In Search of the Extinct in Cave Deposits of Isla de Mona, P.R. by Ángel M. Nieves-Rivera, M.S. and Donald A. McFarlane, Ph.D. Isolobodon portoricensis, the extinct have been domesticated, and its abundant (14C) date was obtained on charcoal and Puerto Rican hutia (a large guinea-pig like remains in kitchen middens indicate that it bone fragments from Cueva Negra, ), was about the size of the surviving formed part of the diet for the early settlers associated with hutia bones (Frank, 1998). (Rodentia: (Nowak, 1991; Flemming and MacPhee, This analysis yielded an uncorrected 14C age Capromydae). Isolobodon portoricensis was 1999). This of hutia was extinct, of 380 ±60 before present, and a corrected originally reported from Cueva Ceiba (next apparently shortly after the coming of calendar age of 1525 AD, (1 sigma range to Utuado, P.R.) in 1916 by J. A. Allen (1916), European explorers, according to most 1480-1655 AD). This date coincides with the and it is known today only by skeletal remains historians. final occupation of Isla de Mona by the Taino from (, , The first person to take an interest in the Indians (1578 AD; Wadsworth 1977). The Île de la Gonâve, ÎIe de la Tortue), Puerto faunal remains of the caves of Isla de Mona purpose of this article is to report some new Rico (mainland, Isla de Mona, Caja de was mammalogist Harold E. Anthony, who paleontological discoveries of the Puerto Muertos, Vieques), the Virgin Islands (St. in 1926 collected the first Puerto Rican hutia Rican hutia in cave deposits of Isla de Mona Croix, St. Thomas) (Rímoli, 1976; Díaz-Díaz, bones. Although his field notes (Anthony, and discuss the recent alleged sightings of 1983; Nowak, 1991; Woods, 1996), and 1926) provide an interesting account of his this rodent on the island and on mainland from San Salvador Island (Mylroie, personal expedition, they are not very useful when it . communication). Some authors believe that comes to defining specific locations. In 1959, the Puerto Rican hutia was native to Puerto Clifford A. Kaye published his Geology of OTHER FAUNAL REMAINS Rico (Nowak, 1991) and was transported to Isla Mona, Puerto Rico, and Notes on the Age Interesting faunal remains continue to be other islands by aborigines, the Taino Indians. of , in which he reported hutia found in various caves of Isla de Mona. Other authors, however, strongly disagree bones (= Puerto Rican hutia) which were Common, rare, and extirpated marine and and considered it native to Hispaniola, collected from several cave deposits and from land such as Red-footed and Brown transported by Indians to Puerto Rico an archaeological site (= Taino site) at Playa Boobies (Sula sula, S. leucogaster), White- (Rímoli, 1976; Díaz-Díaz, 1983; Woods, Sardinera, on the west coast of Mona (Kaye, tailed Tropicbird (Phaethon lepturus), 1985). Ultimately, this can only be resolved 1959). Kingbird (Tyrannus sp.), Black-capped Petrel by paleontology. The Puerto Rican hutia may Recently, the only relevant radiocarbon (Pterodroma hasitala), Audubon Shearwater (Puffinus lherminieri), Puerto Rican Parakeet (Aratinga chloroptera maugei), Puerto Rican (Amazona vittata vittata), reptiles such as the extinct Mona Tortoise (Geochelone [Monachelys] monensis), and a small vertebrae from an undetermined crocodile species have been collected from cave deposits (Frank & Benson, 1998). Specimens of the endemic Mona Coquí (treefrog) (Eleutherodactylus monensis), the Mona Ground Iguana (Cyclura stejnegeri), and a few species of introduced small (eg., Domestic [Felis catus], Black [Rattus rattus of Pippin 1961, and Woods 1996), Brown Rat (?) [R. norvegicus of Wiewandt, 1973], and House [Mus musculus]) may use caves as refugia, but their bone fragments are rare in the cave deposits. A recent paleontological survey has collected a few shark teeth (Carcharodon?) embedded in eroded beachrock, which add the promise of new discoveries. Three bat species, the Antillean Ghost- faced Bat (Mormoops blainvillii), Puerto Rican Mastiff Bat (Molossus molossus fortis), and the Great Bulldog Bat (Noctilio leporinus mastivus) are known to live on Isla de Mona (Woods, 1996), but bat bone fragments are extremely rare in the caves. Chemical and biological activity within fresh guano digests these bones and they do not often survive to become part of the fossil record (Frank & Benson, 1998). Nevertheless, a few small insectivorous bat colonies are present today (eg., a bat cave in Uvero; the bat chamber in Cueva Caballo/Pájaros, and few individuals in Cueva de Chito, both in Playa Reconstruction of the Puerto Rican hutia. Drawing by the senior author. de Pájaros). The Great Bulldog Bat is rarely

92 NSS NEWS, April 2001 seen in Isla de Mona, although on the Puerto Rico mainland it is often seen fishing by low flying between the lagoon and the nearshore between Pt. Guaniquilla and Joyuda (Cabo Rojo), the mangrove channels of La Parguera (Lajas), and on Islas Culebra and Vieques. Invertebrate fossils are common in Isla de Mona cave deposits, and include gastropods (Bulimulus, Cerion, Chondropoma, Lacteoluna, Lucidella, Suavitas), scallops (Pecten), corals (Acropora, Caulastrea, Montastrea, Mussa, Stylophora, Undaria), planktonic foraminiferans, benthic foraminiferans (Amphistegina, Archaias, Nummulites), bryozoans, and echinoderms (spines and test fragments, probably Cidaris). Red and green algae deposits are also common in reef complex deposits through the island (González et al., 1997).

THE CAVES Cueva Negra is an abandoned mined phreatic cave with a shallow sediment floor, carved in the Lirios Limestone unit and located behind Playa Sardinera, west Isla de Mona, at an altitude of 10 m above mean village?). The junior author collected the third caves should be left in place and reported to sea level. At the two entrances, piles of rocks Puerto Rican hutia hemimandible in 1994 local paleontologists, archaeologists, or have been stacked to support tram rails for from a depth of 30 centimeters in an museums. The context in which the bones the transport of bat guano during the mining excavation into the sediment floor of the are found is important to the interpretation operations during the late 19th and early 20th Cathedral Room, Cueva Lirio. Unfortunately, of the finds, and vital information can be lost centuries. Aboriginal pictographs are present none of these specimens has yet been through careless collection. In 1919, Isla de on the soft limestone walls, together with radiocarbon-dated, but the Playa Sardinera Mona was declared “Insular Forest” and a other designs that have obviously been faked specimens are likely to be contemporaneous decade ago it was afforded the status of a by more recent visitors. An early name written with late Taino occupation (<1500 years natural reserve. Today the island is protected in one of the chamber walls reads “John before present) and the Cueva Lirio by local and federal laws, and written Dennis 1745 RD” and pre-dates the guano- specimen is likely to be latest Pleistocene (< permission to collect fossils (or other material) mining period. Wall writings such as these 20,000 years before present) or Holocene for educational or scientific purposes must were previously noticed by Wadsworth (< 10,000 years before present) in age. be obtained a priori from the Puerto Rico (1977). Subfossil bones of the extinct Puerto Rican Department of Natural & Environmental Cueva Lirios is an extensive cave formed hutia are fragile and must be handle with Resources (PRDNER). Unfortunately, guano at the interface of the Lirios Limestone and care. Paleontological remains discovered in mining and decades of careless tourism have the underlying Isla de Mona Dolomite. This cave is located next to El Faro (“The Lighthouse”), eastern Isla de Mona. Cueva Lirios opens spectacularly into the face of a 45-meter-high, overhung seacliff. This

impressive cave was also extensively used for Luis Padilla guano mining operations during the early years of this century. The cave is famous for a spectacular chamber known as the “Cathedral Room,” cave pearls, wind- sculpted speleothems, and spectacular coastal views. Cuevas Lirios and Negra are among the most popular caves visited by hikers, and have been favorite paleontological sites visited by scientists in past years.

THE COLLECTION In 1989 and 1995, the senior author found two Puerto Rican hutia jawbones in the Playa Sardinera area. These hemimandibles were collected incidentally from topsoil while obtaining samples for the study of soil microfungi. One was collected at the entrance of Cueva Negra, the other in the palm forest Tony Nieves (PRDNER) sitting at the bottom of the larger depression of Cuevas del close to Cueva Negra (on the ancient Taino Centro.

NSS NEWS, April 2001 93 Commenting on extinct in the Maricao Commonwealth Forest, he wrote: “However, recent evidence from Maricao suggest that the “Jutía” (Isolobodon portoricensis), a large rodent, believed by Timmy Ortiz (PRDNER). scientists to have been Lower jawbone (hemimandible) of the extinct for approximately extinct Puerto Rican hutia collected in 200 years, may still exist Sardinera. there [Maricao Forest] in small numbers. Every effort should be made to exacted a toll on many of the caves of Isla verify this in the valleys de Mona. Cueva Negra in particular has on the north side of the been heavily vandalized. forest and steps should be Senior author viewing Indian rock art from a cave next to Pt. taken to provide RECENT SIGHTINGS? Capitán. Are they genuine? increased protection to Early accounts of surviving Puerto Rican this zone” (Raffaele, 1977). reduced by excessive human hunting hutias date from 1916, when the hutia Raffaele recorded other sightings of the (Nowak, 1991). The Puerto Rican barn discoverer in person (J. A. Allen) reported Puerto Rican hutia in the gorge of the Río is not known from the fossil record of Isla de fresh remains from caves of mainland Puerto Casey, Las Marías municipality (mainland Mona. Woods (1981) believes that the Rico. Allen also reported archaeological Puerto Rico), reporting: “There are extremely Puerto Rican hutia was a hundred times deposits from the Dominican Republic, strong indications that the “jutía” more common on Hispaniola than was implying a fairly recent (Rímoli, (Isolobodon portoricensis)... survived in this Plagiodontia but disappeared (while 1977). paleontologist Alexander area at least until as recently as the early Plagiodontia survived) because of its less Wetmore reported “an agouti-like ” 1960s. The species could still survive here secretive habits. Before the arrival of (similar to the Puerto Rican hutia) that was in extremely small numbers thus Europeans on Hispaniola, large boas (eg., seen by the grandfathers of old men in Cayey investigations to determine the status of the Epicrates striatus) most likely preyed upon and Mameyes in his Bird remains from the animal should be continued. The valley of birds, now-extinct (Brotomys, Caves of Porto Rico (Wetmore, 1922). Also, the Casei River should be protected from Isolobodon), living rodents (Plagiodontia), Koopman et al. (1953) discussed the development until more information is and extinct insectivores (Nesophontes) possibility that Puerto Rican hutia is still alive gathered on whether the Jutía survives” (Henderson et al., 1987). in western Puerto Rico, and reviewed the (Raffaele, 1977). All of these observations, along with recent genus Isolobodon. In the 1960s, Walter In a series of scientific expeditions sightings of hutia-like mammals from remote Murray Chiesa of the Puerto Rico Institute between 1984-85, Charles A. Woods, J. A. regions of Isla de Mona (eg., Pt. Capitán, of Culture spoke with a number of people Ottenwalder, and W. L. R. Oliver received Booby colony, Bajura de los Cerezos, Cuevas who claimed they had shot and eaten Puerto reports of living in Hispaniola and del Centro, Corral de los Indios, and Playa Rican hutia, and he himself reported having Puerto Rico that resemble the Puerto Rican Cocos) have raised a number of questions. seen one, but was unable to catch the animal hutia. However, these authorities concluded Probably the most important is “is the Puerto (Raffaele, 1977). that Isolobodon is probably extinct, though Rican hutia still alive?” Regrettably, the Certainly, the Puerto Rican hutia was one they suggest it may have persisted in small answer is “probably not”. A variety of of the last Greater Antillean land mammals numbers (eg., in Puerto Rico and Île de la mammals have been introduced to Isla de to become extinct. Modern accounts of hutia Tortue, according to Woods, 1996) until Mona, including , , rabbits, , and sightings from Puerto Rico (including Isla de wiped out by the Small Indian Mongoose mice (Wiewandt 1973; Wadsworth, 1977), Mona) have continued through the 1970s. (Herpestes auropunctatus) introduced into and some of these have survived as feral Two such accounts were originally reported the West Indies beginning in the 1870s populations. Reported hutia sightings must by ornithologist Herbert A. Raffaele while (Woods et al., 1985). Fortunately, the Small always be considered in the context of a studying critical wildlife areas in Puerto Rico. Indian Mongoose was never introduced to potential misidentification. Also, reports of Isla de Mona, so it cannot “fresh” owl pellet remains of hutias from have been a factor in the earlier in the 20th century must now be re- decline of the Mona evaluated in the light of the discovery of fresh population. remains of the extinct insectivore Skeletal remains of the Nesophontes from the Dominican Republic, Puerto Rican hutia are some preserving traces of hair and dried often present in cave tissue, that were shown by radiocarbon deposits deposited by the dating to be more than five centuries old extinct Puerto Rican barn (MacPhee et al., 1999). owl ( cavatica), not T. On the other hand, one should also keep ostolaga of Nowak in mind that the survival of extinct species is (1991). This owl preyed possible. Several such “living fossils” are on the hutias and known. An example is the famous coelacanth Cathedral Room perhaps disappeared (Latimeria chalumna), a huge fish thought of Cueva Lirios (Pérez-Rivera, 1977) extinct but then discovered living. Examples when their numbers were of primitive organisms discovered alive, then

94 NSS NEWS, April 2001 found in the fossil record include the okapi on recent mammalian in Puerto Rico. (Okapia johnstoni), the mountain gorilla American Museum Novitates 3278:1-11. (Gorilla gorilla beringei), the New Zeland Frank, E. F. 1998. A radiocarbon date of 380±60 tuatara (Sphenodon punctatus), the BP for a Taino site, Cueva Negra, Isla de Mona, Puerto Rico. Journal of Caves and Karst Studies

Tasmanian marsupial ‘wolf’ (Thylacinus Luis A. Padilla 60:101-102. cynocephalus), the brachiopod or lamp-shell Frank, E. F. and R. Benson. 1998. Vertebrate (Lingula), and even a Paleozoic plant paleontology of Mona Island, Puerto Rico. (Welwitschia mirabilis) (Heuvelmans, 1958). Journal of Caves and Karst Studies 60:103-106. Many of those species were discovered (or González, L. A., H. M. Ruiz, B. E. Taggart, A. F. Budd rediscovered) in the 20th century, and many and V. Monell. 1997. Geology of Isla de Mona, more await to be discovered (eg., the Puerto Rico. In H. L. Vacher and T. M. Quinn “mapinguari” or mylodontid ground sloth of (eds.), Geology and hydrogeology of carbonate the Amazon; Oren, 1993). Future eyewitness islands, Vol. 54, p. 327-358. Developments in accounts of hutia sightings should be taken Sedimentology, Elsevier. Henderson, R. W., T. A. Noeske-Hallin, J. A. seriously, but every effort should be made to Ottenwalder and A. Schwartz. 1987. On the diet obtain tangible evidence. Current evidence of the boa Epicrates striatus on Hispaniola (Haiti), suggests that the Puerto Rican hutia was with notes on Epicrates fordi and Epicrates probably still alive when the first Europeans gracilis. Amphibia-Reptilia 8:251-258. colonized Puerto Rico, although it cannot Heuvelmans, B. 1958. On the track of unknown have been common or have persisted very animals. Hill & Wang, NY. long because no specimens were ever Kaye, C. A. 1959. Geology of Isla Mona, Puerto returned to the great colonial museums of Rico, and notes on the age of Mona Passage. US Caves next to Cueva de Chito. Europe. Late 20th century sightings of live Geological Survey Professional Paper 317C:141- hutias on mainland Puerto Rico and Isla de 178. Koopman, K. F., T. E. Reynolds and E. E. Williams. Colón (ed.), The scientific survey of Puerto Rico Mona are probably cases of mistaken identity, 1953. A cave faunule from western Puerto Rico and the Virgin Islands: an eighty-year but it is possible that the species survived in with a discussion of the genus Isolobodon. reassessment of the island’s natural history, Vol. small numbers into the 19th or even 20th Breviora 12:1-8. 776, p. 131-148. Annals of the New York centuries in the remotest areas. Only a MacPhee R. D. E., C. Flemming, and D. P. Lunde. Academy of Sciences. rigorous and well-funded program of 1999. “Last Occurrence” of the Antillean Woods, C. A., J. A. Ottenwalder and W. L. R. Oliver. radiocarbon dating can finally establish the insectivoran Nesophontes: new radiocarbon 1985. Lost mammals of the Greater Antilles: the facts. dates and their interpretation. American Museum summarized findings of a ten weeks field survey Novitates 3261:1-20. in the Dominican Republic, Haiti and Puerto Rico. ACKNOWLEDGMENTS Nowak, R. M. 1991. Walker’s mammals of the world. Dodo 22:23-42. We thank Edward F. Frank, Sue Fifth edition. Vol. 2. John Hopkins University Press, Baltimore and London, p. 932. Magdalene (Dept. Geology and Geophysics, Oren, D. C. 1993. Did ground sloths survive to University of Minnesota, Minneapolis), and Recent times in the Amazon region? Goeldiana Clare Flemming (American Museum of Zoologia 19:1-11. Natural History, New York) for their Pérez-Rivera, R. A. 1977. Es posible que [Tyto assistance and for granting us further cavatica] no esté extinto. Science-Ciencia 5:40. literature on the subject. Field work was Pippin, W. F. 1961. The distribution and movement conducted with permission of the Puerto Rico of roof rats on Mona Island, West Indies. Journal Department of Natural and Environmental of Mammalogy 42:344-348. Resources. The senior author wishes to thank Raffaele, H. A. 1977. The status of some endangered species in Puerto Rico with particular emphasis Mrs. Carmen Amorós and her staff of the on Isolobodon (Rodentia). 1976 Symposium of Main Library, University of PR at Mayagüez the Puerto Rico Department of Natural Resources, for their valuable cooperation during the San Juan, PR, p. 100-104. course of this study. This study was partially Rímoli, R. O. 1976. Roedores fósiles de La supported by the University of Puerto Rico Hispaniola. Serie Científica 3, Universidad Sea Grant College Program Budget (90-4) Central del Este, San Pedro de Macorís, RD, p. for small project to the senior author. 35-37. Wadsworth, F. H. 1977. Reseña histórica de la Isla REFERENCES de Mona. Revista/Review Interamericana 6:587- Allen, J. A. 1916. An extinct octodont from the Island 621. of Porto Rico, W.I. Annals of the New York Wetmore, A. 1922. Bird remains from the caves of Academy of Science 27:17-22. Porto Rico. Bulletin of the American Museum of Anthony, H. E. 1926. Field notes of the 1926 Natural History 46:297-333. expedition to Mona and other West Indian islands. Wiewandt, T. A. 1973. Mona amphibians, reptiles, American Museum of Natural History, NY. and mammals. In Junta de Calidad Ambiental (Unpublished manuscript; Department of (ed.), Mona and Monito Islands: an assessment Mammalogy). of their natural and historical resources, Vol. 2, Díaz-Díaz, C. A. 1983. Los mamíferos de Puerto Rico. Appendix L, p. 1-13. Oficina del Gobernador, En J. L. Vivaldi (ed.), Compendio enciclopédico San Juan, PR. de los recursos Naturales, Vol. 1, p. 109-110. Woods, C. A. 1981. Last endemic mammals in Departamento de Recursos Naturales, San Juan, Hispaniola. Oryx 16:146-152. PR. Woods, C. A. 1985. Evolution and systematics of Flemming, C. and R. D. E. MacPhee. 1999. endemic Antillean rodents. Acta Zoologica Redetermination of holotype of Isolobodon Fennica 170:199-200. portoricensis (Rodentia, Capromydae), with notes Woods, C. A. 1996. The land mammals of Puerto Rico and the Virgin Islands. In J. C. Figueroa

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