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12-1-1999 Late Quaternary and Last Occurrence Dates From Caves at Barahona, Donald A. McFarlane Claremont McKenna College; Pitzer College; Scripps College

Recommended Citation McFarlane, D.A. "Late Quaternary Fossil Mammals and Last Occurrence dates from Caves at Barahona, Puerto Rico." Caribbean Journal of Science 35.3 (1999): 238-248.

This Article is brought to you for free and open access by the W.M. Keck Science Department at Scholarship @ Claremont. It has been accepted for inclusion in WM Keck Science Faculty Papers by an authorized administrator of Scholarship @ Claremont. For more information, please contact [email protected]. Caribbean Journal of Science, Vol. 35, No. 3-4, 238-248, 1999 Copyright 1999 College of Arts and Sciences University of Puerto Rico, Mayagu¨ez

Late Quaternary Fossil Mammals and Last Occurrence Dates from Caves at Barahona, Puerto Rico

DONALD A. MCFARLANE

W. M. Keck Science Center, The Claremont Colleges, 925 North Mills Avenue, Claremont CA 91711. [email protected]

ABSTRACT.—Puerto Rico supported at least five genera of endemic terrestrial mammals in the late Qua- ternary, all of which are extinct. Whether these died out in the late , the mid-, or in post-Columbian time has not been established. This paper is the first attempt at radiometrically dating the ‘last occurrences’ of these taxa, together with the first unambiguous descriptions of localities reported by previous workers. Last occurrence dates for , Elasmodontomys and Heteropsomys are shown to be mid-Holocene and overlap with Amerindian occupation of the island. is known only from the . No Puerto Rican has been shown to have survived into the historic (European) era, which contrasts with the situation on some other islands of the .

RESUMEN.—Durante el cuaternario tardı´o Puerto Rico tenı´a por lo menos cinco ge´neros ende´micos de mamı´feros terrestres, todos extintos. No se ha determinado si estos animales se extinguieron durante el pleistoceno tardio, el holoceno medio o luego de la llegada de los colonizadores. Este artı´culo es el primer intento de establecer por medio de pruebas radiome´tricas la u´ ltima presencia de estos ge´neros, ası´ como de proveer la primera descripcio´n precisa de las localidades informadas por otros investigadores. La u´ ltima presencia de Nesophontes, Elasmodontomys y Heteropsomys corresponde al Holoceno medio y coincide con la ocupacio´n indı´gena de la isla. Acratocnus se conoce solamente del pleistoceno tardı´o.Nosehadesmo- strado que alguno de los taxones puertorriquen˜ os hayan sobrevivido durante la era histo´rica (europea), lo que contrasta con la situacio´n en algunas de las otras islas de las antillas.

INTRODUCTION Rico. The purpose of this paper is to defini- tively characterize these sites for the first The late Quaternary endemic time, and to place the faunal remains in a of Puerto Rico as currently recog- radiometric context. nized consists of 15 of (Koop- A remarkable characteristic of the en- man, 1989) and 5 terrestrial genera repre- demic terrestrial mammal fauna of the senting 3 orders. All of the terrestrial taxa West Indies is the severity of the wave of have been extinct since at least the early that decimated it in late Quater- historic period, defined here as beginning nary time. MacPhee and Flemming (1997) in AD 1500. The earliest investigations of noted that 38% of all historic-era mamma- the extinct mammals were undertaken in lian extinctions worldwide impacted the 1916 by Harold E. Anthony, as part of a West Indies, even though these islands ac- wider investigation of the island sponsored count for only 1.4 % of the global land area by the New York Academy of Sciences (An- and less than3%ofrecent global terrestrial thony, 1916, 1918, 1925, 1926), and were fol- mammalian genera. In the Lesser lowed by workers from the University of and in Puerto Rico, this toll Kansas in 1957, the Smithsonian Institution reaches 100% (Table 1). The causes of these in 1976-77, and the University of Kansas extinctions have been widely speculated again in 1978 (Pregill, 1981). Following An- upon, and typically include climate change thony’s success, each expedition focused on (Pregill and Olson, 1981), habitat loss (Mor- a group of caves southwest of Barrio Bara- gan and Woods, 1986), introduced diseases hona, near Morovis in north-central Puerto (MacPhee and Marx, 1997), introduction of 238 FOSSIL MAMMALS FROM PUERTO RICAN CAVES 239

TABLE 1. Extinction statistics for West Indian terres- STUDY SITES AND RESULTS trial mammals, by island. Quaternary Mammal Localities at Barahona Late Pleistocene The Quaternary sites of inter- Diversity est are a group of several dozen caves and Island Group (genera) % Extinction rockshelters cut into the cone karst of the 3 100 Lares Limestone, 2.5 kms northwest of the Puerto Rico 5a 100 town of Ciales (Fig 1). Four of these caves 5 80 have yielded sufficient fossil vertebrate ma- 11 82 terial during this study to warrant further 12b 83 2 100 discussion. Topographic coordinates refer to the Universal Transverse Mercator Grid aexcludes the extinct Isolobodon portoricensis, gener- (North 19), and were obtained with a dif- ally considered to be an Amerindian introduction. ferentially-corrected global positioning sat- bexcludes two nominal taxa of undescribed extinct , whose validity is undetermined. ellite system to a precision of < 1m. Cueva Del Perro 2030686 N; 769482 E. (Fig. 2) feral or exotic predators and competitors (especially rats and the Indian mongoose; J. W. Bee of the University of Kansas first Woods et al., 1985), and direct extermina- investigated this cave in 1957, screening tion by humans (Martin, 1984). The quality some “4 cubic yards of sediment to a depth of evidence used to account for these ex- of four feet.” The bats and ma- tinctions varies tremendously, but in only a terial were described by Choate and Birney few cases can it be considered conclusive. (1968; see Table 2). Unfortunately, Bee did The introduction of rats ( rattus, R. not excavate by stratigraphic unit and his norvegicus, R. exulans) and the Indian mon- collection represents a time-transgressive goose (Herpestes auropunctatus) onto tropi- sequence. cal islands is a case in point. The impact of Examination of the site in 1996 revealed Rattus and Herpestes on ground-nesting that a significant portion of undisturbed is well established (Atkinson, 1985), sediment remains in the cave. The section but their impact on other terrestrial verte- exposed in the face of Bee’s pit consists of brates is much less so. For example, the An- approximately 10 cm of sediment with tigua Racer Alsophis has survived (barely) sparse vertebrate remains, underlain by a on Antigua in the presence of Rattus, Her- distinctive 10 cm thick layer (‘layer B’) of pestes, and almost total clearance of the na- land snails and vertebrate bones. Below, tive habitat, and yet is extinct on Barbuda, the sediment continues to the floor at ap- where Herpestes is absent and the vegeta- proximately 1.2 meters with occasional ver- tion largely uncleared. On Christmas Island tebrate remains. Layer B is clearly an owl- (Indian Ocean), the extinction of two native pellet accumulation dominated by small species of rats is tied closely to the arrival of birds with smaller quantities of (in- R. rattus and, possibly, the introduction of cluding Heteropsomys and occasional Elas- rat-borne disease (Andrews, 1909; Flan- modontomys remains) and material, nery, 1990). The impact of introduced Rat- all of which which may have been trans- tus on native Caribbean has not ported from the entrance chamber. An ac- been properly addressed. celerator radiocarbon date of 5410 80 radio- In Puerto Rico, the loss of one ground carbon before present (rcyrbp) (Beta , at least two native rodents, and an Analytic, Coral gables, Florida, #60893) was insectivore has not been evaluated radio- obtained on bone collagen from the metrically and thus could not hitherto be middle of layer B. reasonably assigned to a cause or causes. The new last-occurrence dates reported Cueva Clara 2030126 N; 769400 E. (Fig 3.) here begin to establish the necessary radio- This cave was first investigated by An- metric framework. thony in 1916, and re-worked by Bee in 240 D. A. MCFARLANE

FIG. 1. Location map of the Barahona caves. Area mapped is the 1 km2 UTM grid whose coordinates are shown. Cave entrances are shown as open circles and asterisks. A: Cueva de la Vaca, B: Cueva Clara, C: Cueva del Perezoso, D: Cueva del Perro, E: Nesophontes Cave (c.f. Pregill, 1981), F: Blackbone Cave (c.f. Pregill, 1981).

1957. of bats, the insectivore Neso- material of rodents and bats (Table 2). A phontes, the rodents Elasmodontomys and radiocarbon date on Pluerodonte shell car- Heteropsomys, and the Acrato- bonate from a depth of 30 cm and immedi- cnus were recovered from different parts of ately adjacent to Elasmodontomys material the cave without stratigraphic control. yielded an ‘infinite’ age of >34,650 rcyrbp When examined in 1996, very little original (Beta # 94478). Previous work has demon- sediment remained, but excavation of an strated that Pluerodonte exhibit a ‘limestone undisturbed pocket yielded well preserved effect’ (Goodfriend and Stipp, 1983) of the FOSSIL MAMMALS FROM PUERTO RICAN CAVES 241

FIG. 2. Plan view of Cueva del Perro. Map symbols and conventions follow Dasher (1994). of 0-3000 excess rcyrbp, or less than Cueva Del Perezoso 2030224 N; 769302 E. 10% of the radiocarbon ages reported here. This is a previously unworked site lo- cated in 1996. A calcite-indurated cave Cueva de la Vaca 2030161 N; 769331 E. mud, some 15-25 cm thick, overlies the (Fig 4.) floor of a shallow cave at its mouth, and is This cave was apparently not excavated protected by large slabs of limestone roof Acratoc- by either Anthony or Bee. It may have been fall. Excavation at this site yielded nus Elasmodontomys one of the caves visited by the Smithsonian and remains (Table 2). party in 1976-77, but no adequate site docu- Acid-dissolution of a bone fragment mentation of this work exists. Several cubic yielded no collagen residue and the site re- meters of sediment had previously been re- mains undated. moved from this site, but undisturbed sedi- ment remains around the periphery. Dur- DISCUSSION ing work in 1996 and 1998, Elasmodontomys, Heteropsomys, Nesophontes and Acratocnus, It is an axiom of that the together with two species of , were re- fossil record is unlikely to yield the last in- covered from the upper 45 cm of the de- dividuals of a lineage, so extinction dates posit (Table 2). The commonest taxon, Elas- must be inferred from ‘last occurrence’ modontomys, occurs to within 5 cm of the dates. Where a sequence of independent surface. A radiocarbon date on Pluerodonte dates is available, it may be possible to es- shell carbonate from 12 cm depth gave an timate the statistical confidence limits age of 33,670 370 rcybp (Beta # 95854). around an inferred extinction date (McFar- 242 D. A. MCFARLANE

TABLE 2. Paleofaunal recoveries from Barahona Holocene, and the historic era (post AD caves. 1500). The mid-Holocene extinctions are particularly poorly constrained by a short- CC CP CV CPZ age of radiometric dates and may prove to EDENTATA be illusionary as additional dates become Acratocnus available. odontrigonus XXX Nesophontes Acratocnus edithae XXX CHIROPTERA The Puerto Rican ground sloth, Acratoc- Monophyllus nus odontrigonus (including Anthony’s A. plethedon X major), is known from numerous poorly Brachyphylla documented cave excavations in north- cavernarum XXX western Puerto Rico. No single sloth or pre- Artibeus cisely defined ‘sloth stratum’ has been ra- jamaicensis X diometrically dated. However, the Stenoderma has not been found in superficial contexts, rufum XX nor has it been found in archaeological de- Phyllonycteris major XX posits that are otherwise common in caves Eptesicus of the Barahona district and elsewhere. An- fuscus XXX thony recovered “one or two fragments of RODENTIA the sloth” from excavations in Cueva Clara Elasmodontomys (field notes in the Department of Mammal- obliquus XXX Xogy, American Museum of Natural His- Heteropsomys tory, 1916); although the stratigraphy of insulans XXX these finds cannot be reconstructed, it is CC = Cueva Clara, CP = Cueva del Perro, CV = unlikely that the specimens postdate the Cueva de la Vaca, CPZ = Cueva del Perezoso. age of >34,650 rcyrbp reported here for the −30 cm stratum. Acratocnus was present in lane, 1999), but in no case is the West In- the Cueva de la Vaca excavation at 33,670 ± dian mammal record currently adequate 370 rcyrbp. No date is available for the for statistical treatment. Cueva Del Perezoso specimens, but Acra- Several authors have attempted to ‘shoe- tocnus is absent from the Cueva Del Perro horn’ West Indian extinctions into a single ‘Layer B’ at 5410 ± 80 rcyrbp, despite the (or at least preeminent) interval, based on presence of Elasmodontomys. The evidence global or continent-wide events. The most available, though inadequate, is consistent popular events have been terminal Pleis- with the view that Acratocnus survived into tocene climate change and human immi- the late Pleistocene but disappeared from gration. Thus, Pregill and Olson (1981) sug- northwestern Puerto Rico before the mid- gested that the spread of mesic habitat Holocene, without trace of anthropogenic during the Holocene may have led to the contact. extinction of a xeric-adapted Pleistocene fauna, an explanation also adopted by Elasmodontomys Woods (1989a). As a reliable database of radiometric dates has begun to accumulate, Elasmodontomys obliquus was recovered it has become apparent that the late Qua- from Cueva Clara, Cueva del Perezoso, ternary extinction crisis cannot be attrib- Cueva de la Vaca at 33,670 ± 370 rcyrbp, uted to a single phenomenon (Flemming et and Cueva del Perro at 5410 ± 80 rcyrbp. al., 1998). Last occurrence dates for West This large bodied (mean body mass = 13.7 Indian late Quaternary mammals are dis- kg, maximum = 17.6 kg, minimum = 9.4 kg; tributed between at least three intervals based on regression equation from Biknev- (Table 3; Fig. 5): the Sangamonian intergla- icius et al., 1993, using 65% distal femur cial (or its immediate aftermath), the mid- anterior-posterior diameter, n= 33) rodent FOSSIL MAMMALS FROM PUERTO RICAN CAVES 243

FIG. 3. Plan view of Cueva Clara. Map symbols and conventions follow Dasher (1994). has never been found in Amerindian mid- abundant charcoal in the sediment record dens or other anthropogenic associations, of a Puerto Rican lake at ∼ 5300 cal-BP is the even though the smaller rodent Isolobodon, signature of Amerindian colonization but believed to be an Amerindian introduction this remains speculative. It is likely that from Hispaniola (Woods, 1989b), is a com- Elasmodontomys persisted at least up to, and mon Amerindian dietary element and ap- perhaps beyond, Amerindian first contact. pears in middens from caves in the Bara- hona district. Heteropsomys The colonization date of Puerto Rico by Amerindians is uncertain, with the oldest Anthony (1926) described three taxa of secure radiocarbon date being 3010 ± 70 rodents form his Puerto Rican collections; rcyrbp (Rouse, 1992). Burney et al. (1994) Isolobodon portoricensis Allen 1916, which argued that the sudden appearance of was considered by Anthony and all subse- 244 D. A. MCFARLANE

FIG. 4. Plan view of Cueva de la Vaca. Map symbols and conventions follow Dasher (1994). quent workers to be an Amerindian intro- Nesophontes duction; Heteropsomys insulans Anthony 1916, and Homopsomys antillensis Anthony of the Nesophontes are 1917. Anthony noted that the latter two known from Cuba, Hispaniola, the Cayman genera were very closely related, and “it Islands, and Puerto Rico; the latter support- may be that the two forms should be con- ing a single large form, N. edithae. The Cay- sidered congeneric” (Anthony, 1926: p151). man Islands form is reported to co-occur Varona (1974) treated Heteropsomys and Ho- with Rattus (Morgan, 1994), implying a mopsomys as congeneric, without elabora- post AD1500 extinction. In the absence of tion, whereas Woods declared them generi- direct radiocarbon date or a description of a cally distinct (1989b) and then synonymous closed stratigraphic context this should be (1993). I follow Woods (1993) in referring considered probable but conjectural. Owl- all late Pleistocene and Holocene Puerto pellet remains of Nesophontes from superfi- Rican echimyid material to Heteropsomys. cial contexts in Hispaniola were reported Heteropsomys was recovered from Cueva by Miller (1930) to retain tissue and hair, Clara, Cueva de la Vaca, and ‘layer B’ of and have been considered as evidence that Cueva del Perro; the latter demonstrating the animal was extant into the 20th century the survival of the taxon into the mid- (e.g., Woods and Eisenberg, 1989). How- Holocene. The limited postcranial material ever, N. hypomicrus specimens preserving available for Heteropsomys suggests that the tissue and hair collected from Cueva Jurg, animal closely matched Isolobodon in size, Parque Nacional Sierra de Baoruco, Do- and thus the absence of this animal from minican Republic, provided a direct accel- Amerindian associations strongly suggests erator radiocarbon date on bone collagen of a non-anthropogenic extinction. 710 ± 50 rcyrbp, demonstrating that preser- FOSSIL MAMMALS FROM PUERTO RICAN CAVES 245

TABLE 3. Last occurrence dates for West Indian late ate without a site description and strati- Quaternary extinct mammals graphic details. A possible Nesophontes as- sociation with an Amerindian midden has Last dated Genus Island occurrence also been reported from Mona Island (A. Nieves, pers. comm.). Presently, it is pru- a Clidomys Jamaica 174,000 ± 12,000 dent to note the marked contrast between Oryzomys Jamaica AD 1877b* k the superficial occurrence of Nesophontes on Xenothrix Jamaica 1870 ± 50 Hispaniola and the Cayman Islands, and its Undescr. rodent Jamaica 11,260 ± 80l Nesophontes Puerto Rico 5410 ± 80c apparent absence from such contexts on Elasmodontomys Puerto Rico 5410 ± 80c mainland Puerto Rico. Heteropsomys Puerto Rico 5410 ± 80c The West Indian radiometric database is Acratocnus Puerto Rico 33,670 ± 370c currently too coarse to confidently identify Nesophontes Cuba 490 ± 50h* patterns in the vertebrate extinction record Cuba 6250 ± 50h* that have pan-West Indian utility. It should Nesophontes Hispaniola 590 ± 50h* be noted that many of the larger-bodied f Rhizoplagiodonita Hispaniola 3755 ± 175 taxa that have been uncritically assumed to Antillothrix Hispaniola 3850 ± 135j l be victims of Amerindian lack any Brotomys Hispaniola 430 ± 60 * such demonstrable association. Amblyrhiza, Isolobodon Hispaniola 710 ± 50l* Capromys Grand Cayman 375 ± 60g the 100-200 kg giant rodent of the Anguilla Amblyrhiza Anguilla 91,000 ± 9800d Bank and its somewhat smaller sister taxon Megalomys St. Lucia AD 1849I* Clidomys of Jamaica, apparently disap- Megalomys Martinique AD 1902b* peared a hundred millennia before human Megalomys Barbuda 750 ± 50e* first-contact (McFarlane et al., 1998a; Megalomys Antigua AD 1200-1260e 1998b). The ∼14 kg rodent Elasmodontomys *signifies direct date on bone collagen, or live- of Puerto Rico provides an especially inter- caught museum specimen. Other dates by strati- esting case, because the ‘last-occurrence’ graphic association. aMcFarlane et al., 1998b; date of 5410 rcybp overlaps (within statis- bMacPhee and Flemming, 1999; cThis paper; dMcFar- tical confidence limits) the ∼5300 rcybp date lane et al., 1998a; eMcFarlane and Flemming, unpub- lished data; fWoods, 1989a; gMorgan, 1994; hMacPhee (Burney et al., 1994) for the appearance of et al., 1999; iAllen, 1942; jRimoli, 1977; kMacPhee, potentially anthropogenic charcoal on the 1996; 1unpublished data, this author. island. Elasmodontomys has not appeared in an archaeological context, even though the vation of organic material is not a reliable Taino are believed to have introduced the indicator of very recent age (MacPhee et al., smaller rodent Isolobodon from Hispaniola 1999). These authors provide a last- and remains of the latter species are com- occurrence date of 680 ± 50 rcybp for N. mon in Puerto Rican midden deposits. paramicrus on Hispaniola, consistent with a The insectivore Nesophontes paramicrus rapid extinction following the circa AD1500 may have survived on Hispaniola into the introduction of Rattus rattus to the island. early historic era, but no comparable evi- Nesophontes has not been recovered from dence of survival beyond the mid- superficial contexts on Puerto Rico. The Holocene has been found for N. edithae on last-occurrence date for a tightly con- mainland Puerto Rico. A similar problem strained stratum containing N. edithae is exists for the Lesser Antillean Megalomys, currently 5410 ± 80 rcyrbp from ‘layer B’ of ‘giant rice rats’ which appear to have dis- Cueva Perro. A single Nesophontes femoral appeared very rapidly after European first- fragment was recovered from the overly- contact on Barbuda, but which persisted on ing, more sparsely fossiliferous ‘layer A.’ St. Lucia into the late 19th century and on Whether this specimen is substantially Martinique into the early 20th century younger that the ‘layer B’ specimens cannot (Flemming and McFarlane, 1997). These be answered on the basis of the available varied extinction chronologies show no ob- material. Morgan and Woods (1986) re- vious correlation with , ported Nesophontes from an Amerindian the introduction of feral mongooses, or site on Vieques, but this is difficult to evalu- Amerindian exploitation. Given that 5 of 246 D. A. MCFARLANE

FIG. 5. Distribution of West Indian mammal ‘last occurrence’ dates. the 8 global, genus-level rodent extinctions the laboratory. Joe Troester provided early of the historic era have impacted West In- encouragement and assistance. Anahi and dian taxa (data from MacPhee and Flem- Abel Vale (Fundación de Investigaciones ming, 1999), the difference of perhaps four Espeleológicas del Karso Puertorriqueño) centuries between different Megalomys spe- collected the high-precision GPS data for cies extinctions in the Lesser Antilles, and the sites. Clare Flemming critiqued an early perhaps four millennia between different version of the manuscript and contributed Nesophontes species extinctions in the systematic advice. , demands explanation. The extinction of the entire suite of greater An- tillean ground sloth genera is currently be- LITERATURE CITED yond explanation, due to the lack of radio- Allen, G. M. 1942. Extinct and vanishing mammals of metric dates. The solutions to these the Western Hemisphere, with the marine species problems can only derive from the accumu- of all the oceans. Amer. Comm. for Int. Wild Life lation of a dense database of radiometric Protection, Special Pub. No.11. New York. 620 p. dates, with careful attention to strati- Allen, J. A. 1916. An extinct octodont from the island of Porto Rico, West Indies. Annals New York Acad. graphic control and the problems of tapho- Sci. 27:17-22. nomy in caves. Andrews, C. W. 1909. On the fauna of Christmas Is- land. Proc. Zool. Soc. Lond., 1909, part 1:101-103. Acknowledgements.—Fieldwork was con- Anthony, H. E. 1916. Preliminary report on fossil ducted under permit # 96-154 issued by the mammals from Porto Rico, with descriptions of a Department of Natural Resources, Puerto new genus of ground sloth and two new genera of hystricomorph rodents. Annals New York Acad. Rico, and funded in part by the W. M. Keck Sci. 27: 193-203. Science Center of The Claremont Colleges. Anthony, H. E. 1916. Preliminary diagnosis of an ap- Caron Bush, Andrea Gessford, and Kirstin parently new family of insectivores. Bulletin Amer. Mewaldt assisted in the excavations and in . Nat. Hist. 35:725-728. FOSSIL MAMMALS FROM PUERTO RICAN CAVES 247

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