AMERICANt Norntates

AMERICANt MUSEUM

NorntatesPUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORYCENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024

Number 3278, 11 pp., 3 figures, 1 table November 12, 1999

Redetermination of Holotype ofIsolobodon portoricensis

(Rodentia, Capromyidae), with Notes on RecentMammalian Extinctions in Puerto Rico*

CLARE FLEMMING1 AND R.D.E. MAcPHEE2

ABSTRACT

While examining material of the Antillean capromyid Isolobodon portoricensis, we discov-ered that the specimen designated by Allen (1916) as the holotype of this species has beenconsistently misidentified in the literature (e.g., Lawrence, 1993). In this paper we correct thiserror and make some observations on Allen's (1916) designated type series in the AMNH-Mcollections. Additionally, we analyze information bearing on the extinction date of I. porto-ricensis and its purported utilization by humans as a food resource.

INTRODUCTIONAs measured by the number of specimens

in collections, the Puerto Rican hutia Iso-lobodon ranks as one of the best knownmembers of the extinct Quaternary mammalfauna of the West Indies. The systematichistory of this taxon and its contents issomewhat confusing. Three species havebeen named: portoricensis, montanus, and

levir (the last two originally placed in dif-ferent monotypic genera by Miller [1922]).Reynolds et al. (1953) doubted the validityof levir, and it now seems to be generallyaccepted that this nomen is a synonym ofportoricensis (Woods, 1993). Samples as-signed to montanus and portoricensis alsodiffer very little when ontogenetic and clinalvariation is taken into account (Reynolds etal., 1953; Woods, 1993), but most authors

* We respectfully dedicate this paper to the memory of our friend and colleague, Karl F Koopman (1922-1997).' Supervisor, Volunteer Services; Research Associate, Division of Vertebrate Zoology; American Museum of Nat-

ural History.2 Curator, Division of Vertebrate Zoology, American Museum of Natural History.

Copyright C) American Museum of Natural History 1999 ISSN 0003-0082 / Price $1.80

AMERICAN MUSEUM NOVITATES

continue to separate the two species. Varona(1974) followed Mohr (1939) in consideringIsolobodon a subgenus of Plagiodontia.However, all Isolobodon can be readily dis-tinguished from other capromyids, includingPlagiodontia, by derived features of cheek-tooth cementum, warranting a genus-leveldistinction (Woods, 1989a; MacPhee andIturralde-Vinent, 1995). As Isolobodonmontanus has been recognized from Hispa-niolan localities only, referral of all PuertoRican specimens discussed here to L por-toricensis is not controversial.

ACKNOWLEDGMENTS

We thank Craig Chesek (AMNH) for hisskillful photography and Patricia J. Wynnefor her interesting rendering of Isolobodonportoricensis "in the flesh." We also thankRichard Tedford (AMNH) for permission tosubmit specimens in his care to radiocarbonanalyses, and Anibal Rodriguez (AMNH) forinformative discussions on the diet of thefirst Puerto Ricans.

ABBREVIATIONS

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

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

AMS 14C accelerator mass spectrometry ra-diocarbon dating

Beta Beta Analytic, Coral Gables, FL(radiocarbon dating lab)

cal-AD solar years AD (used with calibrateddates)

rcyrbp radiocarbon years before present(radiometric datum AD 1950)

yrbp years before present

REDETERMINATION OF HOLOTYPEOF ISOLOBODON PORTORICENSIS

The first scientific collection of the PuertoRican hutia was made in 1915 at Cueva dela Ceiba by the anthropologist Franz Boas,who sent his faunal collection to J. A. Allenwith the notation that

[t]he remains were found in a heavy deposit of ashesin a cave in the Jobo district, between Utuado andArecibo. In the same deposit was the burial of a child.A very large number of shells of crabs and of variouskinds of snails were found. The deposit was undoubt-

edly artificial. I do not believe that it was purely anaccumulation of kitchen refuse. It seems more likelythat it was made for some other purpose. There is noindication of post-Columbian disturbance of the de-posit, but I do not presume that it is more than a fewhundred years old (F Boas, in litt., cited by Allen,1916: 18).

Allen (1916) immediately recognized thatprominent among the ash-encrusted bonessent to him by Boas were the remains of apreviously unknown capromyid-the firstextinct mammal species to be discovered inPuerto Rico. Boas' material included morethan a dozen skulls and large cranial frag-ments. Allen selected one of the more com-plete specimens among these remains as theportoricensis holotype (fig. 1), describing itas follows (Allen, 1916: 19 and pl. 1, figs.3-5):

Isolobodon portoricensis gen. et sp. nov.Type, No. 38409a, from the Cuerva de la Seiba [i.e.,Cueva de la Ceiba], near Utuado, Porto Rico;coll[ector], Dr. Franz Boas. The type skull has thenasals and entire upper dentition complete, but lackspart of one zygoma and the braincase posterior to thefronto-parietal suture.

Allen (1916: 19) also made a critical com-parison to other material in the collectionfrom the type locality: "type skull [is] fullyadult but smaller and evidently younger than[paratype] No. 38409b."The fate of the holotype and type collec-

tion of Isolobodon portoricensis from thispoint onward is obscure. Although the taxonwas certainly not forgotten (e.g., Miller,1929a), Goodwin (1953) failed to include theholotype ofportoricensis in his catalog of theAMNH-M type collection. Lawrence's(1993: 149) later catalog rectified this omis-sion. However, her account of the holotypeand its condition is at variance with Allen's:in her paper, the accession number of theportoricensis holotype is listed as AMNH-M38409, rather than as 38409a, and the con-dition report reads "Cranium partial (nasals,occiput, basiocciput, incisors, both P4, Mls,and left M2, M3 missing)." One might as-sume that Lawrence's description differsfrom that of Allen because of postaccessiondamage to the holotype, but this is not thecase: Allen's (1916) holotype specimen ex-ists in the same state as when he describedit, but to the confusion of subsequent inves-tigators, it was placed in the general AMNH-

NO. 32782

1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE

Fig. 1. Isolobodon portoricensis holotype cranium: AMNH-M 38409a (showing suffix as correctedby authors). Left, dorsal view, showing premortem damage (arrows). Right, closeup of frontal bonesof same specimen. Punctures and cracks are still filled with gray ash (possibly from cooking fires) fromthe deposit in which the specimen was found (see text).

M collections bearing another accession suf-fix (38409b).

It is easy to determine that the specimendescribed by Lawrence is one of the paratypeskulls, as it is illustrated by Allen on thesame plate as the true holotype (Allen, 1916:pl. 1, figs. 6-8; our fig. 2). Evidently, sub-stitution in the collections occurred when thenumbers were physically written on the spec-

imens, whenever that took place; it had cer-tainly happened at or before the time thatH. E. Anthony (1918) had several Isolobodonspecimens photographed for his monographon Puerto Rican mammals. Three skulls hav-ing similar accession numbers (i.e., AMNH-M 38409a, 38409b, and 38409c) are depictedon Anthony's (1918) plate 62. Of these, hisAMNH-M 38409a, listed as the type in the

%--t: i

. ,P I;}

I,

I.

Fig. 2. Isolobodon portoricensis paratype cranium: AMNH-M 38409b (showing suffix as correctedby authors). Left, dorsal view, showing possible premortem damage (arrows). Right, closeup of frontalbones of same specimen. This specimen has been consistently misidentified as the holotype of thisspecies since publication of Anthony's (1918) monograph on Puerto Rican land mammals.

3


figure caption, actually conforms to Allen'sparatype b. Anthony identified another spec-imen as AMNH-M 38409b, but this corre-

sponds to a third member of Allen's paratypeseries. The true holotype is in fact illustratedin Anthony's monograph, but as AMNH-M38409c.Some additional mixups must have oc-

curred, because Allen's paratype b was even-

tually inscribed with the number 38409a andplaced in the type collection; the real holo-type was deposited with the remaining para-types. Anthony's confusion of specimens re-

quired that he use language different fromAllen's when redescribing the holotype (e.g.,Anthony, 1918, 1926), which is the descrip-tion on which Lawrence (1993) relied.

These errors have now been repaired, andthe proper holotype is now installed in theAMNH-M type collection. We correct therelevant entry in the Lawrence (1993) cata-log to read:Isolobodon portoricensis J. A. AllenAnn. New York Acad. Sci. 27: 19, 1916.HOLOTYPE: 38409a. Cranium only.LOCALITY: Puerto Rico; Jobo; Cueva de la

Ceiba, near Utuado.COLLECTOR: F Boas. Original number un-

known.CONDITION: Cranium partial (nasals and

entire upper dentition complete).

DISTRIBUTION AND CULTURALUTILIZATION OF ISOLOBODON

Remains of L portoricensis have been dis-covered at late Holocene occupation sites onmany islands in the northeastern Caribbean(La Gonave, Ile de la Tortue, Hispaniola,Mona, Puerto Rico, Vieques, St. Thomas, St.Croix). No other endemic West Indian rodentspecies enjoyed such an extensive distribu-tion. Indeed, on some islands (e.g., VirginIslands), hutias have been recovered onlywithin human occupation horizons, whichstrongly suggests they could not have beenendemic and had to have been introduced byhumans (Miller, 1918; Woods, 1996).3 For

3Isolobodon may have evolved originally in Hispan-iola, as this is the only context in which it has beenfound in nonanthropogenic settings (Woods, 1996; butsee Anthony, 1918). This point could be settled con-

clusively by finding and directly dating hutia materialthat is truly old (i.e., > 10,000 yrbp), but this projecthas yet to be undertaken.

this reason, the question has been raised fre-quently whether Puerto Rican hutias werekept in a state of captivity or even quasi-domestication by Amerindians (cf. J. A. Al-len, 1916; Anthony, 1918; Miller, 1916,1918; G. M. Allen, 1942; Woods, 1996).

Larger land mammals (various species ofsloths and other rodents) also existed in His-paniola and Puerto Rico during the late Qua-ternary. Oddly, none of these has turned upin any undisturbed archeological contexts; bycontrast, manatees (Trichechus manatus) andeven monk seals (Monachus tropicalis) havebeen found in middens and other coastal sites(Rainey, 1940; personal obs.). One explana-tion for this might be that larger land mam-mals disappeared so rapidly after human col-onization of the Greater Antilles in the earlyto mid-Holocene that their remains have sim-ply not yet been found in archeological con-texts (Martin and Steadman, 1999; but seeBurney et al., 1994; Flenuming et al., 1998).However, explanations of extinction that donot involve overhunting also exist (MacPheeand Marx, 1997).

Although Allen (1916) made reference tocultural modification of Isolobodon bones re-covered at the type locality, his remarks arevery brief and need some amplification to putthem into a useful perspective. Allen's(1916) most perceptive remark was that allskulls recovered from the type locality werefound broken: "... none of them [is] quitecomplete, the occipital and parietal regionsand the nasals being usually lacking, whilemany consist of only the middle portion ofthe skull." This last type of breakage patternis arguably not natural, and suggests humanpreparation to get at the brain, rhinarium,neck musculature, and other soft tissues.

Allen (1916: 17) did not provide a faunalanalysis of the Cueva de la Ceiba material,and his paper contains little quantitative in-formation. It is evident that remains of Iso-lobodon constituted most of the faunal sam-ple, although a few bird bones ("a pigeonand a parrot, the latter probably referable tothe genus Amazona") were also recovered.Remarkably, the Isolobodon material con-sists mostly of hemimandibles (150) andskulls (20), or 43% of the "nearly 400 piec-

NO. 32784


es" identified by Allen to element.4 Postcra-nial bones are thus proportionately underrep-resented, but the reason for this bias is notknown. Bias resulting from butchering tech-niques seems unlikely since the animal'sbody size was relatively small (see below).Allen (1916) noted that "several hundredfragments of little or no scientific value"were also collected-possibly indicating thatpostcranials were broken up or cracked openduring preparation or consumption.A few skulls display unambiguous evi-

dence of human manipulation. On the holo-type skull (AMNH-M 38409a) there is aclear indication of premortem (or immediatepostmortem) damage in the form of a circu-lar, depressed fracture area on the right fron-tal, approximately 5 mm in diameter (fig. 1).The center of the impact site bears a smallpuncture wound. The frontals are incised bya linear cut mark which appears to terminatein or just before the puncture.

In mammals, a sandwich of dense connec-tive tissues, consisting of scalp, periostea,and dura mater, sheathes the tabular bonescomprising the cranial vault. A forceful blowdelivered to the vault of a living individualor intact cadaver will result in radial fractur-ing around the site, but the broken pieces ofskull will tend to remain more or less inplace thanks to the surrounding dense con-nective tissues. If the head is left undis-turbed, the pieces will likely retain their rel-ative positions after soft tissues disappear, al-lowing determination of the time of injury(Brothwell, 1965). Conversely, in skulls bro-ken long after death and decay, there is noth-ing to keep small fragments in their properrelative positions so they tend to simply fallaway. In the portoricensis holotype, the po-sition of fractured pieces at the impact site isconsistent with premortem injury.We infer from this that the depressed frac-

ture and perforation in AMNH-M 38409awere caused by an implement that terminatedin a sharp point or spike. Both cut mark and

4Allen (1916) listed the following elements of L por-toricensis as coming from the same collection as theholotype skull: 20 skulls, 150 mandibular rami, 15 scap-ulae, 1 "clavicle" [= juvenile radius], 15 humeri, 30ulnae, 10 radii, 25 femora, 40 tibiae, 5 fibulae, 1 sacrum,50 ribs, 50 innominate bones, and "several hundredfragments."

puncture wound may have been caused bythe instrument being dragged across the cra-nial roof with some force (perhaps whileskinning the animal, or stabbing it to kill it).Limestone pebbles are unlikely to cause thiskind of damage under any circumstances,whether natural or anthropogenic. A similarlinear cut mark is seen on the frontals of an-other paratype skull, AMNH-M 38409b (fig.2). The feature begins just above the edge ofthe right orbit and slices obliquely over tothe rim of the left infraorbital foramen.

Mandibles also show some signs of ma-nipulation. Allen (1916: 18) stated that "inmany [of the jaws] the condylar portion isdefective or wholly lacking," although wesaw nothing on any jaw that could be con-sidered an unambiguous cut mark. If the con-dyles were removed by human agency, theywere evidently snapped off rather than cutoff-not a difficult undertaking given thegracility of this part of the mandible in theserodents.Of the few long bones represented, none

shows obvious signs of butchering on shaftsor articular ends. A few specimens in the col-lection are deeply calcined, suggesting thathutias were cooked, at least on occasion.Capromyids are relatively large rodents andtherefore a potentially significant source ofmeat (cf. Anthony [1920] concerning hutiahunting in 20th century Jamaica). Allen(1916) estimated the size of Isolobodon por-toricensis to have been roughly comparableto that of Desmarest's hutia (Capromys pi-lorides), an extant Cuban capromyid (adultweight, - 2.6 kg; Silva and Downing, 1995).This compares favorably to guinea pigs (Ca-via, 0.25-0.75 kg) and agoutis (Dasyprocta,2.0-3.8 kg), both of which were introducedto various West Indian islands by Amerin-dians (Miller, 1930), although later introduc-tions are known to have occurred as well(DeVos et al., 1956). Because of its substan-tial size the Puerto Rican hutia would havebeen an animal worth hunting or transport-ing, especially in smaller island contextswhere few or no endemic mammals otherthan bats would have been available. On afinal note, it is worth recording that the Cue-va de la Ceiba sample included a tibia witha healed fracture and a mandible with an ab-scessed molar. This is not an unusual amount

5


Fig. 3. Reconstruction of Isolobodon portoricensis. The cord around the animal's hind leg need notnecessarily imply domestication as such. For example, in reference to the anthropogenically influenceddistribution of I. portoricensis, animals would have to have been caught and kept alive before beingtransported to other islands, whether or not they were kept "in captivity" thereafter.

of pathology to encounter in a natural pop-ulation, of course, but it is commonplace thatinjuries such as these are less likely to bemortal in animals that are kept (and caredfor) in captivity.

It must be admitted on this evidence thatit is not possible to conclude that Isolobodonwas husbanded (as opposed to hunted) inPuerto Rico. Nor does there appear to be anyindependent archeological evidence that canbe recruited to support this notion. If hutiaswere actually kept, they would have to havebeen housed in structures that would not al-low their escape. Quadrilateral areas definedby borders of piled or erected stones (usuallydescribed as juegos de bola or ball courtseven though there is no definitive evidencethat most of them were used as such) havebeen identified at a number of Puerto Ricanarcheological sites (e.g., Rainey, 1940; Ma-son, 1941). However, the areas enclosed bythese features are usually quite large, and ittherefore seems rather unlikely that theycould have functioned as pens or hutches.Wing (1991) reported a high incidence of

dental mutilation (evulsion of fourth premo-lars) in dogs at Sorce, a Saladoid site (-2000-1500 yrbp) on Vieques Island, east ofPuerto Rico. She thinks the mutilation mayhave been practiced in order to insert a muz-zle of some sort, so as to prevent the dogsfrom preying on hutias and ground-nestingbirds that humans wanted themselves.As to the matter of the unusually wide dis-

tribution of hutias, perhaps the best inference

is that Isolobodon individuals were occasion-ally trapped alive by Amerindians, who thenused them to stock any nearby islands lack-ing this convenient resource (cf. similar uti-lization of Dasyprocta in aboriginal West In-dies [Allen, 1942]; cf. also rabbits and hares(Oryctolagus and Lepus) on islands else-where [Nowak, 1999]). This scenario wouldexplain the distribution and frequent utiliza-tion of Puerto Rican hutias by humans with-out requiring that the latter husbanded themto any significant degree (see fig. 3).

EXTINCTION OF ISOLOBODONPORTORICENSIS

EARLY Loss OR LATE SURVIVAL?

Whatever its relationship with humans, thePuerto Rican hutia became extinct neverthe-less, as did two dozen other species of en-demic Antillean land mammals (MacPheeand Flemming, 1999). Indeed, from one per-spective Puerto Rico was hit the hardest ofall the larger islands of the West Indies, as itlost 100% of its endemic nonvolant mam-mals.

In Anthony's (1926: 146) view, I. porto-ricensis was likely to have been "the lastmammal to become extinct on Porto Rico,"although he cited no decisive evidence forthis beyond its occurrence in human occu-pation sites. It has frequently been assumedthat I. portoricensis survived into the early16th century or later, perhaps until the intro-duction of the mongoose in the late 19th cen-

6 NO. 3278


tury (Hoagland et al., 1989; Woods et al.,1985). Recently, Woods (1993: 804) hasgone further and stated that "this species sur-vived in Hispaniola and Puerto Rico until thelast few decades, and may still survive incertain remote areas [such as] La Tortue is-land off the north coast of Haiti." However,as there has been no net increase in the evi-dence supporting late survival (see below),the question of when and under what circum-stances the Puerto Rican hutia disappearedremains open.

The only contemporary reference to hutia-like mammals living in Hispaniola occurs inthe Historia general y natural de las Indiasof Oviedo y Valdes (1535). According toMiller (1929b), the relevant passage in theHistoria may have referred with about equalprobability to (still extant) Plagiodontia as to(extinct) Isolobodon. Although Oviedo yValdes (1535) made reference to the gusta-tory excellence of several of Hispaniola's en-demic rodents (identified as hutia, cori, andmuhoy), his text does not state or even implythat Amerindians kept any of them in con-finement. While his report may be said toconfirm the survival of some native Hispa-niolan species into the early part of the 16thcentury, its value is otherwise limited.

Specimen-based evidence for late disap-pearance of the Puerto Rican hutia is equallylimited, and possibly contradictory. Thereare a few reported associations with intro-duced Rattus rattus (see below), but this es-tablishes only that the "last occurrence" ofIsolobodon portoricensis evidently occurredsubsequent to AD 1500. Radiocarbon datingof critical samples can potentially providemuch finer resolution of extinction times (cf.Steadman et al., 1991). Using this approach,MacPhee et al. (1999) tried to determine a"last occurrence" date for the Antillean is-land-shrew Nesophontes, thought to havesurvived into the 20th century. The latest di-rect date on Nesophontes material retrievedin the course of that investigation was 590 ±50 rcyrbp (cal-AD 1295-1430, with 2-sigmaenvelope), based on a sample from a Cubancave site. All other dated material was older,even though some specimens were reportedas having been found in association withRattus. Radiometric dating provides an in-dependent test of age which, in this instance,

was incompatible with the hypothesis of veryrecent survival of the target taxon. When anadequate sequence of independent dates isavailable, it may be possible to estimate thestatistical confidence limits around an in-ferred extinction date (McFarlane, 1999).The next section discusses our effort to beginto place the "last occurrence" of I. portori-censis on such a basis.

METHODS

Isolobodon specimens selected for datingare from collections made at two localities-Cueva de la Ceiba, described above, and anunnamed cave on Hacienda Jobo near Utu-ado, where H. E. Anthony worked briefly in1916. The Hacienda Jobo specimen selectedfor dating (a mandible) comes from a box ofuncatalogued AMNH-VP specimens thatalso included the bones of the Puerto Ricansloth (Acratocnus odontrigonus) and anotherextinct, endemic rodent (Elasmodontomysobliquus). Judging from paper slips in thebox and Anthony's fieldnotes (Anthony, MS),this collection probably came from the "No.2 cave" on Hacienda Jobo, which he exca-vated during the period June 30-July 2,1916.Specimens selected were submitted for ac-

celerator mass spectroscopy (AMS) dating(table 1). The '3C/'2C ratios as reported bythe lab are normal for recent bone and sug-gest that no contamination has taken place(Geyh and Schleicher, 1990). The '3C-ad-justed 14C dates for these localities (1120 ±50 and 620 ± 60 rcyrbp, respectively) nei-ther prove nor disprove that Puerto Rican hu-tias were still extant at the beginning of Eu-ropean entry into the West Indies. However,the 2-sigma calibrated age range (cal-AD1280-1425) for the Hacienda Jobo jaw is notfar removed from that climacteric, and showsthat the extinction of Isolobodon could nothave occurred earlier than the 14th century(cf. Flemming et al., 1998).

In order to test for contemporaneity of ex-tinct species, 0.5 kg of Elasmodontomysbone from the same box as the Isolobodonjaw was submitted for dating. Although thepretreatment appearance of the Isolobodonjaw was identical to that of the Elasmodon-tomys material (i.e., a thin rind of calcium

7


4.W =3 carbonate encased most bones in the collec-o_m tion prior to cleaning), only the Isolobodon

.0 o,a t. r mandible yielded a date. Evidently, collagenc00<v0;^ o vhad been stripped from the ElasmodontomysmCD=00 N t | bones after deposition, probably as a result

- Ch|| t0 .°of continuous immersion in water within the0 ¢ cave.

s~~~~.0~~ ~%. £ e ~~~~~~~~DISCUSSION

.a oeCD Before discussing dates for IsolobodonaNX> t,4=and their significance, we would like to

XA C\8 .= Oo > < sound a cautionary note concerning the im-portance of dating target taxa directly (and

^Ub X t not relying on referential dates based on 14CX̂=< dating of other taxa supposedly from the

same layer or context). As noted above, theo Z Isolobodon fossils from Hacienda Jobo were

in t z) + a^>vasencrusted and stained as material of otherwo o a,.=;io o : N " endemic taxa. If we had chosen to date only+0 O

Xa -2 CZXc.= > so < <° .= Elasmodontomys remains, we might havet> CZ X Q Q 2 t0+ X jumped to the conclusion that all fossils from

E> = t,4 = this site were undatable and possibly verycov old. Conversely, if we had attempted to date

Isolobodon only, we might have felt justifiedx ; N t et,< in using the resulting date to establish "last

* I occurrence" times for Acratocnus and Elas-modontomys as well. Neither conclusion iswarranted, especially in view of the fact thatour attempt to date the last-named taxon di-

u.z) o £ = > rectly was unsuccessful. When these other0 t+1 +1 Puerto Rican land mammals became extinct

0 o00 is completely unresolved; it need not have0 Y ._been as late as ca. AD 1500 (Flemming et al.,

riS S w 1998). Anthony's (Ms) sketchy fieldnotes are0 041° 8 of little assistance in this regard, since no

stratigraphic data are provided concerningX .> ° the recovery points of individual specimens.

There are few dates to which those re-- ported in table 1 can be usefully compared.

,So > Woods (1989b) reported a number of datesCZ=,X for faunas from cave sites in Hispaniola,

- ° oPDmany of which included Isolobodon. How-ever, as he was not primarily interested incollecting evidence of "last occurrences" of

..<004 < = < Q now-extinct taxa, most of his dates are forQo CZ levels well beneath the surface. All are much.°oX; older than the ones reported here.

00 Z, 'IO0,U 0^ t= e Somewhat more pertinent are dates re-

¢z 64 4o t > o . 0 cently reported by MacPhee et al. (1999) forC6 Monte Culo de Maco, a rock shelter whichX 1) um'0S m > U < yielded owl-pellet material (collected for G.

< < c u S. Miller) containing Nesophontes, unidenti-

8 NO. 3278


fied mammal hair, Rattus bones, and the re-mains of several small extinct mammals, in-cluding Isolobodon (for taxon list see Miller,1930). Despite Miller's (1930) belief that thismaterial was very recent, specimens of theinsectivoran Nesophontes from this siteyielded a '3C-adjusted 14C age of 680 ± 50rcyrbp, while Old World rats from the samesite spanned a range of 480 ± 60 to 310 ±40 rcyrbp (MacPhee et al., 1999). The dis-parity between the insectivore and rat datesimplies that the owl-pellet remains at MonteCulo de Maco are not penecontemporaneousbut were instead built up over several cen-turies. The message here is that one shouldalways be cautious in accepting the validityof claimed associations between endemicsand exotics in such contexts, and always at-tempt to date the target taxon directly.The only conclusion that seems warranted

from the chronometric evidence available atpresent is that, contra Miller (1929a, 1929b)and Woods et al. (1985), there is still no ev-idence to support the view that Isolobodonportoricensis survived long into the period ofEuropean occupation. The documentary in-dications are that these rodents were to beseen only very occasionally even in the ear-liest days of European colonization, and it istherefore entirely plausible that they were inprecipitous decline by or even before AD1500.

Finally, we note that it is somewhat puz-zling that Puerto Rican hutias became extinctif in fact they were kept by pre-ColumbianAmerindians. Human management (as op-posed to overkill) of food animals is usuallyconceived to be a hedge against extinction.It is clear that much remains to be learnedabout the cause/effect relationships that re-sulted in the loss of so many island mammalsduring recent historic times (MacPhee andMarx, 1997; MacPhee and Flemming, 1999).

REFERENCES

Allen, J. A.1916. An extinct octodont from the island of

Porto Rico, West Indies. Ann. Acad.New York Sci. 27: 17-22.

Allen, G. M.1942. Extinct and vanishing mammals of the

Western Hemisphere with the marinespecies of all the oceans. Am. Com-

mittee Int. Wild Life Protection Spec.Pub. 11. New York: Cooper Square.

Anthony, H. E.MS. Fieldnotes of H. E. Anthony, "New

York Academy of Sciences Expedition,Porto Rico, 1916, 31 May to 30 July"[title on first page]. Unpublished type-script, Mammalogy Departmental Li-brary & Archives, American Museumof Natural History.

1918. The indigenous land mammals of PortoRico, living and extinct. Mem. Am.Mus. Nat. Hist., n. ser. 2(2): 331-435.

1920. A zoologist in Jamaica. Nat. Hist. 20:156-168

1926. Mammals of Porto Rico, living and ex-tinct-Rodentia and Edentata. Sci.Surv. Porto Rico and Virgin Is. 9(2):97-238.

Brothwell, D. R.1965. Digging up bones: the excavation,

treatment and study of human skeletalremains. London: Br. Mus. (Nat. Hist.).

Burney, D. A., L. P. Burney, and R.D.E. MacPhee1994. Holocene charcoal stratigraphy from

Laguna Tortuguero, Puerto Rico, andthe timing of human arrival on the is-land. J. Archaeol. Sci. 21: 273-281.

DeVos, A., R. H. Manville, and R. G. Van Gelder1956. Introduced mammals and their influ-

ence on biota. Zoologica, New York41: 163-194.

Flemming, C., R.D.E. MacPhee, and D. A.McFarlane

1998. Thrice bitten: late Quaternary mammalextinctions in the continental and in-sular New World. J. Vertebr. Paleontol.18 (Suppl. 3): 42A.

Geyh, M. A., and H. Schleicher1990. Absolute age determination: physical

and chemical dating methods and theirapplication. New York: Springer.

Goodwin, G.1953. Catalogue of the type specimens of Re-

cent mammals in the American Muse-um of Natural History. Bull. Am. Mus.Nat. Hist. 102: 207-412.

Hoagland, D. B., G. R. Horst, and C. W. Kilpa-trick

1989. Biogeography and population biologyof the mongoose in the West Indies. InC. A. Woods (ed.), Biogeography of theWest Indies; past, present, and future:611-633. Gainesville, FL: SandhillCrane.

Lawrence, M. A.1993. Catalog of Recent mammal types in the

9


American Museum of Natural History.Bull. Am. Mus. Nat. Hist. 217: 200 pp.

MacPhee, R.D.E., and C. Flemming1999. Requiem aternum: the last five hundred

years of mammalian species extinc-tions. In R. D. E. MacPhee (ed.), Ex-tinctions in near time: causes, contexts,and consequences: 333-372. NewYork: Kluwer Academic/Plenum.

MacPhee, R.D.E., and M. A. Iturralde-Vinent1995. Origin of the Greater Antillean land

mammal fauna, 1: new Tertiary fossilsfrom Cuba and Puerto Rico. Am. Mus.Novitates 3141: 30 pp.

MacPhee, R.D.E., and P. A. Marx1997. The 40,000-year plague: humans, hy-

perdisease, and first-contact extinc-tions. In S. Goodman and B. Patterson(eds.), Natural change and human im-pact in Madagascar: 169-217. Wash-ington DC: Smithson. Inst. Press.

MacPhee, R.D.E., C. Flemming, and D. P. Lunde1999. "Last occurrence" of the Antillean in-

sectivoran Nesophontes: new radiomet-ric dates and their interpretation. Am.Mus. Novitates 3264: 20 pp.

Martin, P. S.1984. Prehistoric overkill: the global model.

In P. S. Martin and R. G. Klein (eds),Quaternary extinctions, a prehistoricrevolution: 354-403. Tucson: Univ.Arizona Press.

Martin, P. S., and D. W Steadman1999. Prehistoric extinctions on islands and

continents. In R.D.E. MacPhee (ed.),Extinctions in near time: causes, con-texts, and consequences: 17-55. NewYork: Kluwer Academic/Plenum.

Mason, J. A.1941. A large archaeological site at Capa,

Utuado, with notes on other Porto Ricosites visited in 1914-1915. Sci. Surv.Porto Rico and Virgin Is. 18(2): 209-272.

McFarlane, D. A.1999. A comparison of methods for the prob-

abilistic determination of vertebrate ex-tinction chronologies. In R.D.E.MacPhee (ed.), Extinctions in neartime: causes, contexts, and consequen-ces: 95-103. New York: Kluwer Aca-demic/Plenum.

Miller, G. S.1916. Bones of mammals from Indian sites in

Cuba and Santo Domingo. Smithson.Misc. Collect. 66(12): 11 pp.

1918. Mammals and reptiles collected by

Theodor de Booy in the Virgin Islands.Proc. U.S. Natl. Mus. 54: 507-511.

1922. Remains of mammals from caves in theRepublic of Haiti. Smithson. Misc.Collect. 74(3): 8 pp.

1929a. A second collection of mammals fromcaves near St. Michel, Haiti. Ibid.81(9): 30 pp.

1929b. Mammals eaten by Indians, owls, andSpaniards in the coast region of the Do-minican Republic. Ibid. 82(5): 16 pp.

1930. Three small collections of mammalsfrom Hispaniola. Ibid. 82(15): 10 pp.

Mohr, E.1939. Die Baum- und Ferkelratten-Gattungen

Capromys Desmarest (sens. ampl.) undPlagiodontia Cuvier. Mitt. Hambur-gishen Zool. Mus. Inst. 48: 48-118.

Nowak, R. M.1999. Walker's mammals of the world, 6th

ed., 2 vols. Baltimore: Johns HopkinsUniv. Press.

Oviedo y Valdes, G. F. de1535. Historia general y natural de las Indias.

Juam Cromberger: Seville. [trans. anded. by S. A. Stroudemire, Natural His-tory of the West Indies, 1959, Univ.North Carolina Stud. Romance Lang.Lit. 32: 1-140.1

Rainey, F G.1940. Porto Rican archaeology. Sci. Surv.

Porto Rico and Virgin Is. 18(1): 1-208.Reynolds, T. E., K. F Koopman, and E. E. Wil-

liams1953. A cave faunule from western Puerto

Rico with a discussion of the genus Is-olobodon. Breviora 12, 8 pp.

Silva, M., and J. A. Downing1995. CRC handbook of mammalian body

masses. Boca Raton, LA: CRC Press.Steadman, D. W., T W. Stafford, Jr., D. J. Dona-

hue, and A. J. T Jull1991. Chronology of Holocene vertebrate ex-

tinctions in the Galapagos Islands.Quat. Res. 36: 126-133.

Varona, L.1974. Catalogo de los mamfferos vivientes y

extinguidos de las Antillas. La Habana:Acad. Cienc. Cuba.

Wing, E. S.1991. Dog remains from the Sorce' site on

Vieques Island, Puerto Rico. IllinoisState Mus. Sci. Pap. 23: 379-386.

Woods, C. A.1989a. The biogeography of West Indian ro-

dents. In C. A. Woods (ed.), Biogeog-raphy of the West Indies; past, present,

10 NO. 3278


and future: 741-798. Gainesville, FL:Sandhill Crane.

1989b. A new capromyid rodent from Haiti:the origin, evolution, and extinction ofWest Indian rodents and their bearingon the origin of New World hystricog-naths. In C. C. Black and M. R. Daw-son (eds.), Papers on fossil rodents inhonor of Albert Elmer Wood: 58-89.Los Angeles: Nat. Hist. Mus. Los An-geles Cty.

1993. Suborder Hystricognathi. In D. E. Wil-son and D. M. Reeder (eds.), Mammalspecies of the world: a taxonomic andgeographic reference, second edition:

771-806. Washington DC: Smithson.Inst. Press.

1996. The land mammals of Puerto Rico andthe Virgin Islands. In J. C. FigueroaColon (ed.), The scientific survey ofPuerto Rico and the Virgin Islands, an

eighty-year reassessment of the islands'natural history. Ann. New York Acad.Sci. 776: 131-148.

Woods, C. A., J. A. Ottenwalder, and WL.R. Oliver1985. Lost mammals of the Greater Antilles:

the summarised findings of a ten weeksfield survey in the Dominican Repub-lic, Haiti and Puerto Rico. Dodo, J.Wildl. Preserv. Trust 22: 23-42.

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