AMERICANt MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3278, 11 pp., 3 figures, 1 table November 12, 1999

Redetermination of Holotype of Isolobodon portoricensis (Rodentia, Capromyidae), with Notes on Recent Mammalian 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 been consistently misidentified in the literature (e.g., Lawrence, 1993). In this paper we correct this error and make some observations on Allen's (1916) designated type series in the AMNH-M collections. Additionally, we analyze information bearing on the extinction date of I. porto- ricensis and its purported utilization by humans as a food resource.

INTRODUCTION levir (the last two originally placed in dif- As measured the number of ferent monotypic genera by Miller [1922]). by specimens Reynolds et al. (1953) doubted the validity in collections, the Puerto Rican Iso- of levir, and it now seems to be generally lobodon ranks as one of the best known accepted that this nomen is a synonym of members of the extinct Quaternary portoricensis (Woods, 1993). Samples as- fauna of the West Indies. The systematic signed to montanus and portoricensis also history of this taxon and its contents is differ very little when ontogenetic and clinal somewhat confusing. Three species have variation is taken into account (Reynolds et been named: portoricensis, montanus, and al., 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 2 AMERICAN MUSEUM NOVITATES NO. 3278 continue to separate the two species. Varona edly artificial. I do not believe that it was purely an accumulation of kitchen refuse. It seems more likely (1974) followed Mohr (1939) in considering that it was made for some other purpose. There is no Isolobodon a subgenus of Plagiodontia. indication of post-Columbian disturbance of the de- However, all Isolobodon can be readily dis- posit, but I do not presume that it is more than a few tinguished from other capromyids, including hundred years old (F Boas, in litt., cited by Allen, Plagiodontia, by derived features of cheek- 1916: 18). tooth cementum, warranting a genus-level Allen (1916) immediately recognized that distinction (Woods, 1989a; MacPhee and prominent among the ash-encrusted bones Iturralde-Vinent, 1995). As Isolobodon sent to him by Boas were the remains of a montanus has been recognized from Hispa- previously unknown capromyid-the first niolan localities only, referral of all Puerto extinct mammal species to be discovered in Rican specimens discussed here to L por- Puerto Rico. Boas' material included more toricensis is not controversial. than a dozen skulls and large cranial frag- ments. Allen selected one of the more com- ACKNOWLEDGMENTS plete specimens among these remains as the We thank Craig Chesek (AMNH) for his portoricensis holotype (fig. 1), describing it skillful photography and Patricia J. Wynne as follows (Allen, 1916: 19 and pl. 1, figs. for her interesting rendering of Isolobodon 3-5): portoricensis "in the flesh." We also thank Isolobodon portoricensis gen. et sp. nov. Richard Tedford (AMNH) for permission to Type, No. 38409a, from the Cuerva de la Seiba [i.e., submit specimens in his care to radiocarbon Cueva de la Ceiba], near Utuado, Porto Rico; coll[ector], Dr. Franz Boas. The type skull has the analyses, and Anibal Rodriguez (AMNH) for nasals and entire upper dentition complete, but lacks informative discussions on the diet of the part of one zygoma and the braincase posterior to the first Puerto Ricans. fronto-parietal suture. Allen (1916: 19) also made a critical com- ABBREVIATIONS parison to other material in the collection AMNH-M American Museum of Natural His- from the type locality: "type skull [is] fully tory, Department of Mammalogy adult but smaller and evidently younger than AMNH-VP American Museum of Natural His- [paratype] No. 38409b." tory, Department of Vertebrate Pa- The fate of the holotype and type collec- leontology AMS 14C accelerator mass spectrometry ra- tion of Isolobodon portoricensis from this diocarbon dating point onward is obscure. Although the taxon Beta Beta Analytic, Coral Gables, FL was certainly not forgotten (e.g., Miller, (radiocarbon dating lab) 1929a), Goodwin (1953) failed to include the cal-AD solar years AD (used with calibrated holotype ofportoricensis in his catalog of the dates) AMNH-M type collection. Lawrence's rcyrbp radiocarbon years before present (1993: 149) later catalog rectified this omis- (radiometric datum AD 1950) sion. However, her account of the holotype yrbp years before present and its condition is at variance with Allen's: in her paper, the accession number of the REDETERMINATION OF HOLOTYPE portoricensis holotype is listed as AMNH-M OF ISOLOBODON PORTORICENSIS 38409, rather than as 38409a, and the con- The first scientific collection of the Puerto dition report reads "Cranium partial (nasals, Rican hutia was made in 1915 at Cueva de occiput, basiocciput, incisors, both P4, Mls, la Ceiba by the anthropologist Franz Boas, and left M2, M3 missing)." One might as- who sent his faunal collection to J. A. Allen sume that Lawrence's description differs with the notation that from that of Allen because of postaccession damage to the holotype, but this is not the [t]he remains were found in a heavy deposit of ashes case: Allen's (1916) holotype specimen ex- in a cave in the Jobo district, between Utuado and Arecibo. In the same deposit was the burial of a child. ists in the same state as when he described A very large number of shells of crabs and of various it, but to the confusion of subsequent inves- kinds of snails were found. The deposit was undoubt- tigators, it was placed in the general AMNH- 1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE 3

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

M collections bearing another accession suf- imens, whenever that took place; it had cer- fix (38409b). tainly happened at or before the time that It is easy to determine that the specimen H. E. Anthony (1918) had several Isolobodon described by Lawrence is one of the paratype specimens photographed for his monograph skulls, as it is illustrated by Allen on the on Puerto Rican . Three skulls hav- same plate as the true holotype (Allen, 1916: ing similar accession numbers (i.e., AMNH- pl. 1, figs. 6-8; our fig. 2). Evidently, sub- M 38409a, 38409b, and 38409c) are depicted stitution in the collections occurred when the on Anthony's (1918) plate 62. Of these, his numbers were physically written on the spec- AMNH-M 38409a, listed as the type in the

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Fig. 2. Isolobodon portoricensis paratype cranium: AMNH-M 38409b (showing suffix as corrected by authors). Left, dorsal view, showing possible premortem damage (arrows). Right, closeup of frontal bones of same specimen. This specimen has been consistently misidentified as the holotype of this species since publication of Anthony's (1918) monograph on Puerto Rican land mammals. 4 AMERICAN MUSEUM NOVITATES NO. 3278 figure caption, actually conforms to Allen's this reason, the question has been raised fre- paratype b. Anthony identified another spec- quently whether Puerto Rican were imen as AMNH-M 38409b, but this corre- kept in a state of captivity or even quasi- sponds to a third member of Allen's paratype domestication by Amerindians (cf. J. A. Al- series. The true holotype is in fact illustrated len, 1916; Anthony, 1918; Miller, 1916, in Anthony's monograph, but as AMNH-M 1918; G. M. Allen, 1942; Woods, 1996). 38409c. Larger land mammals (various species of Some additional mixups must have oc- sloths and other ) also existed in His- curred, because Allen's paratype b was even- paniola and Puerto Rico during the late Qua- tually inscribed with the number 38409a and of has turned up the real holo- ternary. Oddly, none these placed in the type collection; in any undisturbed archeological contexts; by type was deposited with the remaining para- manatees manatus) and types. Anthony's confusion of specimens re- contrast, (Trichechus quired that he use language different from even monk seals (Monachus tropicalis) have Allen's when redescribing the holotype (e.g., been found in middens and other coastal sites Anthony, 1918, 1926), which is the descrip- (Rainey, 1940; personal obs.). One explana- tion on which Lawrence (1993) relied. tion for this might be that larger land mam- These errors have now been repaired, and mals disappeared so rapidly after human col- the proper holotype is now installed in the onization of the Greater Antilles in the early AMNH-M type collection. We correct the to mid-Holocene that their remains have sim- relevant entry in the Lawrence (1993) cata- ply not yet been found in archeological con- log to read: texts (Martin and Steadman, 1999; but see Isolobodon portoricensis J. A. Allen Burney et al., 1994; Flenuming et al., 1998). Ann. New York Acad. Sci. 27: 19, 1916. However, explanations of extinction that do HOLOTYPE: 38409a. Cranium only. not involve overhunting also exist (MacPhee LOCALITY: Puerto Rico; Jobo; Cueva de la and Marx, 1997). Ceiba, near Utuado. Although Allen (1916) made reference to COLLECTOR: F Boas. Original number un- cultural modification of Isolobodon bones re- known. covered at the type locality, his remarks are CONDITION: Cranium partial (nasals and very brief and need some amplification to put entire upper dentition complete). them into a useful perspective. Allen's DISTRIBUTION AND CULTURAL (1916) most perceptive remark was that all UTILIZATION OF ISOLOBODON skulls recovered from the type locality were found broken: none of them [is] quite Remains of L portoricensis have been dis- "... covered at late Holocene occupation sites on complete, the occipital and parietal regions many islands in the northeastern Caribbean and the nasals being usually lacking, while (La Gonave, Ile de la Tortue, Hispaniola, many consist of only the middle portion of Mona, Puerto Rico, Vieques, St. Thomas, St. the skull." This last type of breakage pattern Croix). No other endemic West Indian is arguably not natural, and suggests human species enjoyed such an extensive distribu- preparation to get at the brain, rhinarium, tion. Indeed, on some islands (e.g., Virgin neck musculature, and other soft tissues. Islands), hutias have been recovered only Allen (1916: 17) did not provide a faunal within human occupation horizons, which analysis of the Cueva de la Ceiba material, strongly suggests they could not have been and his paper contains little quantitative in- endemic and had to have been introduced by humans (Miller, 1918; Woods, 1996).3 For formation. It is evident that remains of Iso- lobodon constituted most of the faunal sam-

3Isolobodon may have evolved originally in Hispan- ple, although a few bird bones ("a pigeon iola, as this is the only context in which it has been and a parrot, the latter probably referable to found in nonanthropogenic settings (Woods, 1996; but the genus Amazona") were also recovered. see Anthony, 1918). This point could be settled con- the Isolobodon material con- clusively by finding and directly dating hutia material Remarkably, that is truly old (i.e., > 10,000 yrbp), but this project sists mostly of hemimandibles (150) and has yet to be undertaken. skulls (20), or 43% of the "nearly 400 piec- 1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE 5 es" identified by Allen to element.4 Postcra- puncture wound may have been caused by nial bones are thus proportionately underrep- the instrument being dragged across the cra- resented, but the reason for this bias is not nial roof with some force (perhaps while known. Bias resulting from butchering tech- skinning the , or stabbing it to kill it). niques seems unlikely since the animal's Limestone pebbles are unlikely to cause this body size was relatively small (see below). kind of damage under any circumstances, Allen (1916) noted that "several hundred whether natural or anthropogenic. A similar fragments of little or no scientific value" linear cut mark is seen on the frontals of an- were also collected-possibly indicating that other paratype skull, AMNH-M 38409b (fig. postcranials were broken up or cracked open 2). The feature begins just above the edge of during preparation or consumption. the right orbit and slices obliquely over to A few skulls display unambiguous evi- the rim of the left infraorbital foramen. dence of human manipulation. On the holo- Mandibles also show some signs of ma- type skull (AMNH-M 38409a) there is a nipulation. Allen (1916: 18) stated that "in clear indication of premortem (or immediate many [of the jaws] the condylar portion is postmortem) damage in the form of a circu- defective or wholly lacking," although we lar, depressed fracture area on the right fron- saw nothing on any jaw that could be con- tal, approximately 5 mm in diameter (fig. 1). sidered an unambiguous cut mark. If the con- The center of the impact site bears a small dyles were removed by human agency, they puncture wound. The frontals are incised by were evidently snapped off rather than cut a linear cut mark which appears to terminate off-not a difficult undertaking given the in or just before the puncture. gracility of this part of the mandible in these In mammals, a sandwich of dense connec- rodents. tive tissues, consisting of scalp, periostea, Of the few long bones represented, none and dura mater, sheathes the tabular bones shows obvious signs of butchering on shafts comprising the cranial vault. A forceful blow or articular ends. A few specimens in the col- delivered to the vault of a living individual lection are deeply calcined, suggesting that or intact cadaver will result in radial fractur- hutias were cooked, at least on occasion. ing around the site, but the broken pieces of Capromyids are relatively large rodents and skull will tend to remain more or less in therefore a potentially significant source of place thanks to the surrounding dense con- meat (cf. Anthony [1920] concerning hutia nective tissues. If the head is left undis- hunting in 20th century Jamaica). Allen turbed, the pieces will likely retain their rel- (1916) estimated the size of Isolobodon por- ative positions after soft tissues disappear, al- toricensis to have been roughly comparable lowing determination of the time of injury to that of Desmarest's hutia (Capromys pi- (Brothwell, 1965). Conversely, in skulls bro- lorides), an extant Cuban capromyid (adult ken long after death and decay, there is noth- weight, - 2.6 kg; Silva and Downing, 1995). ing to keep small fragments in their proper This compares favorably to guinea pigs (Ca- relative positions so they tend to simply fall via, 0.25-0.75 kg) and agoutis (Dasyprocta, away. In the portoricensis holotype, the po- 2.0-3.8 kg), both of which were introduced sition of fractured pieces at the impact site is to various West Indian islands by Amerin- consistent with premortem injury. dians (Miller, 1930), although later introduc- We infer from this that the depressed frac- tions are known to have occurred as well ture and perforation in AMNH-M 38409a (DeVos et al., 1956). Because of its substan- were caused by an implement that terminated tial size the Puerto Rican hutia would have in a sharp point or spike. Both cut mark and been an animal worth hunting or transport- ing, especially in smaller island contexts 4Allen (1916) listed the following elements of L por- where few or no endemic mammals other toricensis as coming from the same collection as the than bats would have been available. On a holotype skull: 20 skulls, 150 mandibular rami, 15 scap- final note, it is worth recording that the Cue- ulae, 1 "clavicle" [= juvenile radius], 15 humeri, 30 ulnae, 10 radii, 25 femora, 40 tibiae, 5 fibulae, 1 sacrum, va de la Ceiba sample included a tibia with 50 ribs, 50 innominate bones, and "several hundred a healed fracture and a mandible with an ab- fragments." scessed molar. This is not an unusual amount 6 AMERICAN MUSEUM NOVITATES NO. 3278

Fig. 3. Reconstruction of Isolobodon portoricensis. The cord around the animal's hind leg need not necessarily imply domestication as such. For example, in reference to the anthropogenically influenced distribution of I. portoricensis, would have to have been caught and kept alive before being transported to other islands, whether or not they were kept "in captivity" thereafter. of pathology to encounter in a natural pop- is that Isolobodon individuals were occasion- ulation, of course, but it is commonplace that ally trapped alive by Amerindians, who then injuries such as these are less likely to be used them to stock any nearby islands lack- mortal in animals that are kept (and cared ing this convenient resource (cf. similar uti- for) in captivity. lization of Dasyprocta in aboriginal West In- It must be admitted on this evidence that dies [Allen, 1942]; cf. also rabbits and hares it is not possible to conclude that Isolobodon (Oryctolagus and Lepus) on islands else- was husbanded (as opposed to hunted) in where [Nowak, 1999]). This scenario would Puerto Rico. Nor does there appear to be any explain the distribution and frequent utiliza- independent archeological evidence that can tion of Puerto Rican hutias by humans with- be recruited to support this notion. If hutias out requiring that the latter husbanded them were actually kept, they would have to have to any significant degree (see fig. 3). been housed in structures that would not al- low their escape. Quadrilateral areas defined EXTINCTION OF ISOLOBODON by borders of piled or erected stones (usually PORTORICENSIS described as juegos de bola or ball courts even though there is no definitive evidence EARLY Loss OR LATE SURVIVAL? that most of them were used as such) have Whatever its relationship with humans, the been identified at a number of Puerto Rican Puerto Rican hutia became extinct neverthe- archeological sites (e.g., Rainey, 1940; Ma- less, as did two dozen other species of en- son, 1941). However, the areas enclosed by demic Antillean land mammals (MacPhee these features are usually quite large, and it and Flemming, 1999). Indeed, from one per- therefore seems rather unlikely that they spective Puerto Rico was hit the hardest of could have functioned as pens or hutches. all the larger islands of the West Indies, as it Wing (1991) reported a high incidence of lost 100% of its endemic nonvolant mam- dental mutilation (evulsion of fourth premo- mals. lars) in dogs at Sorce, a Saladoid site (- In Anthony's (1926: 146) view, I. porto- 2000-1500 yrbp) on Vieques Island, east of ricensis was likely to have been "the last Puerto Rico. She thinks the mutilation may mammal to become extinct on Porto Rico," have been practiced in order to insert a muz- although he cited no decisive evidence for zle of some sort, so as to prevent the dogs this beyond its occurrence in human occu- from preying on hutias and ground-nesting pation sites. It has frequently been assumed birds that humans wanted themselves. that I. portoricensis survived into the early As to the matter of the unusually wide dis- 16th century or later, perhaps until the intro- tribution of hutias, perhaps the best inference duction of the mongoose in the late 19th cen- 1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE 7 tury (Hoagland et al., 1989; Woods et al., was incompatible with the hypothesis of very 1985). Recently, Woods (1993: 804) has recent survival of the target taxon. When an gone further and stated that "this species sur- adequate sequence of independent dates is vived in Hispaniola and Puerto Rico until the available, it may be possible to estimate the last few decades, and may still survive in statistical confidence limits around an in- certain remote areas [such as] La Tortue is- ferred extinction date (McFarlane, 1999). land off the north coast of Haiti." However, The next section discusses our effort to begin as there has been no net increase in the evi- to place the "last occurrence" of I. portori- dence supporting late survival (see below), censis on such a basis. the question of when and under what circum- stances the Puerto Rican hutia disappeared METHODS remains open. The only contemporary reference to hutia- Isolobodon specimens selected for dating like mammals living in Hispaniola occurs in are from collections made at two localities- the Historia general y natural de las Indias Cueva de la Ceiba, described above, and an of Oviedo y Valdes (1535). According to unnamed cave on Hacienda Jobo near Utu- Miller (1929b), the relevant passage in the ado, where H. E. Anthony worked briefly in Historia may have referred with about equal 1916. The Hacienda Jobo specimen selected probability to (still extant) Plagiodontia as to for dating (a mandible) comes from a box of (extinct) Isolobodon. Although Oviedo y uncatalogued AMNH-VP specimens that Valdes (1535) made reference to the gusta- also included the bones of the Puerto Rican tory excellence of several of Hispaniola's en- sloth (Acratocnus odontrigonus) and another demic rodents (identified as hutia, cori, and extinct, endemic rodent (Elasmodontomys muhoy), his text does not state or even imply obliquus). Judging from paper slips in the that Amerindians kept any of them in con- box and Anthony's fieldnotes (Anthony, MS), finement. While his report may be said to this collection probably came from the "No. confirm the survival of some native Hispa- 2 cave" on Hacienda Jobo, which he exca- niolan species into the early part of the 16th vated during the period June 30-July 2, century, its value is otherwise limited. 1916. Specimen-based evidence for late disap- Specimens selected were submitted for ac- pearance of the Puerto Rican hutia is equally celerator mass spectroscopy (AMS) dating limited, and possibly contradictory. There (table 1). The '3C/'2C ratios as reported by are a few reported associations with intro- the lab are normal for recent bone and sug- duced Rattus rattus (see below), but this es- gest that no contamination has taken place tablishes only that the "last occurrence" of (Geyh and Schleicher, 1990). The '3C-ad- Isolobodon portoricensis evidently occurred justed 14C dates for these localities (1120 ± subsequent to AD 1500. Radiocarbon dating 50 and 620 ± 60 rcyrbp, respectively) nei- of critical samples can potentially provide ther prove nor disprove that Puerto Rican hu- much finer resolution of extinction times (cf. tias were still extant at the beginning of Eu- Steadman et al., 1991). Using this approach, ropean entry into the West Indies. However, MacPhee et al. (1999) tried to determine a the 2-sigma calibrated age range (cal-AD "last occurrence" date for the Antillean is- 1280-1425) for the Hacienda Jobo jaw is not land-shrew Nesophontes, thought to have far removed from that climacteric, and shows survived into the 20th century. The latest di- that the extinction of Isolobodon could not rect date on Nesophontes material retrieved have occurred earlier than the 14th century in the course of that investigation was 590 ± (cf. Flemming et al., 1998). 50 rcyrbp (cal-AD 1295-1430, with 2-sigma In order to test for contemporaneity of ex- envelope), based on a sample from a Cuban tinct species, 0.5 kg of Elasmodontomys cave site. All other dated material was older, bone from the same box as the Isolobodon even though some specimens were reported jaw was submitted for dating. Although the as having been found in association with pretreatment appearance of the Isolobodon Rattus. Radiometric dating provides an in- jaw was identical to that of the Elasmodon- dependent test of age which, in this instance, tomys material (i.e., a thin rind of calcium 8 AMERICAN MUSEUM NOVITATES NO. 3278

4.W =3 carbonate encased most bones in the collec- _mo tion prior to cleaning), only the Isolobodon .0 o,a t. r mandible yielded a date. Evidently, collagen c00

s~~~~.0%.~~ ~ £ e ~~~~~~~~DISCUSSION .a o eCD Before discussing dates for Isolobodon X>aN t,4=and their significance, we would like to XA 8 .= Oo C\> < sound a cautionary note concerning the im- portance of dating target taxa directly (and ^Ub X t not relying on referential dates based on 14C X^=< dating of other taxa supposedly from the same layer or context). As noted above, the o Z Isolobodon fossils from Hacienda Jobo were in t ^>vasz)encrusted+ a and stained as material of other " wo o a,.= ;i+0o Oo : N endemic taxa. If we had chosen to date only Xa-2 Xc.= > CZ so < <° .= Elasmodontomys remains, we might have t> CZ X Q Q 2 t0+ X jumped to the conclusion that all fossils from E> = t,4 = this site were undatable and possibly very ovc old. Conversely, if we had attempted to date Isolobodon only, we might have felt justified x ; N t et,< in using the resulting date to establish "last * I occurrence" times for Acratocnus and Elas- modontomys as well. Neither conclusion is warranted, especially in view of the fact that our attempt to date the last-named taxon di- u.z) o £ = > rectly was unsuccessful. When these other 0 +1t +1 Puerto Rican land mammals became extinct 0 o00 is completely unresolved; it need not have 0 Y been._ as late as ca. AD 1500 (Flemming et al., fieldnotes are riS S w0 0 1998). Anthony's (Ms) sketchy 41° 8 of little assistance in this regard, since no stratigraphic data are provided concerning X .> ° 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 dates =,XCZ for faunas from cave sites in Hispaniola, - ° manyoPD of which included Isolobodon. How- ever, as he was not primarily interested in collecting evidence of "last occurrences" of ..<004 < = < Q now-extinct taxa, most of his dates are for oQ CZ levels well beneath the surface. All are much .°oX; older than the ones reported here. 00 'IO0 Z,^ t= ,U 0e Somewhat more pertinent are dates re- ¢z o t > o .64 4 0 cently reported by MacPhee et al. (1999) for C6 Monte Culo de Maco, a rock shelter which X1) u'0mS m > U < yielded owl-pellet material (collected for G. < < c u S. Miller) containing Nesophontes, unidenti- 1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE 9 fied mammal hair, Rattus bones, and the re- mittee Int. Wild Life Protection Spec. mains of several small extinct mammals, in- Pub. 11. New York: Cooper Square. cluding Isolobodon (for taxon list see Miller, Anthony, H. E. 1930). Despite Miller's (1930) belief that this MS. Fieldnotes of H. E. Anthony, "New material was very recent, specimens of the York Academy of Sciences Expedition, insectivoran Nesophontes from this Porto Rico, 1916, 31 May to 30 July" site [title on first page]. Unpublished type- yielded a '3C-adjusted 14C age of 680 ± 50 script, Mammalogy Departmental Li- rcyrbp, while Old World rats from the same brary & Archives, American Museum site spanned a range of 480 ± 60 to 310 ± of Natural History. 40 rcyrbp (MacPhee et al., 1999). The dis- 1918. The indigenous land mammals of Porto parity between the insectivore and rat dates Rico, living and extinct. Mem. Am. implies that the owl-pellet remains at Monte Mus. Nat. Hist., n. ser. 2(2): 331-435. Culo de Maco are not penecontemporaneous 1920. A zoologist in Jamaica. Nat. Hist. 20: but were instead built up over several cen- 156-168 turies. The message here is that one should 1926. Mammals of Porto Rico, living and ex- always be cautious in accepting the validity tinct-Rodentia and Edentata. Sci. of claimed associations between endemics Surv. Porto Rico and Virgin Is. 9(2): and exotics in such contexts, and always at- 97-238. tempt to date the target taxon directly. Brothwell, D. R. 1965. Digging up bones: the excavation, The only conclusion that seems warranted treatment and study of human skeletal from the chronometric evidence available at remains. London: Br. Mus. (Nat. Hist.). present is that, contra Miller (1929a, 1929b) Burney, D. A., L. P. Burney, and R.D.E. MacPhee and Woods et al. (1985), there is still no ev- 1994. Holocene charcoal stratigraphy from idence to support the view that Isolobodon Laguna Tortuguero, Puerto Rico, and portoricensis survived long into the period of the timing of human arrival on the is- European occupation. The documentary in- land. J. Archaeol. Sci. 21: 273-281. dications are that these rodents were to be DeVos, A., R. H. Manville, and R. G. Van Gelder seen only very occasionally even in the ear- 1956. Introduced mammals and their influ- liest days of European colonization, and it is ence on biota. Zoologica, New York therefore entirely plausible that they were in 41: 163-194. precipitous decline by or even before AD Flemming, C., R.D.E. MacPhee, and D. A. 1500. McFarlane Finally, we note that it is somewhat puz- 1998. Thrice bitten: late Quaternary mammal extinctions in the continental and in- zling that Puerto Rican hutias became extinct sular New World. J. Vertebr. Paleontol. if in fact they were kept by pre-Columbian 18 (Suppl. 3): 42A. Amerindians. Human management (as op- Geyh, M. A., and H. Schleicher posed to overkill) of food animals is usually 1990. Absolute age determination: physical conceived to be a hedge against extinction. and chemical dating methods and their It is clear that much remains to be learned application. New York: Springer. about the cause/effect relationships that re- Goodwin, G. sulted in the loss of so many island mammals 1953. Catalogue of the type specimens of Re- during recent historic times (MacPhee and cent mammals in the American Muse- Marx, 1997; MacPhee and Flemming, 1999). um of Natural History. Bull. Am. Mus. Nat. Hist. 102: 207-412. Hoagland, D. B., G. R. Horst, and C. W. Kilpa- REFERENCES trick Allen, J. A. 1989. Biogeography and population biology 1916. An extinct octodont from the island of of the mongoose in the West Indies. In Porto Rico, West Indies. Ann. Acad. C. A. Woods (ed.), Biogeography of the New York Sci. 27: 17-22. West Indies; past, present, and future: Allen, G. M. 611-633. Gainesville, FL: Sandhill 1942. Extinct and vanishing mammals of the Crane. Western Hemisphere with the marine Lawrence, M. A. species of all the oceans. Am. Com- 1993. Catalog of Recent mammal types in the 10 AMERICAN MUSEUM NOVITATES NO. 3278

American Museum of Natural History. Theodor de Booy in the Virgin Islands. Bull. Am. Mus. Nat. Hist. 217: 200 pp. Proc. U.S. Natl. Mus. 54: 507-511. MacPhee, R.D.E., and C. Flemming 1922. Remains of mammals from caves in the 1999. Requiem aternum: the last five hundred Republic of Haiti. Smithson. Misc. years of mammalian species extinc- Collect. 74(3): 8 pp. tions. In R. D. E. MacPhee (ed.), Ex- 1929a. A second collection of mammals from tinctions in near time: causes, contexts, caves near St. Michel, Haiti. Ibid. and consequences: 333-372. New 81(9): 30 pp. York: Kluwer Academic/Plenum. 1929b. Mammals eaten by Indians, owls, and MacPhee, R.D.E., and M. A. Iturralde-Vinent Spaniards in the coast region of the Do- 1995. Origin of the Greater Antillean land minican Republic. Ibid. 82(5): 16 pp. mammal fauna, 1: new Tertiary fossils 1930. Three small collections of mammals from Cuba and Puerto Rico. Am. Mus. from Hispaniola. Ibid. 82(15): 10 pp. Novitates 3141: 30 pp. Mohr, E. MacPhee, R.D.E., and P. A. Marx 1939. Die Baum- und Ferkelratten-Gattungen 1997. The 40,000-year plague: humans, hy- Capromys Desmarest (sens. ampl.) und perdisease, and first-contact extinc- Plagiodontia Cuvier. Mitt. Hambur- tions. In S. Goodman and B. Patterson gishen Zool. Mus. Inst. 48: 48-118. (eds.), Natural change and human im- Nowak, R. M. pact in Madagascar: 169-217. Wash- 1999. Walker's mammals of the world, 6th ington DC: Smithson. Inst. Press. ed., 2 vols. Baltimore: Johns Hopkins MacPhee, R.D.E., C. Flemming, and D. P. Lunde Univ. Press. 1999. "Last occurrence" of the Antillean in- Oviedo y Valdes, G. F. de sectivoran Nesophontes: new radiomet- 1535. Historia general y natural de las Indias. ric dates and their interpretation. Am. Juam Cromberger: Seville. [trans. and Mus. Novitates 3264: 20 pp. ed. by S. A. Stroudemire, Natural His- Martin, P. S. tory of the West Indies, 1959, Univ. 1984. Prehistoric overkill: the global model. North Carolina Stud. Romance Lang. In P. S. Martin and R. G. Klein (eds), Lit. 32: 1-140.1 Quaternary extinctions, a prehistoric Rainey, F G. revolution: 354-403. Tucson: Univ. 1940. Porto Rican archaeology. Sci. Surv. Arizona Press. Porto Rico and Virgin Is. 18(1): 1-208. Martin, P. S., and D. W Steadman Reynolds, T. E., K. F Koopman, and E. E. Wil- 1999. Prehistoric extinctions on islands and liams continents. In R.D.E. MacPhee (ed.), 1953. A cave faunule from western Puerto Extinctions in near time: causes, con- Rico with a discussion of the genus Is- texts, and consequences: 17-55. New olobodon. Breviora 12, 8 pp. York: Kluwer Academic/Plenum. Silva, M., and J. A. Downing Mason, J. A. 1995. CRC handbook of mammalian body 1941. A large archaeological site at Capa, masses. Boca Raton, LA: CRC Press. Utuado, with notes on other Porto Rico Steadman, D. W., T W. Stafford, Jr., D. J. Dona- sites visited in 1914-1915. Sci. Surv. hue, and A. J. T Jull Porto Rico and Virgin Is. 18(2): 209- 1991. Chronology of Holocene vertebrate ex- 272. tinctions in the Galapagos Islands. McFarlane, D. A. Quat. Res. 36: 126-133. 1999. A comparison of methods for the prob- Varona, L. abilistic determination of vertebrate ex- 1974. Catalogo de los mamfferos vivientes y tinction chronologies. In R.D.E. extinguidos de las Antillas. La Habana: MacPhee (ed.), Extinctions in near Acad. Cienc. Cuba. time: causes, contexts, and consequen- Wing, E. S. ces: 95-103. New York: Kluwer Aca- 1991. Dog remains from the Sorce' site on demic/Plenum. Vieques Island, Puerto Rico. Illinois Miller, G. S. State Mus. Sci. Pap. 23: 379-386. 1916. Bones of mammals from Indian sites in Woods, C. A. Cuba and Santo Domingo. Smithson. 1989a. The biogeography of West Indian ro- Misc. Collect. 66(12): 11 pp. dents. In C. A. Woods (ed.), Biogeog- 1918. Mammals and reptiles collected by raphy of the West Indies; past, present, 1999 FLEMMING AND MACPHEE: ISOLOBODON PORTORICENSIS HOLOTYPE 11

and future: 741-798. Gainesville, FL: 771-806. Washington DC: Smithson. Sandhill Crane. Inst. Press. 1989b. A new capromyid rodent from Haiti: 1996. The land mammals of Puerto Rico and the origin, evolution, and extinction of the Virgin Islands. In J. C. Figueroa West Indian rodents and their bearing Colon (ed.), The scientific survey of on the origin of New World hystricog- Puerto Rico and the Virgin Islands, an naths. In C. C. Black and M. R. Daw- eighty-year reassessment of the islands' son (eds.), Papers on fossil rodents in natural history. Ann. New York Acad. honor of Albert Elmer Wood: 58-89. Sci. 776: 131-148. Los Angeles: Nat. Hist. Mus. Los An- Woods, C. A., J. A. Ottenwalder, and WL.R. Oliver geles Cty. 1985. Lost mammals of the Greater Antilles: 1993. Suborder . In D. E. Wil- the summarised findings of a ten weeks son and D. M. Reeder (eds.), Mammal field survey in the Dominican Repub- species of the world: a taxonomic and lic, Haiti and Puerto Rico. Dodo, J. geographic reference, second edition: Wildl. Preserv. Trust 22: 23-42. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last five years are available at World Wide Web site http://nimidi.amnh.org. Or address mail orders to: American Museum of Natural History Library, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: [email protected]

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