THE HERPETOLOGY OF PUERTO RICO Past, Present, and Future I

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1 THE HERPETOLOGY OF PUERTO RICO Past, Present, and Future I I RICHARD THOMAS AND RAFAEL JOGLAR Biology Department PO. Box23360 University of Puerto Rico Rio Piedras, Puerto Rico W E WISH TO EXAMINE the historical pattern of the development of herpetology, the study of amphibians and reptiles, in Puerto Rico. Although there is some subjectivity in dividing this history into time periods, there are obvious trends that seem to be produced by historical change in society, along with conceptual changes in the fields of systematics and ecology, and technological change that permits new kinds of data to be explored. The Puerto Rico Area (sometimes called Greater Puerto Rico) includes the islands of Monito, Mona, Desecheo, Puerto Rico, the Passage Islands of Culebra, Vieques and associated cays, and the US. and British Virgin Islands. This area has also been called the Puerto Rico Island Shelf.' The islands from Puerto Rico through the Virgin Islands, but excluding Monito, Mona, Desecheo, and St. Croix, have been called the Puerto Rico Bank islands; all were connected as one island as recently as 15,000 years ago during the most recent Pleistocene lowering of sea level.*,2 Even at that point of low sea level, Mona, Monito, Desecheo, and St. Croix were separated from the rest. The herpetofauna of the Puerto Rico Area is highly endemic. Only seven (9.2%) of its naturally occurring species have populations that occur natively on other island shelves. It is very similar in generic composition to that of the other Greater Antillean islands. Five of its genera are shared with all other major Antillean island banks, and the other eleven genera are primarily Greater Antillean. One of these, a genus of toads (Peftophryne), is endemic to the Greater Antilles.3 The Puerto Rico Area has no endemic genera of its own. THE EARLY PERIOD The first herpetological species known to inhabit the Puerto Rico Area were described by Linnaeus in his Systemu Nuturue in 175 i4 (see APPENDIX), but these, with one possible exception, are very widespread species that have 181

2 1 a2 ANNALS New York Academy of Sciences been introduced by man onto Puerto Rico and numerous other islands. Rvo species described in the 18th century, the skink Mubuyu mubouyu (probably more than one species), and the rock iguana Cycluru cornutu occur naturally but are not endemic species (both occur on other islands). The gecko Tbecuductylus rupicuudu, also described in that century, has a wide distribution in the Lesser Antilles and South America, but is known only from St. Croix, US. Virgin Islands and Neker Island, British Virgin Islands, in the Puerto Rico Area. The herpetology of Puerto Rico began formally in 1820, when the first endemic species, the giant anole, Anolis cuvieri, was described by Merrem.5 Between 1837 and 1854, the world s first herpetological monograph, DumCril and Bibron s Erpetologie G6n&rule,6 described five species of Puerto Rican provenance. (These species were initially recorded as having come from Martinique, since that was where the collector PlCe shipped them to France. ) From 1854 until the publication of the results of the Scientific Survey in 1928,8 seventeen species were described by a number of workers (Gray, Hallowell, Gunther, Cope, Reinhardt and Lutken, Peters, Zenneck, Meerwarth, and Stejneger) from various European countries and the United States. Of these species, seven were described by Edward Drinker Cope, The Father of North American Herpetology and curator at the Academy of Natural Sciences in Philadelphia. In the early part of this century, under the directorship of Thomas Barbour, the Museum of Comparative Zoology at Harvard became an important center for West Indian herpetological studies, but Barbour was mostly involved in Cuban and Bahamian field work and did not collect in the Puerto Rico Area, although he described C luru pinguis.9 His monographs of the West Indian members of certain widespread genera, such as Spbueroductylus, Ameivu, Cyclura, and l kacbemys, and his Contribution to the Zoogeogrupby of the West Indies, o along with some shorter papers, dealt with Puerto Rican taxa, but other than his description of Cycluru pinguis, he did not give any especial emphasis to the Puerto Rico Area. AndrC P. Ledru reported on a French natural history expedition to Puerto Rico in 1797, in which he had participated as a naturalist. Ledru s description of the reptiles is rather general, even fanciful, and the species, although referred to by Linnaean binomials, are not all precisely identifiable. There are two intriguing reports by Ledru, One is an iguana-like lizard ( Lucertu iguana ) that was hunted with dogs and eaten and said to be common along the coast around Fajardo in the eastern part of Puerto Rico. This could well have been remnant populations of Cycluru pinguis, now extant on Anegada in the British Virgin Islands but known on Puerto Rico only from fossil and subfossil remains2 Even more interesting is Ledru s report of a lagarto de rabo espiral (lizard with a spiral tail), which strikingly calls to mind the curly-tailed Leiocepbulus of Cuba, the Bahamas, and Hispaniola, but known only from Late Pleistocene to sub-recent material on Puerto Rico.2 Unfortunately, specimens from the French expedition do not seem to be extant, and Ledru s description is inadequate. Later in the 19th century, Dr. Agustin Stahl ( ), a physician who lived in Aguadilla, made natural history collections and in published

3 Herpetology Of Puerto Rico T H 0 M A S IJ 0 G LA R 183,I rf 80 FIGURE 1 Cumulative graph 50- of species over time. Each species is recorded as date 4o of first record for the Puerto Rico Area, if known. u) 3o - i YEARS a list of three species of frogs and 20 reptiles (including four marine turtles); he did not describe any species. Although GrantI2 called Dr. Stahl a herpetologist, it appears that his interest extended broadly into natural history. Similarly, Dr. Francisco del Valle Atiles published a list of Los Animales Vertebrados Utiles y Los Dafiinos a la Agricultura in in which he lists all of the species listed by Stahl, except Diploglossus plei. The number of described species increased steadily until the decade of , when there was a sharp increase followed by a relative lull until the turn of the century (FIG. 1). In the 80 years between the description of Anolfs cuvierf and 1900, 27 species (36%) of the presently known herpetofauna of the Puerto Rico Area had been described. THE STEJNEGER-SCHMIDT PERIOD In 1904 Leonhard Stejneger, curator of Herpetology at the United States National Museum, published The Herpetology of pbrto R ~co,~ in which he described seven new species from Puerto Rico and two from Mona Island. His monograph was the result of accumulated collections plus his own field work of approximately two months in The herpetological results of the Scientific Survey were published in 1928* by Karl Patterson Schmidt of the Field Museum in Chicago who visited the island in 1919 (when he was associated with the American Museum of Natural History) and added nine species, one of them from Mona Island. These combined efforts of Stejneger and Schmidt saw the greatest rise in new species descriptions, increasing the known fauna by 21% of the modern count (FIG. 1). THE CHAPMAN GRANT PERIOD In the early 1930s Chapman Grant, a major in the US. Army stationed in Puerto Rico from 1930 to 1932,l3 personally discovered and described 10 species of Puerto Rico Area amphibians and reptiles. His collections resulted in two additional species later described by 0thers.~*J5 From 1931 to 1934

4 184 ANN A L s New York Academy of Sciences Grant published 25 papers, most of them in the Journal of the Department of Agriculture of Puerto Rico, on various aspects of amphibians and reptiles in the Puerto Rico Area. 16-J9 These ranged from new species descriptions to comments on biology and distribution. From 1937 to 1948 he published six more paper^,^*.^^-*^ five in the same journal, which by 1946 had become the Journal of the Department of Agriculture of the University of Puerto Rico. Despite the work by Stejneger and Schmidt, some of the species discovered by Grant were widespread or conspicuous amphibians and reptiles. Although a number were spectacular species, perhaps the most intriguing was the giant anole, Anolis roosevelti, from the island of Culebra. 9 Aside from the two specimens taken by Grant, it has not been found alive since, and its relationship to the other anoles has been enigmatic. Grant made similar but less extensive contributions to the herpetology of Jamaica and Cuba; he alone was responsible for increasing the number of known species in Puerto Rico by 18%. Grant was a self-taught herpetologist and founder of the journal Herpetologica and later the Herpetologists League, which publishes the journal. Grant was certainly an energetic collector, but his discovery of so many species overlooked by others may have been in part the result of increased efficiency and extent of travel that the automobile provided. THE HIATUS During the 22-year period from 1937 to 1959 only one living species was added, the gecko Phyllodactylus wirshingi, described by Kerster and SmithI5 from material collected in the 1930s by Chapman Grant. In 1952, Williams described a fossil tortoise, Geochelone monensis, from Mona Island.46 Very likely the decline in field activity was in some part due to preoccupation with World War 11. Also, the dogma had arisen among biologists that the West Indies was so depauperate in vertebrate species that there could not be much left to discover, perhaps discouraging continued systematic effort. This belief, evidenced in grant reviews and in conversations with colleagues (A. Schwartz, personal communication) may have been the result of a fallacious deduction: that since the West Indies had limited diversity of its fauna4 compared to the mainland they therefore had few species remaining to be discovered. Beginning in the late 1950s two people were primarily responsible for putting this idea to rest: the late Albert Schwartz of Miami-Dade Community College and Ernest Williams of Harvard. THE LATE PERIOD Up to the 1960s virtually all herpetological publication was systematic in nature, focusing on the discovery and description of species. We will summarize some of the major features of herpetological investigation in the region during the last 30 years. From 1959 to the present, systematic discovery continued, but some studies focused on intraspecific variation, and many others dealt with the ecology and natural history. In 1959 Juan A. Rivero and collaborators at the University of Puerto Rico in Mayaguez began a series of papers on frogs The 1960s also saw the start of a series of studies on the physiological ecology, natural history, systematics, and biogeography of frogs and

5 Herpetology 0 f Puerto Rico T H 0 M A S / J 0 G LA R 185 reptiles by Harold Heatwole and collaborators at the University of Puerto Rico at Rio Piedras (see, for examples, refs. 1 and 54-60). Also in the middle 1960s, Albert Schwartz participated in and sponsored herpetological field work that resulted in the publication by him and Richard Thomas of papers describing intraspecific variation of some species and the addition to six others to the fauna.61-'' By this time the new species being described were, generally, cryptic or sibling species or isolates on peripheral islands, such as two species from Isla Deseche0.5~>.~9,~5 Williams and river^^^ and Thomas66 described the peculiar small anole, Anolis occultus, from the interior forests of Puerto Rico. Thomas'" recognized the existence of two sibling species of Eleutherodactylus and described E. coqui. He also described6' the smallest known amniote, a lizard, Sphaerodactylus parthenopion from the Virgin Islands. Drewry and Jones'* described one of the most unusual of all frogs, the live-bearing Eleutherodactylus jasperi from the Sierra de Cayey, which perhaps had escaped detection by earlier investigators because it is a strict bromeliad-dweller. In 1978 Juan Rivero published Los Anfibios y Reptiles de Puerto Rico/The Amphibians and Reptiles of Puerto Rica53 This book has been a useful reference and an important stimulus to local students to pursue herpetological studies. Gregory Pregill,2 then of the University of Kansas, studied Pleistocene fossils of amphibians and reptiles and added to the Puerto Rican fauna two extinct species of the genus Leiocephalus, a genus widespread in the Greater Antilles.z MacPhee and WYSS'~ documented Early Miocene reptiles: a pleurodiran turtle, a boid snake, a possibly iguanid lizard, and a crocodilian, none of which have been unambiguously allocated to genus or species. In addition to Heatwole's papers on the ecology of Puerto Rican species, students of Ernest Williams, including George Gorman, Paul Hertz, Raymond Huey, Stanley Rand, and Thomas Schoener, studied various aspects of the ecology and physiological ecology of Anolis (see, for example, refs ). Ernest William~,~' synthesizing knowledge gained by him and his students, and using Puerto Rico as an exemplar, published The Origin of Faunas: Euolution of Lizard Congeners in a Complex Island Fauna-A Trial Analysis. In this landmark work he theorized that the Puerto Rican anole radiation occurred when competitive exclusion forced species into a limited set of available niches. Adaptation to these niches produced a limited range of "ecomorphs" that occur convergently in different island radiations of anoles. The ecomorph has proved a fruitful concept. Recently, Williams' last graduate student, Jonathan lo so^,^^-^^ now at Washington University, has extended this line of reasoning by linking ecomorphs to experimentally determined performance abilities and combining the results with a phylogenetic analysis in order to hypothesize parallel sequences of niche invasions for Puerto Rican and Jamaican anoles. Further work on anole ecology by Douglas Reagan has added to our knowledge of food habits, density, and distribution within the canopy of the forest.h3 In the Virgin Islands, reported on the Anegada herpetofauna and on Cyclura pinguis. Rudolfo Ruibal and Richard Philibosian86.87 pub-

6 186 ANNALS New York Academy of Sciences lished studies on the anoles, and Philibosian and Yntemaw-89 updated island records for amphibians and reptiles. The late William P. MacLean studied Sphaerod~ctylus,~~9~ published a book on reptiles and amphibians of the Virgin Islands,g2 and published the eminently useful list of islands and their herpetofaunas.93 Recently, Mayer and LazeP4 have reported distributional records for the Virgin Islands. Beginning in the mid-1970s Peter Narins, then of Cornell University and later of UCLA, began analyzing the acoustic and biological properties of the calls of E. coqui and made the discovery that each component of the call has a different function.959% During the 1980s Harvey Pough of Cornell University and Margaret Stewart of the State University of New York at Albany and their graduate students, Theodore Bigen, Karyn Townsend, Daniel Townsend, Lawrence Woolbright and others, conducted studies on the habits, physiology, and ecology of Eleutherodactylus coquf in the Luquillo Forest They added considerably to our knowledge of ecology, behavior, physiology and other aspects of the natural history of this species. Keith Christian, then of the University of Puerto Rico at Rio Piedras, took advantage of the introduced population of the Cuban ground iguana (Cyclura nubila) living on Magueyes Island in the bay of Parguera, and studied various aspects of its biology. W-102 The revolution in phylogenetic inference in recent years, both in algorithms and in the use of molecular data, has affected our views of the biogeographic relationships of Puerto Rico Area species to species on other island~.~~j~3-~~7 Blair Hedges and Carla Hass at Penn State and collaborators have recently provided molecular-based phylogenies for EIeutherodactyIus, Anolis, and Sphaerodactylus. 103~108~109 The most recent descriptions of new species in the area are those of Lazell,llo describing Anolis ernestwilliamsi from Carrot Rock in the British Virgin Islands and Hedges and Thomas111 describing a new blind snake (ZJJphlops hypometbes) from Puerto Rico. A landmark effort with which it is fitting to close the late period, is the comprehensive West Indian herpetofaunal survey of Schwartz and Henders0n.~5 This work, although not specifically of the Puerto Rico Area, surveys the literature on the natural history of West Indian amphibians and reptiles and summarizes Schwartz s 30+ years of field experience. It is an invaluable introduction to the herpetofauna, a fountainhead for those wishing insight into potential research problems. CURRENT STUDIES AND PROBLEMS Allen Lewis of the University of Puerto Rico at Mayaguez has studied the reproductive and feeding habits of Ameiua exsul and Ameiva wetm0rei~~~j3 and is undertaking other studies of lizards. Fernando Bird Pic0 at the University of Puerto Rico at Mayaguez is studying genetic differentiation of populations of Eleutherodactylus. Juan Rivero is preparing a new edition of his book, Los Anfibios y Reptiles de Puerto Rico/Tbe Amphibians and Reptiles of Puerto Rico.53 Recent work on the Puerto Rican rear-fanged colubrids from Thomas s lab at the University of Puerto Rico has tested theories about

7 Herpetology Of Puerto Rico T H 0 M A 5 IJ 0 G LA R 187 the selective advantage of venom in the early stages of its evolution and has also examined the behavior of the prey.53j14-l16 Thomas also continues to study the systematics, feeding habits and ecology of blind snakes, amphisbaenians and Spbaeroductylus, the distribution and species-level systematics of amphibians and reptiles generally. A study of the herpetofauna of the peripheral islands of Puerto Rico is under way, with a focus on founder-induced differentiation of the sphaerodactyls of the S. nicbolsi/.s. townsendf complex. Gordon Rogowitz, also of the University of Puerto Rico at Rio Piedras, is studying various aspects of the physiology of Anolis and Eleutberodactylus. Jonathan Losos of Washington University is investigating further implications of the ecomorph concept in Anolfs.80-8z Leo Fleishman of Union College is examining the photobiology of AnolisII' James Hanken of the University of Colorado and R. Elinson of the University of Toronto are studying early embryonic development in E. coqui. II8,I19 Margaret Stewart of the State University of New York at Albany and Lawrence Woolbright of Siena College continue to study activity patterns and populations of Eleutberodactylus coqui. There are also programs under way (University of Puerto Rico at Humacao, and several zoos in the U.S.A. and Canada) to develop captive breeding programs for Peltopbryne lemur James D. Laze11 of the Conservation Agency, Jamestown, Rhode Island, has instigated studies of the herpetofauna of Guana Island, British Virgin Islands.lzO Mayer and LazeIP reported new records for the Puerto Rico Bank, and they continue to work on the biogeography of the bank. The Puerto Rican boa Epicrates inomatus has been on the Federal Endangered species list for a number of years. Many people think that this species, which is commonly encountered in a variety of habitats throughout the island, may not be truly endangered; Reaganlz1 and Rodriguez and Reagantzz have published on the biology of E. inomatus. Tol~on~~~-~~~ has studied the populations of the smaller boa, Epicrates monensis, of the Puerto Rico Area, from the viewpoint of conservation and management. Interestingly, only since 1988 has this species been known from Puerto Rico proper,iz6 where it appears to be of very restricted distribution. It has become increasingly apparent that a number of the frog species on Puerto Rico proper have been declining in abundance, and the unusual Eleutherodactylus karlschmidti has very likely become extinct; it appears not to have been observed by herpetologists since 1977 (Joglar, Thomas, personal observation; W. P. MacLean, G. Drewry and others, personal communication); and recent attempts to locate it have been unsuccessful. Another species, the live-bearing Eleutberodactylus jasperi, has also not been found since 1981 and is also suspected of having gone extinct. Moreno,IZ7 and Burrowes and JoglarlZ8 attempted to find it and were unsuccessful.127 There is a strong suspicion that some of the other frogs (E. eneidae, E. locustus, E. wigbtmanae) are also declining. As noted by Hedgeslz9 there exists no quantitative baseline data prior to the putative period of decline, and amphibian populations do fluctuatei3" (Joglar, unpublished data on E. coqui, E. portoricensis, E. gryllus, and E. unicolor). However, for a herpetologist who worked in the Puerto Rican forests in the 1960s, the contrast in diversity with the present

8 188 ANNALS New York Academy of Sciences is very marked (Thomas, personal observation). Rafael Joglar and Patricia Burrowes at the University of Puerto Rico continue investigating the status of populations of Eleutherodacfylus in the Luquillo forest and on other parts of the island. The endemic toad Peltophryne lemur is another amphibian whose populations are very restricted on Puerto Rico (it has not been found in the Virgin Islands in about 30 years), presumably a result of relatively long-term decline in populations throughout the island. Anna Goebel of the University of Colorado at Boulder is studying the mitochondria1 DNA differences among the known populations. Around 10% of the Puerto Rico Area herpetofauna comprises non-native species introduced by man during this century (APPENDIX). Puerto Rico is the only one of the major Greater Antillean island banks lacking hylid frogs, but three species have been introduced: The Cuban Osteopilus septentrionalis on Puerto Rico, St. Thomas, and St. Croix; and the North American Hyla cinerea63 on Puerto Rico. (H. cinerea has not been seen for a while by herpetologists and may no longer be extant.) Scinax rubra was discovered in 1988 (Thomas, unpublished data). A large tropical frog, Leptodactylus fallax, was introduced as a possible food source in 1929,33 but it appears not to have become established. The North American bullfrog, Rana catesbeiana was introduced for similar reasons and is widespread throughout the island; in part it preys on native species such as Leptodactylus albilabris, which occurs in the same habitats (Thomas, personal observation). River053 reported that the toad Bufo marinus was introduced in 1920, 1924, and 1926 to control a beetle larva in the cane fields, although Schmidt8 had heard reports of its introduction as early as The crocodilian, Caiman crocodilus, was apparently introduced as a result of released or escaped pets, probably in the 1950s to 1960s, and now occurs in parts of the northern coastal plain; it is well-known from the Tortuguero Lagoon. The common mainland iguana, Iguana iguana, has become well-established and widespread in Puerto Rico and some nearby cays in the last 20 years; some of the Virgin Islands populations are much older. A population of the Cuban ground iguana Cyclura nubila has existed on Isla Magueyes in the Bay of Parguera for some years.53 The geckos, Hemidactylus mabouia and H. brooki, found on many of the Puerto Rico Area islands, were once thought to have been introduced from Africa during colonial times (for example, ref. 28), but Klugel3' concluded that both probably arrived by natural dispersal from West Africa. On Puerto Rico, H. mabouia is evidently displacing H. brooki (Thomas, personal observation); it also now occurs on other islands (e.g., Mona, Culebra), where GrantZ8 did not find it. Kl~geI3~ noted that the evidence for a natural dispersal of H. mabouia is not as strong as for H. brooki. The tortoise Geochelone carbonaria occurs on some of the Virgin Islands and in the Lesser Antille~;I3~ it is not certain whether the populations are introduced,6s but some may be native. ' 3~~33 Rivero indicates (personal communication) that a natricine snake has become established in the southwestern part of Puerto Rico. It is not uncommon to see press reports of exotic pythons or boas being found in residential areas, and one was found in the Luquillo Forest (Luis Rivera, personal communication), but there is no indication that any of these

9 Herpetology Of Puerto Rico T H 0 M A S / J 0 G L A R c : : Oi1991 Periods FIGURE 2 Histogram of publications in the area of systematics (light bars, including distributional records) and non-systematic publications (hatched bars). have become established. The continuing pet trade in exotics implies that the percentage will increase. Any prospective of the herpetofauna must take this into consideration. The pet trade is extensive, even though much of the importation is not legal. The effects on the native fauna of introduced species are potentially disastrous. As noted by Pr0vine,'3~ however, there is another aspect to introduced species: they provide an opportunity to study the evolution of founder-based populations. Although it may be desirable to try to eliminate or control introduced populations, it is important that they be studied, both for their potential contribution to evolutionary theory and for understanding their impact on the native fauna. THE FUTURE AND CONCLUSIONS It is obvious from the foregoing that a major aspect of future herpetology in the Puerto Rico Area will be conservation oriented. As Puerto Rico becomes more and more overpopulated, the ever increasing pressure on its natural areas will necessitate increased efforts at conservation of the species. Conservation-oriented studies will often draw upon and complement more traditional ecological studies. Yet sadly, the natural histories of all but a few of the species of the Puerto Rico Area are very poorly known. Although the description of new species has slowed, there are probably several undescribed species on the bank (Thomas, unpublished data). The possibility of overlooked sibling species or relict species confined to small areas

10 190 A N N A L s New York Academy 0 f Sciences cannot be discounted. Nevertheless, it is probably fair to say that we are near the end point in additions to the herpetofauna. We can see that a distinct shift in emphasis to studies of ecology and natural history has been occurring (FIG. 2). Equally, with the ongoing development of molecular methods for assessing evolutionary and genetic relationships, 35~3~ we expect continued interest in amphibian and reptile relationships as the specific and infraspecific levels leading to a better knowledge of the biogeography of the Puerto Rico Area. REFERENCES 1 HEATWOLE, H. and E MACKENZIE. Herpetogeography of Puerto Rico. IV. Paleogeography, Faunal Similarity and Endemism. Evolution 21: , PREGILL, G. K. Late Pleistocene Herpetofaunas from Puerto Rico. Miscellaneous Publications of the University of Kansas Museum of Natural History (71): 1-72, PREGILL, G. K. Cranial Morphology and Evolution of West Indian Toads (Salientia: Bufonidae): Resurrection of the genus Peltopbryne Fitzinger. Copeia 1981 (2): , LINNAEUS, C. Systema naturaeper Regna ma Naturae, ed. Norwich and London: Jarrold & Sons, 1st edit., MERREM, 8. Tentamen systematis ampbibiorum. Marburg, DUMBRIL, A. M. C. and G. BIBRON. Erpktologie gcncrale ou histoire naturelle des reptiles, vols Paris, STEJNEGER, L. The Herpetology of Porto Rico. Report of the US. National Museum: , SCHMIDT, K. P. Amphibians and Land Reptiles of Puerto Rico, with a List of Those Reported from the Virgin Islands. Annals of the New York Academy of Sciences lo(1): 1-160, BARBOUR, T. Notes on the Herpetology of the Virgin Islands. Proceedings of the Biological Society of Washington 30: , BARBOUR, T. A Contribution to the Zoogeography of the West lndies with Especial Reference to Amphibians and Reptiles. Memoirs of tbe Museum of Comparative ZOO~O~V 44(2): , LEDRU, A. P. Viaje a la isla de Puerto Rico en el afio 1797, ejecutado por una comisidn de sabtos franceses, de orden de su gobierno bajo la direccibn de capitbn Nicolds Baudin, con object0 de bacer indugaciones y coleccfones relativas a la bistoria natural... Universidad de Puerto Rico: Ediciones del Instituto de Literatura Puertorriquefia, GRANT, C. Two Early Puerto Rican Herpetologists. journal of tbe Department of Agriculture of Puerto Rico 16(4): , SMITH, H. M. Chapman Grant, Herpetologica, and the Herpetologists League. Herpetologica 42(1): 1-32, RUTHVEN, A. G. and H. T. GAIGE. Observations on l)rphlops from Puerto Rico and Some of the Adjacent Islands. Occasional Papers of Tbe Museum of Zoolom, University of Micbigan (307): 1-12, KERSTER, H. W. and H. M. SMITH. The Identity of the Puerto Rican Species of Pbylloductylus (Reptilia, Squamata). Herpetologica ll(4): , GRANT, C. Notes on the Ameivas of Porto Rico. Copeia 1931(2): 62, GRANT, C. Notes on Bufo marinus (Linnaeus). Copeia 1931(2): 62, GRANT, C. The Sphaerodaayls of Porto Rico, Culebra and Mona Islands. Journal of the Department of Agrfculture of Puerto Rico 15(3): , GRANT, C. A Revised List of the Herpetological Fauna of Culebra Island. Journal of tbe Department of Agriculture of Puerto Rfco 15(3): 215, GRANT, C. Reestablishment of a Scincid Lost since Journal of tbe Deparfment of Agriculture of Puerto Rico 15(3): , 1931.

11 Herpetology of Puerto Rico T H 0 M A S I J 0 G LA R GRANT, C. Two New Species and a Subspecies of the Genus Anolis. Journal of the Department of Agriculture of Puerto Rico 15(3): , GRANT, C. Notes on Pseudemys stejnegeri Schmidt. Copeia 1931(2): 142, GRANT, C. A New Sphaerodactyl from Porto Rico. Journal of the Department of Agriculture of Puerto Rico 16(1): 31, GRANT, C. The Herpetology of Vieques Island. Journal of the Department of Agriculture of Puerto Rico 16(1): 37-39, GRANT, C. Bufo lemur, a Rare Puerto Rican Toad. journal of the Department of Agriculture of Puerto Rico 16(1): 41, GRANT, C. Sphaemdactylus grandisquamis, a Valid Species. Journal of tbe Department of Agriculture of Puerto Rico 16(1): 31, GRANT, C. Chart for Determining the Sphaerodactylus of the Porto Rican Region. Journal of the Department of Agriculture of Puerto Rico 16(1): 47-49, GRANT, C. The Hemidactyls of the Porto Rican Region. Journal of the Department of Agriculture of Puerto Rico 16(1): 51-57, GRANT, C. A New Frog from Puerto Rico. Journal of the Department of Agriculture of Puerto Rico 16(2): , GRANT, C. The genus Alsophis in the Puerto Rico Area. Journal of the Department of Agriculture of Puerto Rico 16(2): , GRANT, C. A Redescription of Amphisbaena caeca with a discussion of its Relationship to A. bakeri. Journal of the Department of Agriculture of Puerto Rico 16(2): , GRANT, C. The Large Ameivas of the Puerto Rican Region with One New Species. Journal of the Department of Agriculture of Puerto Rico 16(2): , GRANT, C. Herpetological Notes from the Puerto Rico Area. Journal of the Department of Agriculture of Puerto Rico 16(2): , GRANT, C. A New Frog from the Virgin Islands. Journal of the Department of Agriculture of Puerto Rico 16(3): , GRANT, C. Notes on the Boas of Puerto Rico and Mona. Journal of the Department of Agriculture of Puerto Rico 16(3): , GRANT, C. The Herpetology of St. John and Adjacent Keys, U.S. Virgin Islands. Journal of the Department of Agriculture of Puerto Rico 16(3): , GRANT, C. A Genus of Gecko New to the Greater Antilles. Journal of the Department of Agriculture of Puerto Rico 16(3): , GRANT, C. Herpetology of Tortola; Notes on Anegada and Virgin Gorda, British Virgin Islands. Journal of the Department of Agriculture of Puerto Rico 16(3): , GRANT, C. and C. ROOSEVELT. The Herpetology of Caja de Muertos Island and Cardona Key, Porto Rico. Journal of the Department of Agriculture of Puerto Rico 16(1): 47-49, GRANT, C. and C. R. DE SOLA. Antillean Tortoises and Terrapins: Distribution, Status and Habits of Testudo and Pseudemys. * Copeia 1934(2):73-79, GRANT, C. Herpetological Notes with New Species from the American and British Virgin Islands. Journal of the Department of Agriculture of Puerto Rico 21(4): , GRANT, C. A New Name for Alsophis antillensis. Journal of the Department of Agriculture of Puerto Rico 30(2): , GRANT, C. Cyclura stejnegeri-a Portrait. Herpetologica 2: , GRANT, C. Herpetological Notes on St. Croix, Virgin Islands. Herpetologica 2: , GRANT, C. Pseudemys in the West Indies. Journal of Entomology and Zoology 40: 26-28, WILLIAMS, E. E. A New Fossil Tortoise from Mona Island. Bulletin of the American Museum of Natural History (99): DARLINGTON, P. J., JR. Zoogeography: The Geographical Distribution of Animals. New York: John Wiley & Sons, xi pp,, 1957.

12 192 A N N A L S New York Academy of Sciences 48 RIVERO, J. A. Two New Species of Eleutherodactylus from Puerto Rico. Breuiora, Museum of Comparative Zoology (103): 1-6, RIVERO, J. A. Eleutherodactylus hedricki, a New Species of Frog from Puerto Rico (Salientia, Leptodactylidae). Breulora, Museum of Comparative Zoology (103): 1-6, RIVERO, J. A. Notes on the Distribution of Some Puerto Rican Frogs, with a Discussion of the Possible Origin of Eleutherodactylus locustus. * Caribbean Journal of Sciences 3(2): 81-85, RIVERO, J. A,, J. MALDONADO, and H. MAYORGA. On the Habits and Food of Eleuthemdactylus karlschmidti. Caribbean Journal of Sciences 3(1): 25-27, RIVERO, J. A., H. MAYORGA, E. ESTREMERA, and I. IZQUIERDO. Sobre el Bufo lemut: Caribbean Journal of Science 15(3-4): 33-40, RIVERO, J. A. Los anfibios y reptiles de Puerto Rico/The amphibians and reptiles of Puerto Rico. San Juan, PR: Universidad de Puerto Rico Editorial Universitaria, HEATWOLE, H. Comparison of Critical Thermal Maxima of Two Species of Puerto Rican Frogs of the Genus Eleutherodactylus. * Physiological Zoology 38(1): 1-8, HEATWOLE, H. and F. TORRES. Distribution and Geographic Variation of the Ameivas of Puerto Rico and the Virgin Islands. Herpetogeography of Puerto Rico 111. Studies on the Fauna of Curacao and other Caribbean Islands 24(92): , HEATWOLE, H. Herpetogeography of Puerto Rico. V. Description of a New Species of Sphaerodactylus from Desecheo Island. Breuiora (292): 1-6, HEATWOLE, H., F. TORRES, S. BLASINI DE AUSTIN, and A. HEATWOLE. Studies on Anuran Water Balance-I. Dynamics of Evaporative Water Loss by the Coqui, Eleutherodactylus portoricensis. * Comparative Biochemistry and Physiology 28: , HEATWOLE, H. and R. LEVINS. Biogeography of the Puerto Rican Bank: Species krnover on a Small Cay, Cay0 Ahogado. Ecology 54: , HEATWOLE, H. Herpetogeography of Puerto Rico. VII. Geographic Variation in the Anolis cristatellus complex in Puerto Rico and the Virgin Islands. Occasional Papers of the Museum of Natural History University of Kansas (46): 1-17, HEATWOLE, H., R. LEVINS, and M. D. BYER. Biogeography of the Puerto Rican Bank. Atoll Research Bulletin (251): figures, SCHWARTZ, A. Snakes of the Genus Alsophis in Puerto Rico and the Virgin Islands. Studies on the Fauna of Curacao and Other Caribbean Islands 23(90): , SCHWARTZ, A. A Review of the Genus Dromicus in Puerto Rico and the Virgin Islands. Stahlia 9: 1-14, SCHWARTZ, A. and R. THOMAS. A Check-list of West Indian Amphibians and Reptiles. Pittsburgh: Carnegie Museum Special Publications, Vol pp., SCHWARTZ, A. A New Species of Sphaerodactylus (Sauria, Gekkonidae) from lsla Monito, West Indies. Proceedings of the Biological Society of Washington 90(4): , SCHWARTZ, A. and R. W. HENDERSON. Amphibians and Reptiles of the West Indies: Descriptions, Distributions, and Natural Histov. Gainesville: University of Florida Press, xvi pp THOMAS, R. A New Anole (Sauria: Iguanidae) from Puerto Rico. Part 11. Field Observations on Anolis occultus Williams and Rivero. Breuiora (231): 10-16, THOMAS, R. A New Gecko from the Virgin Islands. Quarterly Journal of tbe Florida Academy of Sciences 28(1): 18-22, THOMAS, R. and A. SCHWARTZ. Spbaerodactylys (Gekkonidae) in the Greater Puerto Rico Region. Bulletin of the Florida State Museum lo(6): , THOMAS, R. Additional Notes on the Amphisbaenids of Greater Puerto Rico. Breviora 249: 1-23, 1966.

13 Herpetology of Puerto Rico T H 0 M A S I) 0 G LA R THOMAS, R. New Species of Antillean Eleutherodactylus. Quarterly Journal of the Florida Academy of Sciences 28(4): , THOMAS, R. The Relationships of Antillean Fjphlops (Serpentes: vphlopidae) and the Description of Three New Hispaniolan Species, in Biogeography of the West Indies, edited by C. A. Woods. Gainesville, FL: Sandhill Crane Press, pp , WILLIAMS, E. E. and J. A. RIVERO. A New Anole (Sauria: Iguanidae) from Puerto Rico. Part I. Description. Breviora, Museum of Comparative Zoology (231): 1-21, DREWRY. G. E. and K. JONES. A New Ovoviviparous Frog, Eleutherodactylus jasperi (Amphibia, Anura, Leptodactylidae) from Puerto Rico. Journal of Herpetology lo(3): , MACPHEE, A. R. and A. R. WYSS. Oligo-Miocene Vertebrates from Puerto Rico, with a Catalog of Localities. American Museum Novitates (2965): 1-45, HERTZ, P. E., A. ARCE-HERNANDEZ, J. RAMIREZ-VAZQUEZ, W. TIRADO-RIVERA, and L. VAZQUEZ-VIVES. Geographical Variation of Heat Sensitivity and Water Loss Rates in the Tropical Lizard, Anolis gundlachi. Comparative Biochemistry and Physiol- OD 63A: , HERTZ, P. E. Evaluating Thermal Resource Partitioning by Sympatric Lizards Anolis cooki and A. cristatellus: A Field Test Using Null Hypotheses. Oecologia 90: , HERTZ, P. E. Temperature Regulation in Puerto Rican Anolis Lizards: A Field Test Using Null Hypotheses. Ecology 73(4): , HUEY, R. B. and T. P WEBSTER. Thermal Biology of Anolis Lizards in a Complex Fauna: The Cristatellus Group on Puerto Rico. Ecology 57: , WILLIAMS, E. E. The Origin of Faunas: Evolution of Lizard Congeners in a Complex Island Fauna-A Trial Analysis. Evolutionary Biology 6: 47-89, Losos, J. B. Ecomorphology, Performance Capability, and Scaling of West Indian Anolis Lizards: An Evolutionary Analysis. Ecological Monographs bo(3): , Losos, J. B. Concordant Evolution of Locomotor Behaviour, Display Rate and Morphology in Anolis Lizards. Animal Behaviour 39: , LOSOS, J. B. The Evolution of Convergent Structure in Caribbean Anolis Communities. Systematic Biology 41(4): , REAGAN, D. I? Congeneric Species Distribution and Abundance in a Threedimensional Habitat: The Rain Forest Anoles of Puerto Rico. Copeia 1992(2): CAREY, W. M. The Herpetology of Anegada, British Virgin Islands. Caribbean Journal of Sciences 12: CAREY, W. M. The Rock Iguana, Cyclura pinguis, on Anegada, British Virgin Islands, with Notes on Cyclura ricordi and Cyclura cornuta of Hispaniola. Bulletin of the Florida State Museum of Biological Sciences 19(4): , RUIBAL, R. and R. PHILIBOSIAN. Aggression in the Lizard Anolis acutus. Copeia 1974(2): , RUIBAL, R. and R. PHILIBOSIAN. The Population Ecology of the Lizard, Anolis acutus. Ecology 55(3): , PHILIBOSIAN, R. and J. A. YNTEMA. Records and Status of Some Reptiles and Amphibians in the Virgin Islands. I Herpetologica 32(1): 81-85, PHILIBOSIAN, R. and J. A. YNTEMA. Records and Status of Some Reptiles and Amphibians in the Virgin Islands Herpetologica 34(1): 47-51, MACLEAN, W. P. and R. HOLT. Distributional Patterns in St. Croix Sphaerodactylus Lizards: The Taxon Cycle in Action. Biotropica ll(3): , MACLEAN, W. I? Water-loss Rates of Sphaerodactylus parthenopion (Reptilia: Gekkonidae), the Smallest Amniote Vertebrate. Comparative Biochemistry and Physiology 82A(4): , MACLEAN, W. I? Reptiles and Amphibians of the Virgin Islands. Caribbean, London: MacMillan, vii + 54 pp., 1982.

14 194 AN N A L S New York Academy of Sciences 93 MACLEAN, W. P., R. KELLNER, and H. DENNIS. Island Lists of West Indian Amphibians and Reptiles. Smithsonian Herpetological Information Service (40): 1-47, MAYER, G. C. and J. J. D. LAZELL. Distributional Records for Reptiles and Amphibians from the Puerto Rico Bank. Herpetological Review 19(1): 23-24, NARINS, P. M. and R. R. CAPRANICA. Sexual Differences in the Auditory System of the Treefrog, Eleutheroductylus coqui. I Science 192: , NARINS, P. M. and R. R. CAPRANICA. Communicative Significance of the bo-note Call of the Treefrog, Eleutherodactylus coqui. Journal of Comparative Physiology 127: 1-9, STEWART, M. M. and F. H. POUCH. Population Density of Tropical Forest Frogs: Relation to Retreat Sites. Science 221: , POUCH, F. H., T. L. TAIGEN, M. M. STEWART, and P. F. BROUSSARD. Behavioral Modification of Evaporative Water Loss by a Puerto Rican Frog. Ecology 64(2): , TOWNSEND, D. S. The Costs of Male Parental Care and Its Evolution in a Neotropical Frog. Behavforal Ecology and Sociobiology 19: , CHRISTIAN, K. A. Physiological Consequences of Nighttime Temperature for a Tropical, Herbivorous Lizard (Cyclura nubila). Canadian Journal of Zoology 64: , CHRISTIAN, K. A., I. E. CLAVIJO, N. CORDERO-LOPEZ, E. E. ELIAS-MALDONADO, M. A. FRANCO, M. V. LUGO-RAMIREZ, and M. MARENGO. Thermoregulation and Energetics of a Population of Cuban Iguanas (Cylura nubfla) on Isla Magueyes, Puerto Rico. Copeia 1986(1): 65-69, CHRISTIAN, K. A. and D. TORREGROSA. Effect of Diet on Nitrogenous Wastes of the Iguana, Cyclura nubila. Comparative Biochemistry and Physiology 85A: , SCHOCHAT, D. and H. C. DESSAUER. Comparative Immunological Study of Albumins of Anolis Lizards of the Caribbean Islands. Comparative Biochemfstry and Phy~iology 68A: 67-73, GORMAN, G. C., B. DONALD, M. SOUL& and S. Y. YANG. The Relationships of the Puerto Rican Anolis: Electrophoretic and Karyotypic Studies, in Advances in Herpetology and Evolutfonay Bfology, edited by A. G. J. Rhodin and K. Miyata. Cambridge, MA: Museum of Comparative Zoology, pp , JOCLAR, R. A. Estudio fenetico del genero Eleutherodactylus en Puerto Rico. Curibbeun Journal of Scfence 19(3-4): 33-40, JOGLAR, R. A. Phylogenetic Relationships of the West Indian Frogs of the Genus Eleuthemdactylus: A Morphological Analysis, in Bfogeography of the West Indies: Past, Present, and Future, edited by C. A. Woods. Gainesville, FL: Sandhill Crane Press, pp , GUYER, C. and J. M. SAVAGE. Cladistic Relationships among Anoles. Systematic ZOO~O~Y 35: , HEDGES, S. B. Evolution and Biogeography of West Indian Frogs of the Genus Eleuthemductylus: Slow-evolving Loci and the Major Groups, in Biogeography of the West Indies: Past, Present, and Future, edited by C. A. Woods. Gainesville, FL: Sandhill Crane Press, pp , HAS, C. Evolution and Biogeography of West Indian Sphaerodactylys (Sauria: Gekkonidae): A Molecular Approach. Journal of Zoology (London) 225: , LAZELL, J. D., JR. Biogeography of the Herpetofauna of the British Virgin Islands with Description of a New Anole, in Advances In Herpetology and Evolutionary Biology, edited by A. G. J. Rhodin and K. Miyata. Cambridge, MA: Museum of Comparative Zoology, pp , HEDGES, S. B. and R. THOMAS. Cryptic Species of Snakes (qphlopidae: vphlops) from the Puerto Rico Bank Detected by Protein Electrophoresis. Herpetologica 47(4): , 1991.

15 Herpetology of Puerto Rico THOMASIJOGLAR LEWIS, A. R. Body Size and Growth in Tbo Populations of the Puerto Rican Ground Lizard (Teiidae). Journal of Herpetology 20(2): , LEWIS, A. R. and J. F. SALIVA. Effects of Sex and Size on Home Range, Dominance, and Activity Budgets in Ameiva exsul (Lacertilia: Teiidae). Herpetologica 43(3): , RODRIGUEZ-ROBLES, J. A. and R. THOMAS. Venom Function in the Puerto Rican Racer, Alsophis portoricensfs (Serpentes: Colubridae). Copeia 1992(1): 62-68, RODRIGUEZ-ROBLES, J. A. and M. LEAL. Effects of Prey vpe on Feeding Behavior of Alsophis portoricensis (Serpentes: Colubridae). ]ournal of Herpetology 27(2): , THOMAS, R. and M. LEAL. Feeding Envenomation by Arrhyton exiguum (Serpentes: Colubridae). Journal of Herpetology 27(1): , FLEISCHMAN, L. J., E. R. LOEW, and M. LEAL. Ultraviolet Vision in Lizards. Nature 365(6445): 397, HANKEN, J., M. W. KLYMKOWSKY, C. H. SUMMERS, D. W. SEUFERT, and N. INGEBRIGTSEN. Cranial Ontogeny in the Direct-Developing Frog, Eleutbemdactylus coqui (Anura: Leptodactylidae), Analyzed Using Whole-mount Immunohistochemistry. Journal of Morphology 211: , ELINSON, R. P., E. M. DEL PINO, D. S. TOWNSEND, F. C. CUESTA, and P. EICHHORN. A Practical Guide to the Developmental Biology of Terrestrial-breeding Frogs. Biological Bulletin Woods Hole 179: , LAZELL, J. D., JR. The Herpetofauna of Guana Island: Diversity, Abundance, Rarity, and Conservation, in Status y Dfstribucidn de 10s Reptiles y Anfibios de la region de Puerto Rico, edited by J. Moreno. San Juan: Departamento de Recursos Naturales de Puerto Rico, pp , REAGAN, D. P. Ecology of the Puerto Rican Boa (Epicrates inornatus) in the Luquillo Mountains of Puerto Rico. Caribbean Journal of Science 20(3-4): , RODRIGUEZ, G. A. and D. P. REAGAN. Bat Predation by the Pueno Rican Boa, Epicrates inornatus. I Copeia 1984(1): , TOLSON, P. J. Critical Habitat, Predator Pressures and the Management of Epfcrates monensis (Serpentes: Boidae) on the Puerto Rico Bank: A Multivariate Analysis, in Management of Amphibians, Reptiles, and Small Mammals in North America. U.S. Department of Agriculture, General Technical Report, pp , TOLSON, P. J. Breeding the Virgin Islands Boa Epicrates monensis granti at the Toledo Zoological Gardens. Intmational Zoo Yearbook 28: , TOLSON, P. J. The Conservation Status of Epicrates monensis (Serpentes: Boidae) on the Puerto Rico Bank, in Status y distribucidn de 10s reptiles y anfibios de la region de Puerto Rico, edited by J. Moreno. San Juan: Departamento de Recursos Naturales de Puerto Rico, pp , HEDGES, S. B. and R. THOMAS. The Importance of Systematic Research in the Conservation of Amphibian and Reptile Populations, in Status y distrlbucidn de 10s reptiles y an/tbios de la region de Puerto Rico, edited by J. Moreno. San Juan: Departamento de Recursos Natunles de Puerto Rico, pp , MORENO, J. Status Survey of the Golden Coqui Eleuthemdactylusjasperi, in Statusy distribucidn de 10s reptilesy anfibios de la regton de Puerto Rico, edited by J. Moreno. San Juan: Departamento de Recursos Naturales de Puerto Rico, pp , BURROWES, P., and R. JOGLAR. A Survey of the Population Status and Ecological Evaluation of Three Pueno Rican Frogs, in Status y distribucidn de 10s reptiles y anfibios de la region de Puerto Rico, edited by J. Moreno. San Juan: Departamento de Recursos Natunles de Puerto Rico, pp , HEDGES, S. B. Global Amphibian Declines: A Perspective from the Caribbean. Biodiversity and Conservation 2: , 1993.

16 196 ANN A LS New York Academy of Sciences 130 PECHMANN, J. H. K., D. E. SCOTT, R. D. SEMLISCH, J. P. CALDWELL, L. J. VITT, and W. GIBBONS. Declining Amphibian Populations: The Problem of Separating Human Impacts from Natural Fluctuations. Science 253: , KLUGE, A. The Evolution and Geographical Origin of the New World Hemidactylus mabouia-bmoki complex. (Gekkonidae, Sauria). Miscellaneous Publications, Museum of Zoology, University of Michigan (138): 1-78, CENSKY, E. J. Geochelone carbonaria (Reptilia: Testudines) in the West Indies. Florida Scientist 50(2): , LAZELL, J. D., JR. Tortoise, cf. Geochelone carbonaria, from the Pleistocene of Anguilla. Northern Lesser Antilles. Journal ofherpetology 27(4): , PROVINE, W. B. Founder Effects and Genetic Revolutions in Microevolution and Speciation: An Historical Perspective, in Genetics, Speciation, and the Founder Principle, edited by L. V. Giddings, K. Y Kaneshiro, and W. W. Anderson. New York: Oxford University Press, pp , I35 HILLIS, D. M. and C. MORITZ, Editors. Molecular Systematics. Sunderland, MA: Sinauer Associates, xvi pp AVISE, J. J. A,, R. M. BALL, E. BIRMINGHAM, T. LAMB, J. E. NEIGEL, C. A. REEB, and N. C. SAUNDERS. Intraspecific Phylogeography: The Mitochondria1 Bridge between Population Genetics and Systematics. Annual Review of Ecology and Systematics 18: , 1987.

17 APPEND I X First Record List of Species known from the Puerto Rico Area with Date of Original Description and, if Different, Date of Date PRArea Species Author Prov Range Group s Bufo murinus L i n n a e u s I Bank + St. Croix Amph: Bufonidae Iguana iguana Linnaeus I, N? Bank + St. Croix Rept: Iguanidae s Cayman crocodvlus Linnaeus I PR Rept: Alligatoridae Tbecadactylus rupicaudu Houttuyn N VIs Rept: Gekkonidae Mubuyu mubouyu Lacepede N PR Area Rept: Scincidae Cvcluru cornutu Bonnaterre N Mona ReDt: Iauanidae Hvlu cinerea Schneider I PR Rept: Iguanidae 1802 I988 Scinuxrubru Daudin I PR Amph: Hylidae s Runu cutesbeiunu Shaw I PR s Hemiductvlus mabouiu Moreau de lonnes I PR Area Amph: Hylidae ReDt: Gekkonidae 1820 Anolis cuvieri Merrem N PR Rept: Polychridae Geocbelone curbonariu Spix I? VIS Rept: Testudinidae 1829 Amphisbaena caeca Cuvier N PR + Vieques Rept: Amphisbaenidae s? Cycluru nubila Gray I PR (Magueyes I.) Rept : lguanidae 1837 Anolis Dulcbellus Dumeril and Bibron N PR Bank Rept: Polychridae 1837 Anolis cristatellus Dumeril and Bibron N PR Bank Rept: Polychridae 1839 Diploglossus plei Dumeril and Bibron N PR Rept: Anguidae s Osteopilus septentrionalis Dumeril and Bibron I PR Amph: Hylidae 1843 Epicrutes inornutus Reinhardt N PR Rept: Boidae 1844 Typblops plutycepbulus Dumeril and Bibron N PR Rept: Typhlopidae (confinued) B 2 1 B r 3

18 A P P E N D I X (continued) Date PRArea Species Author Prov Range Group 1844 Tvpbfops ricbardi Durneril and Bibron N Passage Is. + VI Rept: Typhlopidae 1845 Hemidactylus brooki Gray N PR Rept: Gekkonidae 1857 Anolis acutus Hallowell N St. Croix Reot: Polvchrididae ~~ ~~ 1859 LeDtodactvlus aqilabris Gunther N PR Bank Arnph: Leptodactvlidae 1859 Spbaemdactylus macrolepis Gunther N PR Bank Rept: Gekkonidae 1862 Amphisbaena fenestrata cope N Vls Rept: Amphisbaenidae 1862 Anolis stratulus cow N PR Bank Reot: Polvchridae 1863 Eleutherodactylus antillensis Reinhardt and Luetken N PR Bank Amph: Leptodactylidae 1863 Eleutherodactylus Ientus cope N Vls + St. Croix Amph: Leptodactylidae 1863 Ameiva exsul cope N PRBank Rept: Teiidae 1863 Ameiva DoloDs cow N St. Croix Rept: Teiidae 1863 Alsopbis portoricensis Reinhardt and Luetken N PR Bank + Mona + Desecheo Rept: Colubridae 1863 Alsopbis sancticrucis cope N St. Croix Rept: Colubridae 1863 Avbyton emguum cope N PR Bank Rept: Colubridae 1868 PeltoDhme lemur cow N PR Bank Amph: Bufonidae 1876 Anolis RUndhCbi Peters N PR Rept: Polychridae 1876 Anolis krugi Peters N PR Rept: Polychridae 1898 Epicrates monensis Zenneck N PRBank + Mona Rept: Boidae 1901 Eleut&ero&ctylccs monensis Meerwarth N MOM Amph: Leptodactvlidae 1901 Spbaerodactylus monensis Stejneger N Mona Rept: Gekkonidae 1904 Eleutherodactylus ricbmondi Stejneger N PR Amph: Leptodactylidae 1904 Eleutberoductvlus unicolor Steineaer N PR Amoh: Leotodactvlidae L W x P R 2

19 1904 Amphisbaena bakeri Stejneger N PR Rept: Amphisbaenidae 1904 Typhlops rostellatus Stejneger N PR Rept: Typhlopidae i$ 1904 Anolis poncensis Stejneger N PR Rept: Polychridae Anolis monensis Stejneger N Mona, Monito Rept: Polychridae 1904 Anolis evermanni Stejneger N PR Rept: Polychridae 1913 Ameiva wetmorei Stejneger N PR Rept: Teiidae % 1917 Cyclura pinguis Barbour N PR Bank Rept : lguanidae 1920 Eleutherodactylus locustus Schmidt N PR Amph: Leptodactylidae Eleutherodactylus brittoni Schmidt N PR Amph: Leptodactylidae Eleutherodactylus gryllus Schmidt N PR Amph: Leptodactylidae 0 -I 1920 Eleutherodactylus wightmanae Schmidt N PR Amph: Leptodactylidae 1926 Typhlops monensis Schmidt N Mona Rept: Typhlopidae z 1927 Eleutherodactylus portoricensis Schmidt N PR Amph: Leptodactylidae ~1 s' 1928 Trachemys sfejnegeri Schmidt N PRBank Rept: Emydidae Sphaerodactylus klauberi Grant N PR Rept: Gekkonidae l Sphaerodactylus nichoki Grant N PR Rept: Gekkonidae *I a 8 'tr f r, 0 * 0 > SDhaerodactvlus roosevelti Grant N PR Reot: Gekkonidae 1931 SDhaerodactvlus townsendi Grant N PR + Vieques Rept: Gekkonidae 1931 Anolis cooki Grant N PR Rept: Polychridae 1931 Anolis roosevelti Grant N Passage + VIs Rept: Polychridae 1931 Eleutherodactylus karkchmidti Grant N PR Amph: Leptodactylidae 1932 Eleutherodactvlus cochranae Grant N PR Bank Amph: Leptodactvlidae 1932 Eleutherodactvlus cooki Grant N PR Amph: Leptodactvlidae 1932 Sphaerodacfylus gaigeae Grant N PR + Vieques Rept: Gekkonidae e (continued) %

20 A P P E N D I X (continued) Date PRArea Species Author Prov Range Group 1935 Typblops canti Ruthven and Gaige N PR Rept: Typhlopidae 1937 Spbaerodactylus beattyi Grant N St. Croix Rept: Gekkonidae 1952 Geocbelone monensis Williams F Mona Reot: Testudinidae 1955 Pbyllodactylus wirsbingi Kerster and Smith N PR Rept: Gekkonidae 1959 Eleutberoductylus eneidae Rivero N PR Amph: Leptodactylidae 1963 Eleutherodactylus bedricki Rivero N PR Amph : Leptodact ylidae 1964 AmDbisbaena scbmidti G W N PR Reot: Amohisbaenidae 1965 Spbaerodactylus partbenopion Thomas N VIs Rept: Gekkonidae 1965 Anolis occultus Williams and Rivero N PR Rept: Polychridae 1966 Eleutbaodactylus coqui Thomas N PR + Passage Is. Amph: Leptodactylidae 1966 Eleutbaodactvlus scbwartzi Thomas N VIs Amoh: LeDtodactvlidae 1966 AmDbisbaena xera Thomas N PR Reot: Amphisbaenidae 1966 Typblops catapontus Thomas N VIs Rept: Typhlopidae 1- Spbaerodactylus levinsi Heatwole N Desecheo Rept: Gekkonidae 1976 Eleutberodactylus jasperi Drewry and Jones N PR Amph: Leptodactylidae 1976 Anolis desechensis Heatwole N Desecheo Reot: Polvchridae 1977 Spbaerodactvlus micropithecw Schwartz N Monito ReDt: Gekkonidae 1981 Leiocepbalus etheridpi Pregill F PR Rept: Tropiduridae 1981 Leiocepbalus partitus Pregill F PR Rept: Tropiduridae 1983 Anolis ernestwilliamsi Lazell N VIs (Carrot Rock) Rept: Polychridae 1991 TvDblops hvdometbes Hedges andthomas N PR Reot: Tvvhlooidae ~ ~ Prov = provenance; N = natively occurring; F = fossil records only; VIs = Virgin Islands; Passage Is. = Vieques, Culebra, and associated islets. PR Area indicates that a species iswidespread throughout the area encompassed by all islands listed. PR Bank refers to Puerto Rico and associated cays, the Passage Islands, and the Virgin Islands, except St. Croix. N h a 5 B 1 ~~