antillanos, yestos tiene importantes implicaciones para la in-terpretacion de su biogeografia historica.

INTRODUCTION

How and when the ancestors of the landvertebrate fauna ofthe Greater Antilles man-aged to reach those islands are topics per-ennially debated in biogeography (for recentexamples, see Hedges et al. [1992, 1994] andPage and Lydeard [1994]). Although agree-ment concerning the method(s) of coloniza-tion is evidently not in sight, there is a grow-ing fossil tally (table 1) which provides un-deniable evidence that founder clades werealready well represented in the islands by themid-Cenozoic. As noted elsewhere (MacPheeand Wyss, 1990), fossils are empirical doc-uments with temporal as well as phylogeneticimplications, and as such are critically im-portant to the reconstruction ofbiogeograph-ical history.

In this report we announce the discoveryof the second and third land mammals ofTertiary age from Cuba, and bring attentionto the first such discovery made in Puerto

Rico. We also extend and modify our modelof the land-mammal colonization of theGreater Antilles (MacPhee and Iturralde-Vi-nent, 1994). In the text, the age of stage andepoch boundaries cited in millions of years(Ma) follows the framework ofHarland et al.(1990).

ABBREVIATIONS

AMNH American Museum of Natural His-tory

AMNH M American Museum of Natural His-tory, Department of Mammalogy

AMNH VP American Museum of Natural His-tory, Department of Vertebrate Pa-leontology

e estimate (in measurement, due tobreakage)

Fm formationJD Juana Diaz Fm (sensu Frost et al.,

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MAcPHEE AND ITURRALDE-VINENT: TERTIARY MAMMALS

1983), not including "upper carbon-ate member"

Ma millions of years (ago)MNHNH Museo Nacional de Historia Natural

(La Habana)MNHNH P Museo Nacional de Historia Natural

(La Habana), paleontological collec-tions of Departamento de Colec-ciones

MNHNH V Museo Nacional de Historia Natural(La Habana), vertebrate collectionsof Departamento de Colecciones

rev. reversed (as applied to figures, to fa-cilitate comparison)

UC "upper carbonate member," in olderdefinitions of Juana Diaz Fm

ACKNOWLEDGMENTSFieldwork at Domo de Zaza was conducted

under the auspices ofagreements between theMuseo Nacional de Historia Natural, theformer Academia de Ciencias de Cuba (pres-ent Ministerio de Ciencias, Tecnologia y Me-dio Ambiente) and the American Museum ofNatural History, and was financed by grantsfrom the National Science Foundation (NSFBSR 900002) and the AMNH. Fieldwork atYauco was conducted with the assistance ofthe Departamento de Geologia, Universidadde Puerto Rico (Recinto de Mayagiiez) andVictor Gonzalez ofSan Juan, and was fundedby a grant from the Caribbean Scientific Ini-tiative ofthe RARE Center for Tropical Con-servation. For assistance at Domo de 7aza,we extend special thanks to Stephen DiazFranco, Osvaldo Jimenez Vasquez, LuisOlmo, Reynaldo Rojas Consuegra, and PavelValdes Ruiz. In connection with the PuertoRican expedition, we thank Profs. JamesJoyce and Hernan Santos (Universidad dePuerto Rico, Mayagiiez) for information con-cerning the Yauco site, and VeronicaMacPhee and Clare Flemming for assistancein the field. We also thank Veronica and Clarefor assistance in editing the manuscript ver-sion of this paper. RDEM took the photo-graphs used in figures 9 and 11; MIV drewfigures 8 and 10; all other illustrations are thework of our talented artists Lorraine Meeker(figs. 1, 2, 3 and 5) and Clare Flemming (figs.4, 6, and 7). Dr. Hans Schellekens (UPR Ma-yagiiez) and Dr. Jennifer White (SimpsonCollege) read and commented upon an earlierdraft of this paper. Finally, we are extremely

grateful to Prof. Edward Robinson (Depart-ment of Geology, University of the West In-dies) and Lic. Gena Fernandez (CIDP-Cu-pet, Cuba) for identifying foraminifers in ourYauco samples (see text).

EARLY MIOCENE RODENT ANDPRIMATE FROM CUBA

A new capromyid rodent, referable to sub-family Isolobodontinae, is named here on thebasis of dental material from the importantEarly Miocene locality of Domo de Zaza inthe province of Sancti Spiritus. A platyrrhinemonkey is also represented in the Zaza ma-terial by a single, well-preserved astragalus.Although the astragalus appears to representa species new to science, it makes a poorholotype. Accordingly, until specimens ca-pable ofproviding a secure diagnosis are dis-covered, we shall simply refer to the new Zazaanthropoid as "unnamed genus and speciesA (primate)." Locality information is keptpurposely brief, since this site and its contextis fully described in another paper (MacPheeet al., in prep.).

RODENTIA

CAPROMYIDAE: ISOLOBODONTINAE

Zazamys veronicae, new genus and speciesHOLOTYPE: Left mandibular ?M3

(MNHNH P 3072; fig. IA, F-G), found inlag deposit near south end ofDomo de Zazasection by 0. Jimenez in February 1994.ETYMoLoGY: Zaza (locality name) + ,uva

(mouse). Species name for Veronica Mahan-ger MacPhee, in gratitude for her unflagginghelp and sharp editing pencil.TYPE LocALiTY AND AGE: Domo de Zaza,

a locality in late Early Miocene Lagunitas Fmof south-central Cuba. For detailed localityinformation and age assessment, see Mac-Phee and Iturralde-Vinent (1994) andMacPhee et al. (in prep.).REFERRED MATERIAL: Right mandibular

Ml or M2 (MNHNH P 3058, fig. 1B, rev.)and left mandibular Ml or M2 (MNHNH P3071, fig. 1 C), found in lag by (respectively)P. Valdes in November 1993 and R. MacPheein February 1994.

DiAGNosIs: The type and only species ofZazamys is Z. veronicae-, genus and species

31995


TABLE 1Reported Tertiary Freshwater and Terrestrial Vertebrates of the Greater Antillesa

Island Age/Formation/Locality Reference

CubabImagocnus zazae (Megalonychidae,

Xenarthra)Zazamys veronicae (Capromyidae,

Rodentia)Unnamed genus and species A

(Platyrrhini, Primates)Unidentified bird

Unidentified pelomedusid turtle

Hispaniolac.deCichlasoma woodringi (Cichlidae,

Perciformes)Anolis dominicanus (Iguanidae,

Sauria)Sphaerodactylus sp. (Gekkonidae,

Sauria)Eleutherodactylus sp. (Leptodactyli-

dae, Anura)cf. Nesoctites (Picidae, Aves)

Unidentified mammal (?rodent)

Puerto RicoBoid (Ophidia)?Iguanid (Sauria)Unnamed genus and species B

(Tardigrada)Unidentified pelomedusid turtle

E. Miocene; Lagunitas Fm; Domo deZaza, S. Spinitus

E. Miocene; Lagunitas Fm; Domo deZaza, S. Spiritus




?M. Miocene; Las Cahobas Fm; Mire-balais, Haiti

?Miocene; amber deposits; DominicanRepublic

?Oligo-Miocene; amber deposits; Do-minican Republic

?Late Eocene; amber deposits; Domin-ican Republic

?U. Eocene-E. Miocene; amber depos-its; Dominican Republic

?Miocene; amber deposits; DominicanRepublic

E. Miocene; Cibao Fm; AguadillaE. Miocene; Cibao Fm; AguadillaE. Oligocene; Juana Diaz Fm; Yauco

E. Miocene; Cibao Fm; Aguadilla

MacPhee & Iturralde-Vinent,1994

This paper

This paper

MacPhee et al., in prep.

MacPhee et al., in prep.

Cockerell, 1924

Rieppel, 1980

Kluge, 1995

Poinar & Cannatella, 1987

Laybourne et al., 1994

Poinar, 1988

MacPhee & Wyss, 1990MacPhee & Wyss, 1990This paper

MacPhee & Wyss, 1990a This table updates table 2 of MacPhee and Iturralde-Vinent (1994), which should be consulted for additional

remarks.b Intact carapaces and plastrons of pelomedusid turtles have been recovered in situ from lagoonal deposits in

Lagunitas Fm. This strongly implies that they were freshwater-adapted (cf. MacPhee and Iturralde-Vinent, 1994,table 2); the habitat of Puerto Rico pelomedusids is still ambiguous, but we include them in this paper.

c See Grimaldi (in press) for discussion of the general problem of dating some ambers; listed ages follow authors'reports, with question marks expressing level of doubt.d Cichlasoma woodringi may not be distinguishable from extant species (M. L. Smith, personal commun.).e "Sphaerodactylus" dommeli (B6hme, 1984) may be an anole rather than a sphaerodactyl according to Kluge

(1995, citing D. Frost, personal commun.).

diagnoses are therefore the same. Amongknown Antillean taxa, Zazamys veronicaemost closely resembles the Quaternary An-tillean genus Isolobodon, representatives ofwhich are known from Hispaniola, PuertoRico, Mona Island, and several ofthe VirginIslands, but not from Cuba (Anthony, 1926;Reynolds et al., 1953; Varona, 1974; Woods,

1989b). Whether this multi-island range isoriginal or the result ofhuman introductionshas long been debated (cf. Allen, 1942; Mor-gan and Woods, 1986), but for the purposesof this paper we accept the occurrence of Is-olobodon on both Hispaniola and Puerto Ricoas natural. Also, in agreement with Woods(1989b) we accept Isolobodon portoricensis

4 NO. 3141

MACPHEE AND ITURRALDE-VINENT: TERTIARY MAMMALS

and L montanus as good species, but followReynolds et al. (1953) in considering L leviras synonymous with I. portoricensis.

Occlusal pattern, root shape, and arc ofcurvature of the teeth in the Zazamys hy-podigm indicate that all are lower molars.Lower molars of Zazamys have two lingualreentrant folds (mesoflexid and metaflexid)and one buccal fold (hypoflexid), as is gen-erally characteristic of capromyids (Woods,1989b). Zazamys differs from all capro-myines and hexolobodontines sensu Woods(1989b, 1992) in that the thick continuousshell ofcementum encircling the molariformsof taxa in these two subfamilies is absent.Zazamys resembles the plagiodontines Pla-giodontia and Rhizoplagiodontia in that ce-mentum is limited to reentrant folds, but dif-fers in that the folds themselves are notobliquely oriented (see below). Zazamys re-sembles the isolobodontines Isolobodon por-toricensis and I. montanus in the patterningof occlusal surfaces. Howevei, there are dif-ferences which indicate that separation at thegeneric level is warranted. The lower molarsof Zazamys differ from those of L portori-censis in the following respects: (1) Reenter-ing metaflexid and hypoflexid meet at thecenter of the occlusal surface, enclosingroughly equal areas of dentine between theirarms; in Isolobodon, metaflexid-hypoflexidjuncture is lingually displaced, so that me-taflexid arms are considerably shorter thanhypoflexid arms and area of dentine enclo-sure is considerably smaller. (2) Ridged ce-mentum is essentially restricted to reentrantfolds and is barely discernible on tooth col-umns (see fig. 1); in Isolobodon, ridged ce-mentum is easily distinguishable outside ofreentrant folds. (3) Reentrants are much more

compressed (also more compressed than inL montanus, which is intermediate in thisregard).

DEsCRnPrION: Because these teeth are iso-lated finds, their orientation in thejaw duringlife is not certain. However, judging from thelocation of intraproximal wear facets andother features, they were probably seated insuch a way that the reentrants were more orless normal to the axis of the jaw.The teeth arejudged to be hypselodont (ev-

ergrowing, with open roots) and probablyhypsodont, although ontogenetic data and

complete jaws would be needed to confirmthese points. Cementum forms noticeableridges only within reentrant folds (fig. 1 F, F'),but in MNHNH P 3072 there are barely dis-cernible, quasiparallel lineaments detectableon external surfaces (fig. lF, arrows). As faras may be determined without a histologicalexamination, these lineaments are not sepa-rately composed of cementum but seem tolie within the enamel. It is unclear whetherthis condition in Zazamys can be validly re-garded as antecedent to the highly pro-nounced version of cementum ridging seenin Isolobodon (fig. 1E). In capromyines, ce-mentum within reentrant folds has a muchless organized, frothy texture.

Because cementum layers on capromyidteeth are applied to smooth enamel surfaces,they can and do scale off. However, small"islands" of cementum usually remain, ex-cept in specimens that have been severelyabraded. The teeth from Zaza are not signif-icantly abraded and we therefore concludethat complete sleeves ofcementum were nev-er present.PHYLOGENETIC RELATIONSHIPS: Isolobo-

dontines and plagiodontines are closely re-lated and are often included in the same sub-family (e.g., by Varona, 1974). Woods (1 989a)defined 14 character state differences betweenIsolobodon and Plagiodontia, but ofthese onlyone (ridged cementum on external surfacesof tooth) can be evaluated on the basis oftheexisting Zazamys hypodigm. Within Ca-viomorpha, loss or modification of externalcementum on molars is not distributed in away that is consistent with the likeliest phy-logenetic reconstructions of the group; mul-tiple convergences across the suborder havetherefore occurred. Ridging, however, ap-pears to be a unique synapomorphy of iso-lobodontine capromyids, occurring (as far asis now known) in no other caviomorph clade.Possession of a version of this character de-cisively places Zazamys next to known iso-lobodontines, a fact which can be suitablyunderlined by including the Zaza rodent insubfamily Isolobodontinae.

It is widely agreed that capromyids andechimyids are closely related, although thereis rather less agreement concerning within-group relationships and the closeness oftheiraffiliation with other caviomorphs (cf. Corbet

51995


A1

Fig. 1. Holotype ofZazamys veronicae (MNHNH P 3072), a left mandibular M3, is seen in occlusal(A), lingual (F), lingual detail (F'), and buccal (G) views, from Early Miocene locality ofDomo de Zazain south-central Cuba. Referred specimens seen in occlusal view are (B), a right (rev.) lower Ml or M2(MNHNH P 3058); and (C), a left lower Ml or M2 (MNHNH P 3071). Compare to left mandibulardentition of Isolobodon portoricensis (AMNH M 38409), in occlusal (D) and buccal (E) aspects (anterioris to the left). Note, in F, presence of periodic banding on enamel (arrows), mimicking lines of ridgedcementum in Isolobodon; note also, in F' (cast), the organization of interflexid cementum. All scales are5 mm.

NO. 31416.


and Hill, 1980; Reig, 1981; Patterson andWood, 1982; Woods, 1989a, 1989b). Woods(1989b) has recently considered these issuesin some detail, and has concluded that cap-romyids are derived from echimyids (or acommon ancestor of echimyids, myocasto-rids, and octodontids). Since capromyids havenot been identified paleontologically or neon-tologically in surrounding mainlands, this ar-gument carries the implication that capro-myids must have evolved wholly within theGreater Antilles. If Capromyidae as usuallydefined is a holophyletic group restricted tothe Greater Antilles and some nearby islands,then based on our reconstruction ofAntilleanfaunal history this clade could not have orig-inated earlier than about 42 Ma- i.e., the be-ginning of the Late Eocene, the earliest timeat which it would have been feasible for landmammals to take up permanent residency(see Discussion).

PRIMATES

ANTHROPOIDEA: PLATYRRHINI

Unnamed Genus and Species A (Primate)

MATERIAL: Right astragalus (MNHNH P3059; table 2, fig. 2), found in lag deposit nearsouth end of Domo de Zaza section by 0.Jimenez in November 1993.PRELIMINARY EVALUATION: At the time of

its discovery in 1993, MNHNH P 3059 wasthought to be morphologically distinct fromastragali of all known mainland New Worldprimates, including near-coevals from Pata-gonia like Carlocebus and Soriacebus(MacPhee and Iturralde-Vinent, 1995). Morerecently, a partial astragalus (MNHNHV 205)attributable to the Cuban monkey Paralouat-ta varonai (fig. 3) has turned up in a faunalcollection from Cueva Alta (for site descrip-tion, see Jaimez Salgado et al., 1992). Thetwo elements are extraordinarily similar insize and details of shape, which suggests (un-surprisingly) that they represent the samenarrow monophyletic group.The following morphological description

uses the terminology and measurementmethods of Meldrum (1990); table 2 shouldbe consulted for measurement values and ac-ronyms. In astragalar length, the Zaza spec-imen places near the upper end of the size

distribution in extant platyrrhines as pre-sented by Meldrum (1990), being exceededin this regard only by Ateles, Alouatta, andLagothrix (Brachyteles was not included).Comparable dimensions of the astragalusfrom Cueva Alta are slightly larger. The headofthe Zaza specimen is slightly damaged me-dially, but the navicular facet was clearlyovoid in shape with its long axis (as measuredby HDW) somewhat obliquely set. The headis missing from the Cueva Alta specimen.The necks of both specimens are compara-tively short but robust. The angle describedby the neck on astragalar body is approxi-mately 300 in each specimen (Meldrum'smethod). The anterior calcaneal facet of theZaza anthropoid is extensive proximally,showing no tendency to taper in the directionofthe deep astragalar sinus. Distally, it meetsthe navicular facet along a raised margin; anonarticular strip between facets is not pres-ent. The anterior calcaneal facet on the as-tragalus assigned to P. varonai is damaged,but there is no doubt that the facet was justas extensive proximally. The posterior cal-caneal facet is very large in both specimens."Waisting" in the medial third of the facet isslight, especially in the Paralouatta astraga-lus. Indeed, the ratios offacet length to width(PCL/PCW) in these specimens are consid-erably higher than in any ofthe taxa or fossilssampled by Meldrum (1990). The astragalarsinus is much deeper in the Zaza specimenthan in the one from Cueva Alta. By contrast,trochlear height (TH) is somewhat higher inP. varonai.

In both specimens, the trochlear articularsurface is high, parallel-sided (not "wedged"),and extensive from front to back. The troch-lear spool is moderately deep, and in proxi-mal aspect the medial rim is slightly higherthan the lateral. The tibial articular surfaceterminates proximally in a long, fluted shelfthat would have guided the tibial and fibularflexors. In P. varonai this area is broken andabraded. Distally, the tibial articular surfaceends more or less abruptly in both specimens.Medially, a shallow stop for the tibial medialmalleolus encroaches on the neck. The distalmargin of the lateral trochlear rim projectsbeyond the level of the astragalar body inboth specimens, but especially so in the CuevaAlta astragalus. The articular surface for the

71995


Fig. 2. Astragalus (MNHNH P 3059) of an anthropoid (unnamed genus and species A), from EarlyMiocene locality of Domo de Zaza, south-central Cuba. Views are A, dorsal; B, ventral; C, proximal;D, distal; E, medial; and F, lateral.

fibula is flat, but ventrad it spreads out lat-erally to a marked degree. The posterolateralprocess of Meldrum (1990) seems to havebeen absent in both specimens, unless thetiny eminence in the Cueva Alta specimencan be so regarded.The fact that there are only minor differ-

ences between these two astragali (e.g., slight-

ly higher trochlea, wider posterior calcanealfacet, shallower astragalar sinus in P. varonai)underlines the high probability that the spe-cies which they represent are closely related.In fact, this degree of similarity begs a ques-tion: if Zaza is confirmably late Early Mio-cene in age (MacPhee et al., in prep.), are thefossils from Cueva Alta truly as young as

8 NO. 3141

MAcPHEE AND ITURRALDE-VINENT: TERTIARYMAMMALS9

Fig. 3. Astraglus (MNHNH V 205) attributed to Paralouatta varonai, from Cueva Alta, Sierra delos Organos, western Cuba. Views are A, dorsal; B, ventral; C, proximal; and D, lateral. Shading denotesabraded areas.

Quatemary? Sloth bones from Cueva Alta(and Cueva del Mono F6sil, to which it isconnected by a fossiliferous fissure) werefound to be beyond the effective range of 14Cdating (M. Tamers, personal commun.), andno usable calcite-bone associations have been

found in these caves to make U/Th datingworthwhile. Accordingly, the Quaternary ageassessment for the monkey caves has to bebased on the character of the recovered fau-nule, which largely consists of taxa knownfrom localities of proven or suspected Qua-

TABLE 2Measurements (in mm) of Zaza and Cueva Alta Astragalia

Specimen L HDW HDH ACF PCL PCW TW TH ATW PTW W

MNHNH P 3059 20.0 9.7e 7.3e 12.7 9.5 5.5 11.7 10.8 9.2 9.1 16.5MNHNH V 205 - - - - 10.Oe 6.9 12.0e 11.9 10.5 10.5e 16.9e

a For landmarks used in measurement, see Meldrum (1990). Key: ACF anterior calcaneal facet; ATW anteriortrochlear width; HDH head height; HDW head width; L length; PCL posterior calcaneal facet length; PCW posteriorcalcaneal facet width; PTW posterior trochlear width; TH trochlear height; TW trochlear width; W width. The lettere indicates that the measurement is estimated (due to breakage).,

1995 9


Callicebus Cebus Ateles Alouatta

tA-CH~~

.Th

Q'Fig. 4. Astragali of representative extant platyrrhines. Sequence of views (top to bottom of page) is

dorsal, ventral, proximal, distal, lateral, and medial. Note individual scales.

ternary age elsewhere in Cuba (see JaimezSalgado et al., 1992). However, there are someunique occurrences: in addition to Par-alouatta, a previously unknown owl and slothhave also been recovered, from localities

within these caves (Oscar Arredondo, per-sonal commun.). Since the fissures from whichthe faunule was collected are not connectedwith the present sedimentary floors of thecaves, and since the cave entrances are sit-

10 NO. 3141

'N-1I


Saimiri Aotus

aXccs

QL_XAX

,(X

Fig. 4. Continued.

uated high above the present valley floor, itis distinctly possible that the fissure fillingssample a much earlier fauna.

Figure 4 provides schematic views of theastragali of some extant platyrrhines and aspecimen usually referred to Dolichocebusgaimanensis, an extinct Patagonian species

(cf. Meldrum, 1990). Several points emergefrom a comparison ofthese astragali with theZaza and Cueva Alta specimens. First, theCuban monkeys are not especially similar toany of the species illustrated (but see below).This applies in particular to Alouatta, the ge-nus originally thought to be the closest living

Callithrix Dolichocebus

1995

.001).......

.......................


relative ofParalouatta (Rivero and Arredon-do, 1991). Among other distinctive features,the astragalus ofAlouatta exhibits a "wedged"trochlea, low trochlear relief, indistinct mar-gins, vertical medial articular surface for thetibia, and a large posteromedial process. TheCueva Alta specimen lacks these features, asdoes the better-preserved astragalus fromZaza. What, then, is the proper sister groupof the Cuban monkeys?

In a cladistic analysis of a selection of cra-niodental characters, MacPhee et al. (1995)found that, among sampled Quaternary plat-yrrhines, the extinct Hispaniolan anthropoidAntillothrix (="Saimiri") bernensis groupedmost closely with Paralouatta varonai. TheseAntillean taxa joined Callicebus at the nextnode. By contrast, no close relationship be-tween the Antillean monkeys and atelines(including Alouatta) was indicted. Consistentwith the trend of the craniodental evidence,Callicebus shows more astragalar resem-blances to the Cuban fossils than atelines do.In extant titi monkeys, the trochlea is highand only slightly "wedged," trochlear wallsare subvertical, and calcaneal facet shape issubstantially similar to the Zaza and CuevaAlta specimens. However, correspondencesto Dolichocebus and other extant platyrrhinessuch as Saimiri (fig. 4) also exist, raising thedistinct possibility that much of this mor-phology is simply primitive (see also MacPheeand Iturralde-Vinent, 1995).We find that the Zaza and Cueva Alta as-

tragali fail to support an affiliation ofthe Cu-ban monkeys with either Alouatta or atelinesin general (cf. Ateles, fig. 4).3 The hypothesisthat the closest living relative of the Cubanand Hispaniolan monkeys is Callicebus, assuggested by craniodental characters(MacPhee et al., 1995), is plausible but re-quires additional evidence and testing.

3The only biogeographical reason for considering ate-lines in the context of platyrrhine evolution in Cuba hasdisappeared. The holotype and only known specimen ofAteles (=Montaneia) anthropomorphus-the supposedlyendemic "Cuban spider monkey"-is less than threecenturies old. It probably represents an imported A. fus-ciceps, as has long been suspected on morphologicalgrounds (see MacPhee and Rivero de la Calle, 1995).

AN EARLY OLIGOCENE SLOTHFROM PUERTO RICO

While engaged in a preliminary survey insouthern Puerto Rico in March 1994, ourfield party discovered a fossiliferous site nearthe town of Yauco, in the southwestern partof the island. Among the vertebrate fossilswe recovered was the proximal part of amammalian femur, here referred to the xe-narthran clade Tardigrada. This discovery of-fers the hope that Puerto Rico can be tiedinto the paleobiogeographical picture beingdeveloped for the other Greater Antilles(MacPhee and Wyss, 1990; MacPhee andIturralde-Vinent, 1994). Of equal interest isthe apparent age of the Yauco sediments. Aswe note in greater detail below, fossil evi-dence indicates that these sediments are EarlyOligocene (Globigerina ampliapertura zone,or about 33-34 Ma according to the timezonation utilized by Berggren et al. [1992]).Yauco may therefore be older than the Zazalocality by 15 Ma or more, making it by farthe oldest land vertebrate site with a well-constrained age in the West Indies. Oddlyenough, if the femur is that of a megalony-chid, as we think is highly probable, it be-comes the oldest specimen so far referred tothat family (cf. Pascual et al., 1990). Thismeans very little by itself, other than to un-derline the fact that tardigrades had to havebeen well diversified and widely distributedin the Neotropics before the Oligocene.

XENARTHRA

TARDIGRADA: ?MEGALONYCHIDAE

Unnamed Genus and Species B (Tardigrade)MATERIAL: Left proximal femur (AMNH

VP 129883, fig. 5), preserving head, neck,and trochanters. Found by C. Flemming inMarch 1994.PRELIMINARY EVALUATION: Our criteria for

assigning this specimen to Mammalia are asfollows: (1) greater and lesser trochanterslarge, projecting, and situated on a markedoblique; (2) medullary cavity not densely filledwith cancellous bone; (3) proximal articularend ovoid, with well-defined border markingattachment ofjoint capsule; and (4) large fo-vea in the center of the femoral head. Cor-porately, these features indicate that AMNH

NO. 314112


VP 129883 represents a mammal, and dis-tinguish it from conditions encountered inchelonians, crocodylians, and squamatans.

Referral to Tardigrada (i.e., the monophy-letic taxon containing all sloths, living andextinct) is considered reasonable because ofstrong resemblances to Quaternary and Re-cent sloths and because there is no other plau-sible candidate among other land mammaltaxa having Antillean representation (i.e., Li-potyphla, Primates, Rodentia). On biogeo-graphical grounds it is reasonable to believethat the Yauco sloth will prove to representa member of Megalonychidae, the only slothfamily that provably managed to colonizeWest Indian islands (for recent treatments ofAntillean and mainland members of thisfamily, see White [1993]). Consistent withthis is the fact that the specimen bears a par-ticularly close resemblance to the proximalfemora of certain living and extinct mega-lonychids (fig. 6). These resemblances in-clude (1) markedly open angle formed byfemoral neck and diaphysis, (2) position andorientation of the trochanters, (3) cross-sec-tional shape of the shaft, and (4) sphericalproximal articular surface. A minor differ-ence from extant Choloepus (and Bradypus)is the presence of a fovea for ligamentumteres; Quaternary Antillean megalonychidsare usually foveate (Fischer, 1971). The chiefdifference is size: the Yauco femur representsan animal as small as or smaller than livingBradypus, while Neocnus, Acratocnus, andCholoepus are considerably larger. Indeed, theYauco sloth would appear to be one of thesmallest fossil tardigradans on record (seeWhite [1993] for evidence ofvery small slothsin the Santacrucian of Argentina). In this re-gard it stands in marked contrast to EarlyMiocene Imagocnus, which appears to havebeen nearly the size ofMegalocnus (- 200 kgaccording to Paula Couto [1979]).The Yauco femur is morphologically closer

to Neocnus and Choloepus than it is to Acra-tocnus, the only Quaternary sloth known fromPuerto Rico. The phylogenetic significance ofthis (if any) is hard to assess in the absenceof adequate material. There are several dif-ferent and mutually exclusive views concern-ing the interrelationships of the Antilleansloths (e.g., Varona, 1974; Paula Couto, 1979;Arredondo, 1988), although we believe that

there is some support for brigading PuertoRican Acratocnus, Hispaniolan Synocnus, andCuban Neocnus and Miocnus in one unit(equivalent to tribe Acratocnini, Varona[1974]). If this grouping is monophyletic,acratocnins, like some other assemblages ofclosely related Antillean taxa, had a multi-island distribution; this regularity requiresexplanation (see Discussion).

LocALITY AND AGE: The sloth femur wasrecovered at LocalityAMNH PR 1994/1, sit-uated just south of Yauco on Highway 3133(figs. 7, 8). Sediments in this region are ap-portioned to Early Oligocene Juana Diaz Fmand an overlying but unnamed carbonate unit,as described in more detail below.The Yauco locality is situated within the

former "lower clastic member" of the JuanaDiaz Fm (Monroe, 1980). The Juana DiazFm has long been informally divided into twomembers, an upper carbonate (UC) unit anda lower clastic unit (Moussa and Seiglie, 1970,1975; Monroe, 1980). Frost et al. (1983) haveproposed that the Juana Diaz Fm (hereafterabbreviated as JD) be restricted to the lowerclastic member, a proposal that we follow inthis paper. The UC, allocated to a new for-mation as yet unnamed, correlates with theGloborotalia opima opima-Globigerina an-gulisuturalis zone and is presumed to be ofLate Oligocene age. The JD correlates withthe Globigerina ampliapertura zone and istherefore older (Pessagno, 1963; Seiglie andBerm(udez, 1969; Moussa and Seiglie, 1970,1975; van den Bold, 1971; Todd and Low,1979; Frost et al., 1983). Abundant plank-tonic forams reported from the JD includeGlobigerina ampliapertura, G. haoi, G. an-gustiumbilicata, G. ciperoensis, G. rohri, G.venezuelana, and Cassigerinella chipolensis(Seiglie and Berm(udez, 1969). Facies sam-pling very shallow water and terrestrial en-vironments have no planktonic forams, butdo yield Amphisteginafloridana, Angulogeri-na vicksburgensis, Baggina cojimarensis,Pararotalia mexicana and Lepidocyclina un-dosa, all ofwhich are typical Oligocene mark-ers (Todd and Low, 1979).The Yauco section exposes both JD and

UC (figs. 8,9). The JD belongs to the "inshoreshelf sandstones and conglomerates" faciesof Frost et al. (1983, their fig. 2). This latterunit, 60-70 m thick, is composed of clays,

131995


Fig. 5. Proximal part of a left femur (AMNH VP 129883) of a megalonychid sloth (unnamed genusand species B), from Early Oligocene Locality AMNH 1994/1, near Yauco, SW Puerto Rico. Views areA, anterior; B, posterior; C, lateral; and D, medial. Scale is 1 cm.

sands and gravels with some marly coquinaintercalations. The top of the unit is an ero-sional horizon where conglomeratic lime-stones of a few centimeters thickness mergeinto coralgal reef limestones of the UC. Thebase of the section was not observed.

Fine-grain sands and sandy clays, withabundant oxidized plant remains, comprisethe most common lithology. These beds formunits tens of centimeters thick, distinctly fin-ing upward and incorporating caliche andother indications of soil-forming processes.Small concentrations of gypsum, apparentlyepidiagenetic, occur as fracture fillings in someof the clays and sandy clays. Such featuresare consistent with terrestrial conditions andan oscillating freshwater table, as might occurin relation to swamps and lagoons close to ashoreline. Alluvial beds (figs. 8, 9) occur as

small, laterally overlapping river channels ofa few meters thickness, and are more frequentin the lower half of the exposed section.Near the base ofthe outcrop, poorly sorted

coarse-grained conglomerates occur as inter-calations, two to three meters in thickness,with clasts up to 15 cm in diameter. Theseconglomerates were probably deposited dur-ing flood events. In localities near Yauco, andalso to the east of the locality, conglomeratesat the base ofthe JD reach thicknesses greaterthan 50 m and are intercalated with red pa-leosols. It is possible that the basal parts ofthese conglomerates are as old as latest Eo-cene (cf. figs. 9, 10).Three marly coquinite horizons, two to

three meters thick and located near the base,middle, and top of the section, are formedmainly by fragmentary or complete tests of

NO. 314114


a b~~~~~~~~~~~~~

El 1SS @(

Fig. 6. Left femora of (a) Neocnus sp. (Arre-dondo coil.), (b) Acratocnus ondontrigonus (AMNHVP 17716.-i), and (c) Choloepus hoffmanni(AMNH M 139772), in anterior (top) and poste-rior (bottom) views.

Lepidocyclina undosa. Occurring in smallernumbers are shells ofgastropods (turritelids),bivalves (pectinids, ostreids, etc.), echino-derms, and crustaceans, with occasional sharkteeth and isolated coral fragments. TheseLepidocyclina coquinites have been inter-preted as forereef deep-shelf facies by Frostet al. (1983), but at Yauco the coquinite bedsare embedded in sediments deposited in anear-shore environment, with an inverte-brate faunal assemblage characteristic ofveryshallow water. Consistent with this interpre-tation is the discovery offreshwater/brackish

aSI-111 20

N -

Y JD67° 66°w

Fig. 7. Location ofreferenced localities in Cuba(top) and Puerto Rico (bottom). Key: CA, CuevaAlta; DZ, Domo de Zaza; Y, Yauco; and JD, JuanaDiaz.

water vertebrates (sirenians, crocodiles, andturtles) in the same sediments. The coquinitebeds probably represent short-lived, shallow-water marine invasions of the coastal plain.Further east, similar beds intercalated withsandy lenses outcrop along the Rio Jacaguas,from which sirenian fossils have been recov-ered (Matthew, 1916).

It is not yet possible to determine whethersome horizons at Yauco are more productiveof vertebrate remains than others. The slothfemur was discovered in the upper halfofthesection, within a lag developed from a clay/sandy clay/caliche sequence. A number ofar-ticulated sirenian bones, found in the middleofthe measured section, were associated withsandy clays evidently deposited in brackishconditions. A few well-preserved crocodileskull elements and fragments of turtle shellwere found in alluvial channel deposits nearthe base of the section.

In order to test age determinations of theJD in the Yauco section, four samples werecollected and analyzed for microfossil con-tent. Sampling horizons are indicated in fig-ure 8 as 94/1 6D (coquinite near base of JD),94/16C (clays 10 m above), 94/16B (middleportion ofJD), and 94/16A (base ofUC). Thesamples yield benthic forams, rare planktonicforams, fish teeth, echinoderm spines, and

1995 15


'(0).0Z

Co

Zo

0

EN

CaNE

<CS

Z o

NCZ

ILQN E- coD .C

<:z 0

< (D

meters0-

20 -

40 -

60

CONGLOMERATES

E2 SANDSTONES

SANDY CLAYS 80 -

LIMESTONES

COQUINITES

Fig. 8. Columnar section of Loc1994/1, near Yauco, SW Puerto Rfeatures are alluvial channels (see alsindicated are approximate positicyielding vertebrate fossils (O) andsapling horizons (0). Tardigradan fcovered from float.

arated by an erosional surface associated witha thin conglomeratic bed (figs. 8, 9).

DISCUSSION

GEOLoGICAL CONSTRAINTS AND LANDMAMMAL BIOGEOGRAPHY OF THE

94/16A GREATER ANTILLESIn an earlier paper (MacPhee and Itur-

ralde-Vinent 1994), we proposed that bothloth femur dispersal and vicariance have occurred in theAMNHVP12883) formation of the Antillean mammal fauna,

and that this point might be tested by anappropriate program of paleontological in-vestigations. It was inferred that the dispersal

,irenian skeleton phase occurred first and involved the move-ment of species en bloc out ofnorthern SouthAmerica (whence all known major taxa ex-cept Antillean lipotyphlans derive) and onto

94/16B subaerial landmasses in the central Carib-bean. How this was accomplished in detailis of course uncertain. However, unlike most

94/16C other proponents ofdispersal, we assume thatsome critical event occurred in the early Ce-nozoic which briefly altered the configurationof lands and the amount of open water that

5 94/16D had to be crossed. That is, dispersals werenot spread out over a long time span, but

crocodilian jaw instead occurred within a very narrow inter-val. The later, vicariant phase involved sub-division offounder clades4 by the erection of

cality AMNH water barriers produced by tectonic dismem-Lico. Lenslike berment and reconfiguration of the existingso fig. 9). Also chain of islands. This phase is elsewhere de-ins of factes scribed as "island-island" vicarianceminr wt reb- (MacPhee and Wyss, 1990); it differs from

ostracods (this last very abundant in sample94/16C).

In each horizon the assemblage of forams(table 3) is typical Lower Oligocene, becausethe Upper Oligocene markers Miogypsina andMiogypsinoides are absent. The absence ofthese marker taxa is consistent with the in-terpretation that these beds were depositedin the Early Oligocene, as indicated by theplanktonic foram data referenced above. Nolarge depositional hiatus is evident betweenthe JD and the UC units, because foramin-iferal assemblages are similar (table 3, sam-ples 94/16A and B), although they are sep-

4 Optimally, the number of "founder clades" recog-nized for Antillean land mammals ought to be equal tothe number of initiators (propagules) that were phylo-genetically separate from one another (i.e., separate spe-cies) at the time of original colonization. Our presentview is that a minimum of eight initiators has to beinferred to explain the existing land mammal evidencefor the Greater Antilles, distributed as follows: Primates(two initiators), Rodentia (three), Insectivora (two), Xe-narthra (one). There could, of course, have been morethan one initiator per lower-level monophyletic group,as some authors have inferred (e.g., Morgan and Woods,1986). For the Greater Antilles as geographically defined,accounting for late-evolving oryzomyins (Rodentia) re-quires a ninth initiator-but only for Jamaica, which isnot considered part of GAARlandia as defined in thispaper.

16 NO. 3141

c

A,111 L& YV" 1%,-


Fig. 9. Top, View of Locality AMNH PR 94/1, near Yauco, southwestern Puerto Rico; sectionexposing Early Oligocene coastal plain environment ofJuana Diaz Fm (sensu Frost et al., 1983). Bottom,View of ?latest Eocene-Early Oligocene poorly sorted alluvial conglomerates at base ofJuana Diaz Fm,Highway 52 near town of Juana Diaz, southern Puerto Rico; red sandy clays exposed beneath uncon-formity represent a paleosol. Coeval conglomerates onlapping older deposits are found in Puerto Rico,Cuba, and Hispaniola, where they represent subaerial erosion products formed during the phase ofactivelate Paleogene uplift discussed in text.

17.1995


TABLE 3Locality AMNH PR 1994/1 (Yauco): Identified Foraminifera

Samplesa

Taxa 94/16A(UC) 94/16B (JD) 94/16C (JD) 94/16D(JD)

Baggina, aff. B. xenoulab XLepidocyclina undosa'c X X XLepidocyclina gigasc XLepidocyclina, cf. L. canelleic XLepidocyclina, cf. L. giraudiac XLepidocyclina, cf. L. vaughani(?)c XLepidocyclina, cf. L. canellei or L. vaughanic XPararotalia mexicanab X XPararotalia, cf. P. guantanamensisb XGypsina sp.c XPalaeonummulites diusc X cf. XAmphistegina sp.c XGlobigerina, cf. G. angustiumbilicatab X

a For location of sampling horizons, see fig. 8.b Identified by Lic. Gena Fernandez (CIDP-Cupet, Cuba).c Identified by Prof. Edward Robinson (University of the West Indies, Mona, Jamaica).

Rosen's (1975, 1985) concept, which as-sumes that most of the critical vicariantly-controlled events in Caribbean biogeograph-ical history were "continent-island." New pa-leontological discoveries and reconsiderationofpertinent geological evidence permit someadditional elaborations, presented below.In our model we specifically proposed that

land mammal dispersal to the Greater An-tilles could have occurred across portions ofthe Aves Rise, if (as is often assumed) partsof this feature were subaerial for a limitedtime in the mid-Tertiary. Other possibilities,such as movements across the YucatanChannel, Nicaragua Rise, or Lesser Antilles,cannot be completely excluded but are muchless plausible for land mammals for one rea-son or another (briefly summarized below).Holcombe and Edgar (1990) considered

fairly elaborate scenarios in which islands ofvarious sizes and interconnectivities were re-constructed for the spine of the Aves Rise,assuming differing subsidence regimes. Theynoted in particular that the thin sedimentarydeposits covering large parts ofthe Aves Rise(<500 m of Late Cretaceous and Cenozoicsediment combined) imply that the Rise waspositive with respect to the sea floor duringmost of the Cenozoic. Other facts also sup-port the contention that a major part of theAves Rise must have once been emergent.

The simultaneous and extensive develop-ment ofcoarse-grained clastic deposits in theGreater Antilles and northern South Americaduring the latest Eocene-Early Oligocene (fig.10) indicates that active uplift took placethroughout the positive portions of the Ca-ribbean, including the Aves Rise, at essen-tially the same time. Likewise, recovery ofLate Eocene and Early Miocene shallow wa-ter limestones from the walls ofthe Aves Rise(Fox et al., 1971) confirms the positive char-acter ofthe Aves during the early and middleTertiary. An intriguing feature of the dredgesamples described by Fox et al. (1971) is thatno Oligocene rocks were identified in any ofthem. Although the evidence is of a negativequality, the absence of Oligocene rocks inthese samples correlates with the presence ofsame-age hiatuses and subaerially depositedconglomerates elsewhere in the Greater An-tilles (fig. 10). Taken together, this evidencetends to confirm the view that the Aves Risewas uplifted around the time of the Eocene-Oligocene boundary, although it is to be hopedthat some day this inference will be con-firmed by a coring program.The late Paleogene uplift event in the Ca-

ribbean region, which was coincidental withthe worldwide Pyrenean phase of tectogene-sis (Leonov and Khain, 1987; Brezsnyanszkyand Iturralde-Vinent, 1978, 1987), resulted

18 NO. 3141


MID-TERTIARY SEDIMENTARY ENVIRONMENTS AND STRATIGRAPHIC UNITS OF THE GREATER ANTILLES AND SOUTH AMERICA

Conglomerates: poorly sorted Sands and clays: shallow marine Limestones: shallow marine 1 Deep marine

related to active uplift and coastal plain environments shelf environments

Fig. 10. Mid-Tertiary sedimentary environments and stratigraphic units for selected areas in theGreater Antilles, Aves Rise, and northern South America (Venezuela). Note (a) correlated depositionof coarse conglomerates in the latest Eocene-Early Oligocene throughout the area, suggesting regionaluplift, followed by (b) marine sedimentation from mid-Oligocene through Middle Miocene, indicatingregional transgression. Under the dispersal hypothesis favored in this paper, the optimal period for landmammal colonization of the Greater Antilles was at or near the Eocene-Oligocene boundary; by theEarly Miocene transgression, the window was closed. Sources: Cuba-Iturralde-Vinent (1969, 1972),Nagy et al. (1983), Brezsnyanszky and Iturralde-Vinent (1978), Cobiella et al. (1977); Hispaniola-Lewiset al. (1990), Mann et al. (1992); Puerto Rico-Monroe (1980), Frost et al. (1983); Aves Rise-Fox etal. (1971), Holcombe et al. (1990); northern South America (Venezuela)-Anonymous (1970), Dallmusand Graves (1972).

in the creation of a large positive feature on

the Caribbean sea floor-GAARlandia,formed by the Greater Antilles Ridge and theAves Rise. Ridge and Rise are also related as

members of the original "Great Arc of theCaribbean" formed in the Cretaceous (Burke,1988). At present the chief subaerial com-

ponents of GAARlandia are Cuba, Hispan-iola, Puerto Rico, Virgin Islands, Aves Is-land, and some islands on the northern fringeof South America (e.g., parts of the Vene-zuelan Antillas Menores). Prior to the LateEocene, Cuba, northern Hispaniola, PuertoRico and the Virgin Islands were not linearly

distributed as they are today, but insteadformed a much more compact array, with theeastern islands being situated considerably tothe west of their present locations (Malfaitand Dinkelman, 1972; Ross and Scotese,1988; Van Fossen and Channell, 1988; Pin-dell and Barrett, 1990).

In addition to general uplift, we originallyposited that dispersals could be more pre-cisely correlated with the dramatic loweringof global sea level ca. 29 Ma identified byHaq et al. (1987), since a major regressionwould have increased the total land area overwhich animals might have dispersed. A nec-

T

-1ii-

II- :u 1.

T

I I

1995 19


essary corollary of this proposition is thatmammal fossils much older than mid-Oli-gocene time would not be expected to occurin Antillean localities. However, the Yaucosloth demonstrates that tardigradans were al-ready in GAARlandia by the Early Oligo-cene. Evidently, the mid-Oligocene sea leveldrop of -160 m did not initiate dispersal, atleast in the case of sloths. Either the sea leveldrop under discussion is not accurately dated,or was not global, or for some other reasondid not affect GAARlandia in the way orig-inally imagined.By contrast, the evidence for the wide-

spread effect ofuplift correlated with the Pyr-enean tectogenetic phase is consistentthroughout the sampled region (fig. 10). InCuba, there is considerable evidence for de-positional hiatuses and extensive develop-ment of coarse-grained, poorly sorted con-glomerates during the Late Eocene-Early Oli-gocene (Iturralde-Vinent, 1972; Nagy et al.,1983; Brezsnyanszky and Iturralde-Vinent,1978, 1985). The same is true for Hispaniola,where coeval uplift produced most of thepresent Cordillera Central and CordilleraOriental. At this time, northern Hispaniolaand the eastern end of Cuba were physicallyconnected as subaerial landmasses (Calais etal., 1992; Lewis et al. 1990; Mann et al., 1992;Iturralde-Vinent, 1994b). Coarse-grainedconglomerates and coastal depositional en-vironments certainly appear in Early Oligo-cene facies in Puerto Rico (fig. 9; Monroe,1980), and, as noted above, it is possible thatthe basal conglomerates of the Juana DiazFm are even older (latest Eocene). These ob-servations demonstrate that correlated upliftalso occurred in the eastern Greater Antilles,creating subaerial connections between theseislands as Meyerhoff (1933) originally sug-gested. One implication of this is thatGAARlandia might have acted as a barrierbetween Atlantic and Caribbean circulations,which would in turn help to explain such ap-parent anomalies as the striking differencebetween Oligocene faunal assemblages on thenorth and south sides of Puerto Rico (Bold,1971; Kaye, 1959). These observations areconsistent with Donnelly's (1989, 1990) in-ference, from analysis ofbiological silica con-tent ofpelagic sediments, that the Aves Ridge(and the nascent Lesser Antilles) acted as a

barrier to the circulation ofintermediate anddeep water at -40 Ma. Donnelly noted thatnothing more than a discontinuous island arcwas implied by this evidence. Whether or notsurface water circulation was broadly inter-rupted at this time has not been investigated.

If terrestrial colonizations could have oc-curred in the latest Eocene or Early Oligo-cene, why could they not have occurred ear-lier, such as in the Middle Eocene? Earliercolonizations cannot be ruled out, but evenif they occurred, it seems unlikely that anycolonizing clades of that age could have per-sisted into the later Cenozoic. Although sub-aerial landmasses ofsome sort existed withinthe Caribbean Sea from time to time fromthe Cretaceous onward, they would have beensubjected repeatedly to episodes of subsi-dence and transgression (most notably inMaastrichtian, Late Paleocene/Early Eocene,and Middle Eocene [Khudoley and Meyer-hoff, 1971; Iturralde-Vinent, 1982; Maur-rasse 1990]). Our conclusion is that mam-mals, at least, could not have successfully col-onized and survived on the nascent islandsof the Greater Antilles until these bodies be-camepermanently subaerial subsequent to thebeginning ofthe Late Eocene, when the wholebelt was tectonically deformed (Khudoley andMeyerhoff, 1971; Lewis et al., 1990; Itur-ralde-Vinent, 1982, 1994a).The last or NeoCaribbean tectonic phase

of Greater Antillean evolution, from Mio-cene to Recent, has been described in severalpapers (Mann and Burke, 1984; Larue et al.,1990; Mann et al., 1990). During this phase,uplift was followed by a general transgression(fig. 9). In addition, several pull-apart basinsand related features opened or expanded alongthe northern Caribbean plate boundary,among them the Cayman Trough southeastof Cuba (Iturralde-Vinent, 1988, 1991; Ca-lais and Lepinay, 1991), Mona Canyon (La-rue et al., 1990; Masson and Scanlon, 1991),and the Sombrero Basin-Anegada Passagesystem east of the Virgin Islands (Mann etal., 1990; Jany et al., 1990). As a result, deep-water channels and basins now divide theblocks comprising the Greater Antilles Ridgeinto several isolated tectonic units, some ofwhich have large subaerial portions. Themarked subsidence ofthe Aves Rise (by 400-1400 m from Miocene onward) would have

20 NO. 3141


Fig. 11. Section of slightly deformed Early-Middle Miocene clastic carbonate deposits, SE Cuba;viewpoint is from east. Sandwiched between subhorizontal beds, a thick horizon composed of largeclinoforms dips south (i.e., in direction of the Cayman Trough) at an angle of -50°.

prevented this structure from acting as a con-nector at any time in the Neogene (Holcombeand Edgar, 1990; Holcombe et al., 1990).Evidence for the factors that produced the

NeoCaribbean tectonic event is well repre-sented in southeastern Cuba. Along the is-land's southern flank (=north wall ofthe Cay-man Trench), various compressional featuresare recorded in the rock record, includingoverthrusting, reverse faulting, tight folding,and strong fracturing of latest Eocene andolder rocks (Lewis and Straczek, 1955; Cob-iella et al., 1977). By contrast, Miocene andyounger deposits are undeformed, showingonly a few degrees of tilting and deepeningtoward the south (Lewis and Straczek, 1955;Cobiella et al., 1977), probably in relation toa transtensional tectonic environment. Low-er-Middle Miocene clastic rocks outcroppingin fault-bounded grabens in southeasternCuba display very large south-dipping cli-noforms, indicating the existence nearby ofa steeply walled deep (fig. 11). The only pos-sible candidate is the Cayman Trough, whichmust therefore have come into existence aboutthis time (Iturralde-Vinent 1991, 1 994b). Thechange in tectonic regime from transpres-sional (pre-Miocene) to transtensional (Mio-cene and thereafter) recorded in southeastern

Cuba probably represents a rotation of thedirection of stresses throughout the Carib-bean, from ENE-WSW to more or less di-rectly E-W. The changeover correlates wellwith the Savic tectogenetic event at the Oli-gocene-Miocene boundary recorded else-where in the world (Schwan, 1980; Leonovand Khain, 1987). A significant consequenceofCaribbean stress-field rotation was disrup-tion and dispersion ofthe terranes that formedHispaniola-Puerto Rico-Virgin Islands (Lewiset al., 1990; Mann et al. 1990; Larue and Ryan,1990; Larue et al., 1990) along the northernboundary of the Caribbean Plate. Equivalentdisruptions occurred along the southernboundary ofthe Caribbean Plate, affecting Cu-ra,ao, Aruba, and Venezuelan Antillas Me-nores (Dailmus and Graves, 1972; Pindell andBarrett, 1990; Mann et al., 1990).

Finally, note must be made of the severeconstraints operating on other apparentlyplausible routings for biotic immigration:

Central America(Nicaragua Rise-Cayman Ridge)

The Nicaragua Rise and Cayman Ridgewere evidently a single geological entity be-fore the opening ofthe Cayman Trough; bothhave vulcano-plutonic arc rocks oflatest Cre-

211995


taceous-Paleogene age as partial basement(Holcombe et al., 1990; Holcombe and Ed-gar, 1990). The Nicaragua Rise is sometimescited as a possible avenue for mammals intothe Greater Antilles via Jamaica (see discus-sion by Buskirk, 1985), often in connectionwith an assumed North American origin forAntillean lipotyphlans. However, emergenceof the entire Nicaragua Rise-Cayman Ridgecomplex seems unlikely because each com-ponent is draped by later Cenozoic marinedeposits, implying submergence during theperiod of interest. These sediments are Eo-cene and younger in the case of the CaymanRidge, Eocene through Quaternary on theNicaragua Rise near the Central Americanmainland, and Eocene through Miocene inJamaica (Holcombe et al., 1990; Maurasse,1990). Although there are certain faunal sim-ilarities between Jamaica and the rest of theGreater Antilles, there are also substantialdifferences (e.g., absence of insectivores)which imply that its contacts with other is-lands have been limited (Buskirk, 1985; Mor-gan, 1993).Yucatan Channel

Offshore seismic and sonar research in thewestern Caribbean Sea (see especially Case etal., 1990; Holcombe et al., 1990) conclusivelydemonstrates that a land connection did notexist between western Cuba and the YucatanPeninsula during the Tertiary. The topographyof the basin between westernmost Cuba andCozumel consists of a series of unconnectedridges separated by deep trenches, some ofthem>2 km below sea level. Therefore, sea-leveldrops, even ofgreat magnitude, would not haveappreciably affected the paleogeography ofthisarea. In any case, latest Eocene and Oligocenesinistral strike-slip motion along the Pinar Faultwould have acted to widen, not shorten, theseparation between Cuba and the Yucatan Pen-insula (Iturralde-Vincent, 1994a). It is possiblethat a much older connection (?late in the Mid-dle Cretaceous) between Central America andwesternmost Cuba once existed, since meta-morphic rocks recovered from ridges in theYucatan Channel are similar to those outcrop-ping in the Maya Mountains of Belize and inwestern Cuba (Pindell and Dewey, 1982).However, even ifthis connection involved sub-aerial exposure of lands, it was far too early to

have influenced the dispersal ofthe land mam-mal groups known from the Greater Antilles.

Other Possibilities

Although the Lesser Antilles seem to bewell situated to act as a discontinuous cor-ridor across which mammals might have mi-grated from South America, the evidence isagainst any taxa having done so at a pointearly enough to comport with the fossil evi-dence now coming out of the Greater Antil-les. Most ofthe evolutionary development ofthis arc occurred subsequent to the latest Eo-cene-Oligocene (Maury et al., 1990). Nev-ertheless, a very limited suite ofmammaliantaxa got partway up the chain by some mech-anism. This suite includes a sloth and a capy-bara in Grenada during the Pliocene (R. Singer,personal commun.) and sigmodontines in thenorthernmost Lesser Antilles during the Qua-ternary (Morgan and Woods, 1986). Sloths, ofcourse, occurred in the Greater Antilles, butsigmodontines did not, except in Jamaica whereone species is known from Recent deposits(Morgan, 1993). Another imponderable is thatAnguilla and St. Martin, but no other LesserAntillean islands, bore a heptaxodontid rodent(Amblyrhiza) related to Elasmodontomys fromPuerto Rico. Each of these cases represents aninterpretative problem, but it needs to be notedalso that several ofthese "anomalous" taxa arevery poorly described (or even undescribed)and additional information on their phyloge-netic relationships is sorely needed.The Beata Rise, south of Hispaniola, does

not physically extend as far as South America(Case et al., 1990). Its age is uncertain, al-though Santonian and younger sediments oc-cur on the structure (Holcombe et al., 1990).Furthermore, its aspect as a submarine prom-ontory of Hispaniola may be a relatively re-cent phenomenon (Mann and Burke, 1984;Mann et al., 1990). The Rise has subsided- 1000 m since the Middle Eocene; but sincemost of the Ridge lies at -2500 to -3000m, its opportunity for subaerial exposure musthave been limited (Holcombe et al., 1990).

SIGNICANCE OF FOSSILLAND-MAMMAL DIscovERIEs

Three ofthe four major clades ofAntilleanmammals known from the Quaternary- Tar-

22 NO. 3141


digrada, Caviomorpha, and Platyrrhini- arenow known, for certain, to have existed onlandmasses in the central Caribbean Basinduring the mid-Tertiary (table 1). The onlymajor taxon now missing from the Tertiarypicture is Lipotyphla, and its absence is surelydue to the vagaries of fossil discovery, notactual absence. It is equally important to notethat, by contrast, nothing of an unexpectednature has yet shown up -no marsupials, nocarnivores, no condylarths, and so on (cf.Perfit and Williams, 1989). In short, to thedegree that the few fossil discoveries madeto date can be considered representative ofmid-Tertiary diversity, the Antillean landmammal fauna was essentially the same thenas now in terms of major taxa. A similarpattern appears to be emerging for other ver-tebrate groups, in the sense that specimenscomplete enough to permit confident assign-ment to named lower-level groups have allturned out to be referable to extant cladeshaving Antillean representation (table 1). TheTertiary arthropod fauna conflicts with thisneat picture to some degree, for it includesseveral "unexpected" taxa that are not partof the modern Neotropical fauna (Wilson,1988; D. Grimaldi, personal commun.). Inaddition, according to Graham (1990), theTertiary flora of the islands has a distinctlyNorth American aspect, quite unlike themodern flora which is overwhelmingly SouthAmerican in affinity. However, as Borhidi(1985) notes, in addition to North and SouthAmerican components there are also ele-ments ofAfrican, Pacific, and Tethyan origin.It is to be hoped that additional discoverieswill add to the database needed to resolve thecomplex botanical history of these islands.

ZAZAMYS FROM CUBA

Zazamys veronicae is a true biogeograph-ical "ghost"-that is, an extinct member ofa clade that lived outside the clade's supposeddistribution area (and whose existence wasunsuspected prior to the discovery of fossilmaterial). Quaternary representatives of Iso-lobodontinae are known from Hispaniola,Puerto Rico, and some of the Virgin Islands,but because no members of this clade sur-vived into Pleistocene/Recent time in Cuba,they were assumed never to have occupiedthat island (e.g., Simpson, 1956).

How did isolobodontines reach Cuba? Onepossibility is that Zazamys veronicae repre-sents a unique occurrence of isolobodontinesin Cuba, its proximate ancestor having beenrafted in from Hispaniola, the alleged centerof isolobodontine diversity (Woods, 1989a).A difficulty with this interpretation is thatZazamys is morphologically more primitivethan Isolobodon, its apparent sister taxon,which -reversing the logic implicit in center-of-diversity argumentation -indicates thatHispaniolan isolobodontines are derived froma Cuban source.A more interesting possibility is that the

complementary distribution ofZazamys andIsolobodon is strictly due to vicariance, i.e.,the subdivision of populations through theerection of a barrier, in this instance the sub-division ofland. Island-island vicariance is ro-bust in that it provides a common-cause ex-planation for a number of distributional fea-tures and potential sister-group pairings. Asnoted above, there is solid geological supportfor the contention that the Windward andMonaPassages did not exist as such prior to the Mio-cene. Thus, during the critical periodwhen landmammals were most likely to have entered theGreater Antilles- latest Eocene through EarlyOligocene (fig. 10) -Cuba, northern Hispan-iola, and Puerto Rico probably constituted asingle landmass (MacPhee and Iturralde-Vi-nent, 1994). However, this situation could nothave prevailed for a lengthy period: Cuba wasalready separated into three large archipelagosby the beginning of the Early Miocene (Itur-ralde-Vinent, 1969), and eastern Cuba andnorthern Hispaniola would no longer have beenin contact by this time (Calais et al., 1992).This information suggests that the "orderly"distribution ofmany Quatemnary Antillean taxamay not have been due to dozens ofindividualrafting events among the various islands, cre-ating a superficial, ahistorical faunal similarity,but instead to the dissociation of the islandsthemselves as they progressively parted com-pany with their resident biotas already in place.Some sister-group pairings already recognizedor commented upon in the literature that maybe a result of vicariant events affecting Cubaand Hispaniola include Miocnus + Synocnusin Megalonychidae (cf. Varona, 1974), Neso-phontes micrus + N. paramicrus in Neso-phontidae (cf. Patterson, 1962), and Par-

1995 23


alouatta varona + Antillothrix bernensis in Pla-tyrrhini (MacPhee et al., 1995). For the reasonscited above, island-island vicariance also ac-counts for the existence of Zazamys in Cuba.It can be predicted that other "ghosts" havingclosest relatives on adjacent islands will befound, as the Antillean fossil record continuesto improve.Zazamys has a direct bearing on another

problem, which is the estimation of the an-tiquity of certain clades ofAntillean rodents.Previously, Woods (1982, 1989a) inferredthat the rodent colonization(s) took place inthe medial to later Oligocene, which is inreasonable agreement with the evidence pre-sented here. However, in his most detailedanalysis of the problem, Woods (1989b: 83)concluded that Antillean capromyids andechimyids are derived from an echimyid an-cestor that

... invaded the Greater Antilles from SouthAmerica either via island-hopping through theLesser Antilles or by rafting directly to PuertoRico or eastern Hispaniola, which has servedas the center of evolution for capromyids. Thetime of this invasion is unknown but all avail-able data suggest it was after the Oligocene andprobably as recently as the late Miocene. Thefirst radiation gave rise to plagiodontine, hex-olobodontine, and isolobodontine lineages onHispaniola, and later (Pliocene and Pleistocene)to the capromyine lineage that inhabits Cuba,Jamaica, and the Bahamas.

I propose that spiny rats (Heteropsomyinaetentatively retained in the Echimyidae), hutias(Capromyidae sensu stricto), and giant hutias(Heptaxodontidae sensu lato) all invaded theGreater Antilles sometime after the middleMiocene.

Zazamys proves that the minimum age forthe start ofthe invasion, however it may haveoccurred, could have been no later than EarlyMiocene, not Middle or Late Miocene, andfor geological reasons was assuredly earlierstill. Furthermore, because it is identifiablyisolobodontine, the morphology ofZazamysdemonstrates that the cladistic split betweenisolobodontines and their sister taxon withinCapromyidae must also have occurred beforethe late Early Miocene. Since there are noTertiary fossils for other groups of Antilleanrodents, it might still be maintained in a dis-

persalist hypothesis that they arrived later.However, this introduces new complications,since in the case of Capromyidae, the non-isolobodontine subfamilies as defined byWoods must have originated as early as or ear-lier than Isolobodontinae. Where were th. y, ifnot in the Antilles? Similarly, with the discov-ery ofan Early Oligocene sloth in Puerto Rico,Woods' (1989a: 771) speculation that slothsmay not have entered the Greater Antilles untilthe Late Miocene can also be rejected.

ZAZA ANTHROPOID

Too little is known of the Zaza primate tobuild a picture of its phylogenetic relation-ships, but its similarity to Paralouatta (andthe latter's distinctiveness within Platyrrhini)raises one issue that needs brieftreatment. Inthe systematic literature concerning endemicWest Indian monkeys, it is frequently as-sumed that these monkeys have to be thelittle-differentiated, insular representatives ofRecent mainland clades. Thus although Wil-liams and Koopman (1952) recognized themorphological distinctiveness of the holotypeof Xenothrix mcgregori and were ultimatelyunable to fit it within the confines of any ex-isting subfamily, they still concluded that itcould be comfortably regarded as "cebid." Ina similar vein, Rosenberger (1977) and Rosen-berger et al. (1990) argued that Xenothrix is akind ofpitheciine because ofsimilarities injawshape, despite the absence of detailed resem-blances in molar morphology. Likewise, Ri-moli (1977) preferred to deposit the unusualmonkey maxilla from Cueva Berne within Sai-miri (as S. bernensis) rather than consider themore radical alternative that it represented adistinct clade. In a rather different way Fordand Morgan (1986) and Ford (1990) have alsoperpetuated this approach, although they haveemphasized morphology over systematic affil-iations.The assumption that -ifwe just look long

and hard enough -every Antillean fossilmonkey can be shoehorned into one or an-other extant clade of mainland platyrrhinesat a low categorical level is no longer defen-sible in either theory or practice. Discoveryof the Zaza anthropoid demonstrates that

NO. 314124


platyrrhines were certainly present in theGreater Antilles at the start of the Neogene,and probably considerably earlier. Ifthe Zazaanthropoid is a member of the same mono-phyletic group as the one recently recognizedfor Paralouatta and Antillothrix by MacPheeet al. (1995), the implication is that the cla-distic event separating this group from itsclosest mainland relatives must have been atleast as early-making it one of the oldestnodes yet recovered within Platyrrhini.

SUMMARY

Recent discoveries in Cuba and Puerto Ricoestablish that the initiators ofmost (and per-haps all) founder clades of Antillean landmammals were emplaced on subaerial por-tions of GAARlandia by the mid-Tertiary.Evidence for a specific horizon in the mid-Tertiary is still thin, but in our opinion theimmigration event(s) occurred at or near theEocene-Oligocene boundary, probably with-in a very narrow interval. We consider it un-likely that successful emplacements couldhave occurred much earlier than Late Eocene

because ofthe lack ofpermanent landmasseswithin the Caribbean Sea. Since the emergentparts of the Greater Antilles Ridge were in aclose-packed array until the commencementof the Neogene, it is reasonable to believethat initiator populations could have beenwell distributed across GAARlandia. Withthe fragmentation of GAARlandia, progres-sive divergence of vicariated lineages wouldhave begun; also, some island lineages wouldlater become extinct (e.g., isolobodontines inCuba). It is possible that fossil members offounder clades will eventually be discoveredin each of the modem Greater Antilles, withthe possible exception ofJamaica, whose pa-leoposition during the Eocene-Oligoceneneeds to be better constrained in our opinion.Other mammalian clades may have dis-persed at the same time as the familiar An-tillean clades that survived into the Quater-nary, but if so they must have died out soonafter arrival. No such clades have yet beenidentified. Our model may or may not begeneralizable to other Antillean vertebratetaxa; this will have to be settled by the ap-propriate specialists in these groups.

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