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Home , Bananaquit, Clusia, Elfin woods warbler, Maricao State Forest, Puerto Rican tanager, Puerto Rican tody

STATUS, ABUNDANCE, AND DISTRIBUTION OF OF ,

1 2 ADRIANNE G. TOSSAS AND CARLOS A. DELANNOY 1Department of Biology, University of Puerto Rico, Río Piedras, Puerto Rico 00931; and 2Department of Biology, University of Puerto Rico, Mayagüez, Puerto Rico 00681

Abstract.—We report 73 birds species in 26 families, including 31 new records, based on observations made during visits to Maricao State Forest from 1978 to 2000. The best-represented family was the Parulidae, with 19 species, including some rare migratory species in Puerto Rico, such as Swainson’s (Limnothlypis swainsonii) and Kentucky Warblers (Oporornis formosus). The species total in Maricao corresponded closely with the number of species in other montane forests in Puerto Rico. Maricao State Forest seems to be marginal for the endemic and endangered Puerto Rican Nightjar (Caprimulgus noctitherus) and Puerto Rican Broad-winged Hawk (Buteo platypterus brunnescens), but is prime habitat for the endangered Puerto Rican Sharp-shinned Hawk (Accipiter striatus venator) and the endemic Puerto Rican Vireo (Vireo latimeri). Populations of the resident Adelaide’s War- bler (Dendroica adelaidae) and (D. angelae) occur in sympatry only in Maricao State Forest. Resumen.—ESTADO, ABUNDANCIA Y DISTRIBUCIÓN DE LAS AVES DEL BOSQUE DE MARICAO, PUERTO RICO. Re- portamos 73 especies de aves, incluyendo 31 nuevos registros, basados en observaciones hechas durante visitas al Bosque Estatal de Maricao de 1978 a 2000. La familia mejor representada fue la Parulidae, con 19 especies, in- cluyendo algunas especies migratorias raras en Puerto Rico como la Reinita de Swainson (Limnothlypis swainsonii) y la Reinita de Kentucky (Oporornis formosus). El número total de especies en el Bosque Estatal de Maricao es similar al informado para otros bosques montanos en Puerto Rico. El Bosque Estatal de Maricao aparenta ser hábi- tat marginal para el Guabairo Pequeño de Puerto Rico (Caprimulgus noctitherus) y el Guaraguao de Bosque (Buteo platypterus brunnescens), ambas formas endémicas y en peligro de extinción, pero es hábitat primario para el Hal- cón de Sierra (Accipiter striatus venator), en peligro de extinción, y para el endémico Bien-te-veo (Vireo latimeri). Poblaciones residentes de la Reinita Mariposera (Dendroica adelaidae) y la Reinita del Bosque Enano (D. angelae) coexisten en simpatría únicamente en el Bosque Estatal de Maricao. Key words: abundance, biodiversity, populations, Puerto Rico, status

INTRODUCTION Grande, and Maricao, in the westernmost part of the LITTLE IS KNOWN ABOUT the avian species richness Cordillera Central. Elevations range from 150 to 875 of Maricao State Forest, the second largest forest m above sea level. Annual rainfall and temperature reserve in Puerto Rico. An inventory and species list average 2326 mm and 21.7°C, respectively (Silander of the avifauna is important for developing manage- et al. 1986). ment plans, implementing conservation measures, as Over 85% of the soils in Maricao State Forest are well as serving as baseline information for develop- derived from ultramafic rocks. These rocks represent ing research projects at the community level. Fur- 1% of the total area of Puerto Rico, limited in distri- thermore, a species list can be used to compare rich- bution to the southwestern part of the island ness among different forests in the island and with (Figueroa Colón 1992). Even with high levels of other regions. rainfall, the soils are dry because of their fast drain- In this report we present an updated account of the age and high permeability. Soils derived from ul- avifauna present in Maricao State Forest compiled tramafic rocks are characterized by high contents of from visits made during two decades. The present iron, chromium, nickel, and cobalt; low calcium/ study is the only comprehensive effort to update the magnesium ratio; and low contents of clay (Figueroa list of species in the Forest since Raffaele (1975). Colón 1992). High levels of endemism are pro- We also classify species according to their relative moted by these nutritional imbalances and high lev- abundance, and comment on their status and distribu- els of phytotoxics, such as nickel and magnesium tion. (Silander et al. 1986; Figueroa Colón 1992). Three life zones are found in Maricao Forest. The subtropical moist forest comprises 32.9% of the For- STUDY AREA AND METHODS est, the subtropical wet forest 65.2%, and the lower Maricao State Forest comprises 4150 ha in the mu- montane wet forest 1.9% (Ewel and Whitmore 1973; nicipalities of Mayagüez, San Germán, Sabana Colón et al. 1976). The vegetation can be classified

El Pitirre 14(2) Page 47 TOSSAS AND DELANNOY — BIRDS OF MARICAO STATE FOREST into five associations: dry slope forest, slope forest, Buchenavia capitata, Ocotea acuneata, and Magno- mixed hardwood, exposed ridge woodland, and Po- lia portorricensis. The palm, montana, and docarpus mixed hardwood (Colón et al. 1976). The the ferns, Cyathea arborea and C. aquiline, are moist life zone is found at elevations up to approxi- common in the understory. mately 600 m on the southern slopes of the Forest. The headwaters of various important rivers are The woody vegetation in the moist life zone is char- found in Maricao State Forest. Five rivers originate acterized by narrow crowns, slender stems, and an in the southern slopes of the Forest and three to the average canopy height of 6–9 m. The most common north, including the río Maricao (Colón et al. 1976). are Bursera simaruba, Homalium racemosum, The pools of a fish hatchery are filled with the water Plumeria obtusa, rosea, Eugenia confusa, of the río Maricao. Pimenta racemosa, and Dipholis cubensis. In areas Species list.—We compiled a bird species list from of high humidity with moderate slopes and deep soils observations made during approximately 1500 visits (e.g., along the narrow drainage basins), a more to Maricao State Forest from 1978 to 2000. Delan- mesic vegetation grows. This vegetation resembles noy repeatedly visited Maricao from 1978 to 1995, that which grows in the wetter life zones. whereas Tossas visited from 1998 to 2001. Most vis- The wet life zone receives 2000–4000 mm of rain- its were associated with ornithological research we fall per year and in Maricao State Forest occurs at have conducted over the years. The list includes all approximately 550–750 m elevation. The Forest in bird species we observed, as well as those reported the wet life zone has a poorly developed emergent by Raffaele (1975). We used field guides (Biaggi tree layer (Colón et al. 1976), and the canopy height 1970, Raffaele 1989, Raffaele et al. 1998) to identify ranges from 12 to 20 m. Dominant trees include: the birds observed with binoculars or trapped in mist Buchenavia capitata, Zanthoxylum martinicense, nets. Turpinia paniculata, Pithecellobium arboreum, and The frequency in which birds were encountered Coccoloba pubescens. Parts of this forest were previ- during our visits was used as a measure of their rela- ously cleared and reforested in the 1920s and 1930s tive abundance. Accordingly, species were catego- (Colón et al. 1976). rized as very common, if observed five or more times The lower montane wet life zone occurs from 700 in a single visit; common, if at least one individual to 900 m elevation and receives approximately 2550 was observed in more than half of the visits; uncom- mm of precipitation yearly. Two forest types domi- mon, if observed in less than half the visits; and rare, nate this zone. A dwarf spinescent vegetation domi- if two or fewer individuals were observed per year. nates the narrow ridges and peaks exposed to strong Accidentals or vagrants were those species that oc- winds and along the steep slopes with shallow ser- curred in Maricao State Forest only once every dec- pentine soils. The trees range from 2 to 5 m in height ade or greater. We also categorized birds according and 5 to 20 cm in diameter at breast height (dbh). to their status. Resident species, either permanent or The principal species are Dipholis cubensis, Guet- seasonal, are those known to breed in Maricao State- tarda pungens, G. scabra, Randia aculeata, Ilex Forest, non-breeding residents do not breed in the riedlaei, Coccoloba pirifolia, and Comocladia Forest, but do breed elsewhere on the island, and glabra (Colón et al. 1976). Where the slopes are not Neotropical migrants are birds that breed in the steep and the ridges are wide, more soil and humus Nearctic and spend several months in the Forest. A accumulate to produce a more mesic habitat. Here naturalized bird is an thriving in the are sclerophyllous but large, and the trees the wild. larger (5–10 m in height and 10–40 cm in dbh) than We include information on the distribution of each on the narrow ridges, peaks, and steep slopes with species in the Forest. The species list follows the shallow soils. The principal trees are Clusia griseba- taxonomic classification and nomenclature of the chiana, Coccoloba sintenisi, Eugenia confusa, and American Ornithologists’ Union (1998). Endemic Tabebuia schumanniana. A different forest type species appear in bold. grows on deep serpentine soils of the moderate slopes that characterize the wide upper ridges at ele- vations of 800–850 m (Colón et al. 1976). There the RESULTS forest type forms a more or less continuous canopy Ardeidae at 15–20 m. Epiphytic growth is not abundant, how- ever are common. Dominant tree species are: Snowy Egret (Egretta thula).—Common non- Micropholis chrysophyloides, Matayba dominguen- breeding resident observed feeding in creeks, fish sis, coriacea, Sloanea amygdalina, hatchery ponds, and margins of the río Maricao.

Page 48 El Pitirre 14(2) TOSSAS AND DELANNOY — BIRDS OF MARICAO STATE FOREST

Little Blue Heron (Egretta caerulea).—Common common resident observed in dense forest at all non-breeding resident observed feeding along the elevations. The species becomes uncommon in the margins of the río Maricao and ponds in the fish months following the breeding season. It forms hatchery. monospecific flocks in July and presumably mi- Cattle Egret (Bubulcus ibis).—Common non- grates to lowland forests during fall and winter. breeding resident observed feeding from the mar- This flocking behavior was reported by Rivera- gins of the fish hatchery ponds and the río Mari- Milán (1992) in wet and moist zones throughout cao. the island. Green Heron (Butorides virescens).—Common resi- White-crowned Pigeon (Columba leucocephala).— dent observed feeding along the margins of the Accidental non-breeding resident in the Forest. fish hatchery ponds and the río Maricao. Not observed by us, but reported by Raffaele (1975). Cathartidae White-winged Dove (Zenaida asiatica).— Turkey Vulture (Cathartes aura).—Very common Uncommon non-breeding resident observed in for- non-breeding resident observed flying over the est edges and disturbed . Three individuals Forest canopy from low to high elevations, mainly were seen by AGT, from February to April 1999, above the drier southern slopes. Turkey Vultures at 650 m elevation in the Forest interior in an area commute back and forth from the southwest to the heavily affected by Hurricane Georges (1998). The Añasco valley in western Puerto Rico, crossing species is common in disturbed habitats around over the Maricao State Forest. The vultures take Maricao State Forest. advantage of the strong slope and thermal updrafts in the Forest that create a corridor. They stop to Zenaida Dove (Zenaida aurita).—Uncommon resi- feed in Maricao when carrion is available. dent in winter, spring, and summer; found in dry slope forest, plantation forest, and Podocarpus– Accipitridae mixed hardwood forest at different elevations. Sharp-shinned Hawk (Accipiter striatus venator).— Common Ground-Dove (Columbina passerina).— Locally common resident in dense mesic forest Uncommon resident found at middle and low ele- and maría (Calophyllum brasiliense) plantations vations in the dry southern slopes. from middle to high elevations. Maricao holds one of five known breeding populations. Other popula- Key West Quail-Dove (Geotrygon chrysia).—A tions are in Toro Negro, Carite, Luquillo, and Gui- rare breeding resident at lower elevations in the larte Forests. Only 130 individuals remain of this dry southeastern slopes. Locally common near endangered endemic to Puerto Rico Cerro Avispa (450–500 m elevation) in the west- (Delannoy 1997). ern slopes. However, AGT captured two adults in mist nets at 750 m elevation on 2 April 1999, and Broad-winged Hawk (Buteo platypterus brunnes- found a pair building a at 770 m on 1 June cens).—Accidental in the Forest. Not observed by 2000. The nest was depredated. the authors, but reported by Raffaele (1975). It is an endangered subspecies with extant populations Ruddy Quail-Dove (Geotrygon montana).—Very in Luquillo, Carite, and Río Abajo Forests, totaling common resident in dense forests at all elevations 125 individuals (Delannoy 1997). before Hurricane Georges, but uncommon thereaf- ter. The species seems to be highly affected by Red-tailed Hawk (Buteo jamaicensis).—Common canopy loss from hurricane winds, as reported af- resident widespread throughout the Forest, ob- ter (1989) in the Luquillo Experi- served at all elevations and habitats. mental Forest, in eastern Puerto Rico (Wunderle Falconidae 1995). American Kestrel (Falco sparverius).—Uncommon Cuculidae resident found in disturbed habitats (e.g., vacation Mangrove Cuckoo (Coccyzus minor).—Rare resi- center) and forest edges at all elevations. dent, found mainly at middle and low elevations (Falco columbarius).—Accidental migrant near forest edges, in disturbed habitats, and dry in the Forest. One individual observed hunting in forests of the southern slopes. the edge of a mesic forest at 700 m elevation on 28 Puerto Rican Lizard-Cuckoo (Saurothera vieil- October 2000 by AGT. loti).—Common resident, more frequently heard Columbidae than seen. Widespread throughout the Forest. Scaly-naped Pigeon (Columba squamosa).—Very

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Strigidae comm.). Puerto Rican Screech-Owl (Otus nudipes).—A Tyrannidae common resident, widespread throughout the For- (Contopus latirostris).— est. Common breeding resident found at all elevations, Short-eared Owl (Asio flammeus portoricensis).— mainly in the understory of dense woodland. Rare resident, one individual was observed on 1 Puerto Rican Flycatcher (Myiarchus antil- August 1982 by CAD at 770 m in dense brush larum).—Rare resident found mainly in disturbed dominated by fern mats in the dry southeastern areas and forest edges at all elevations. Common slopes. in shade coffee plantations around the Maricao Caprimulgidae State Forest. Puerto Rican Nightjar (Caprimulgus noc- Gray Kingbird (Tyrannus dominicensis).—Common titherus).—Vagrant, one individual was observed breeding resident found in open and disturbed and photographed on 10 January 1988 by CAD in woodlands at all elevations. a maría plantation, 300 m north of the west fork of Loggerhead Kingbird (Tyrannus caudifasciatus).— the río Maricao, at 600 m above sea level. Possibly Rare resident found mainly in secondary forests, it was a stray bird from the nearby population in disturbed woodland, and edges at all elevations. Susúa State Forest. Vireonidae Apodidae Puerto Rican Vireo (Vireo latimeri).—Very com- Black Swift (Cypseloides niger).—Seasonal resi- mon breeding resident widespread throughout the dent, common in spring and summer at high eleva- Forest. tions. The breeding status in the Forest is un- Black-whiskered Vireo (Vireo altiloquus).—Very known. Observed in groups of four to six individu- common resident widespread throughout the For- als, feeding with swallows. est. Breeds in Maricao State Forest but presumably Trochilidae migrates to northern from Septem- Antillean Mango (Anthracothorax dominicus).— ber to February (Raffaele et al. 1998). Rare resident species, observed at low elevations Hirundinidae in the south- and east-facing slopes. Martin (Progne dominicensis).— Green Mango (Anthracothorax viridis).— Common non-breeding resident observed flying Common breeding resident widespread throughout over the Forest canopy during spring and summer. the Forest. Possibly migrates to South America in the winter Puerto Rican Emerald (Chlorostilbon mau- months (Raffaele et al. 1998). gaeus).—Common breeding resident widespread Cave Swallow (Petrochelidon fulva).—Common throughout the Forest. non-breeding resident observed in groups of five Todidae or more flying over the Forest canopy at all eleva- Puerto Rican ( mexicanus).—Very tions. common breeding resident found in forested habi- Turdidae tats at all elevations. Red-legged Thrush (Turdus plumbeus).—Very Alcedinidae common resident found at all elevations and habi- Belted (Ceryle alcyon).—Common tats in the Forest. Neotropical migrant, observed mainly in the pools Mimidae of the fish hatchery and along the río Maricao. Pearly-eyed Thrasher (Margarops fuscatus).— Picidae Uncommon resident found mainly in disturbed Puerto Rican Woodpecker (Melanerpes por- woods, forest edges, and forest stands along rivers toricensis).—Very common resident present at all from low to high elevations. elevations and habitats. Commonly seen and heard Sturnidae in groups of up to six individuals. Hill Myna (Gracula religiosa).—Accidental exotic Yellow-bellied Sapsucker (Sphyrapicus varius).— in the Forest. CAD observed one individual from Accidental Neotropical migrant in the Forest. An March to May 1993 near the Department of Natu- adult male was observed on 19 February 2001 at ral and Environmental Resources headquarters at 880 m elevation (José A. Colón López, pers. an elevation of 770 m.

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Parulidae Swainson’s Warbler (Limnothlypis swainsonii).— Northern Parula (Parula americana).—Common Accidental Neotropical migrant in the Forest. One Neotropical migrant widespread in the Forest from record: CAD trapped and banded an adult on 16 fall until early spring. January 1992 in dense forest at 770 m elevation. Magnolia Warbler (Dendroica magnolia).—Rare Ovenbird (Seiurus aurocapillus).—Common Neotropical migrant observed in dense forest. Neotropical migrant frequently observed near the CAD observed one adult male 10 April 1981 at ground in dense forest over a wide range of eleva- 650 m elevation. tions from fall to spring. Cape May Warbler (Dendroica tigrina).—Common Northern Waterthrush (Seiurus noveboracensis).— Neotropical migrant observed in a wide range of Neotropical migratory wood warbler not observed habitats and elevations from fall to spring. by the authors, but reported by Raffaele (1975). Black-throated Blue Warbler (Dendroica caerules- Louisiana Waterthrush (Seiurus motacilla).— cens).—Common Neotropical migrant widespread Common Neotropical migrant observed along in the Forest from fall until early spring. creeks and the río Maricao from fall until early spring. Yellow-rumped Warbler (Dendroica coronata).— Rare Neotropical migrant observed in dense forest Kentucky Warbler (Oporornis formosus).— and edges. CAD observed a pair of adults on 7, 13, Accidental Neotropical migrant in the Forest. One and 25 February 1981 at 800 m elevation. record: an adult male trapped and banded by CAD on 9 March 1992 in dense forest at 770 m eleva- Black-throated Green Warbler (Dendroica tion. virens).—Accidental Neotropical migrant observed in dense forest. Two adult males observed by CAD Hooded Warbler (Wilsonia citrina).—Accidental from 5 January until 7 April 1981 at 875 m eleva- Neotropical migrant observed in dense woodland. tion. AGT observed an adult male on 1 April 1999 One adult male trapped and banded by AGT on 22 at 700 m. March 1999 in dense forest at 700 m elevation. Yellow-throated Warbler (Dendroica ).— Coerebidae Accidental Neotropical migrant in the Forest. Ob- (Coereba flaveola).—Very common served by AGT on 11 and 26 November 2000 at resident found at all elevations and habitats. 650 m elevation, in pine (Pinus caribaea) and Thraupidae eucalyptus (Eucalyptus robusta) plantations, re- Puerto Rican (Nesospingus specu- spectively. liferus).—Very common resident widespread Adelaide’s Warbler (Dendroica adelaidae).— throughout the Forest. Common resident found in vegetation at all Scarlet Tanager (Piranga olivacea).—Accidental elevations. Neotropical migrant in the Forest. One record: an Prairie Warbler (Dendroica discolor).—Rare adult female trapped and banded by AGT in dense Neotropical migrant observed in dense forest from forest at 700 m elevation on 25 October 1998, fall to spring. probably passing through in its southward migra- Blackpoll Warbler (Dendroica striata).— tory route. Uncommon Neotropical migrant, observed mainly Puerto Rican (Spindalis portoricen- in dense forest at high elevations during its south- sis).—Very common resident widespread through- ward migration. out the Forest. Elfin Woods Warbler (Dendroica angelae).— Antillean Euphonia (Euphonia musica).—Common Common resident found mainly in the Podocar- resident observed at different elevations and habi- pus–mixed hardwood association from 700–900 tats. The species became uncommon in the Forest m, but also can be observed in dense forest at after Hurricane Georges (Tossas, pers. obs.). lower elevations. Emberizidae Black-and-white Warbler (Mniotilta varia).— Yellow-faced Grassquit (Tiaris olivacea).— Common Neotropical migrant widespread Uncommon resident observed mainly in openings throughout the Forest from fall until early spring. and forest edge over a wide range of elevations. American Redstart ( ruticilla).— Black-faced Grassquit (Tiaris bicolor).—Very com- Common Neotropical migrant widespread mon resident in openings, disturbed areas, and for- throughout the Forest from fall to early spring.

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est edge over a wide range of elevations. in vertical structure. Adelaide's Warbler prefers Puerto Rican Bullfinch (Loxigilla portoricen- shorter shrub vegetation, whereas Elfin Woods War- sis).—Very common resident widespread through- bler prefers taller and dense forest (Cruz and Delan- out the Forest. noy 1984). These congeners occur in sympatry over a wide elevational gradient in Maricao State Forest. Cardinalidae This is the only reported habitat in Puerto Rico Rose-breasted Grosbeak (Pheucticus ludovi- where the two species coexist (Cruz and Delannoy cianus).—Accidental Neotropical migrant in the 1984). Forest. CAD observed two adults, male and fe- Wood warblers are also the most represented group male, on 4 March 2001 at 750 m elevation. in other montane forests in Puerto Rico. Miranda- Icteridae Castro et al. (2000) reported 13 species, including Shiny Cowbird (Molothrus bonariensis).— the resident Adelaide’s Warbler in Los Tres Picachos Uncommon resident. Observed in disturbed habi- State Forest. Wiley and Bauer (1985) reported 18 tats, along edges, and forest fragments from low to species in Luquillo Forest, including the resident high elevations. Elfin Woods Warbler. Among the migratory wood Greater Antillean Oriole (Icterus dominicensis).— warblers found in Maricao, several are rare in Puerto Uncommon resident found in a variety of habitats Rico. These include the Kentucky, Swainson’s, Yel- at all elevations. We have seen Shiny Cowbird fe- low-throated, and Black-throated Green warblers. males entering oriole in disturbed habitats in The presence of these rare warbler species, along the Forest. with Scarlet Tanager, can be related to the vast ex- tension of this forest reserve, and high quality and Estrildidae diversity of habitats available. Moreover, shade cof- Indian Silverbill (Lonchura malabarica).— fee plantations around Maricao State Forest act as Naturalized species in Puerto Rico, rarely ob- buffer zones and further increase the amount of for- served in Maricao State Forest. CAD observed a est cover. flock of 14 on 11 January 1986 at 650 m in the dry Leopold (1963) and Raffaele (1975) reported the south-facing slopes. Broad-winged Hawk in Maricao State Forest. How- Tricolored Munia (Lonchura malacca).— ever, we have not seen any in this forest or adjacent Naturalized species in Puerto Rico, rarely ob- habitats in 20 years. We conclude that this bird is not served in Maricao State Forest. On 8 September a regular member of the Forest avifauna. Its status 1985, CAD observed a flock of 15 in a forest edge better fits the category of a vagrant species. Simi- at 650 m in the dry south-facing slopes. larly, Raffaele (1975) reported White-crowned Pi- geons in Maricao State Forest, and Wiley (1979) DISCUSSION sighted an unreported number in March 1975 while conducting monthly and bimonthly censuses in 1974 In this work we report 73 bird species in 26 families and 1975. However, we did not see or hear this spe- for Maricao State Forest, including 13 species and cies in Maricao during our study period. We suggest two subspecies endemic to Puerto Rico. This number the White-crowned Pigeon should be considered a increases the size of the previous list (Raffaele 1975) vagrant in Maricao. by 31 species. More species, however, might be added with further effort, particularly from censuses Maricao State Forest may be at the upper limit of or mist-netting when Neotropical migrants are pre- the endemic and endangered Puerto Rican Nightjar’s sent or passing through. Other published bird lists for range. Vilella and Zwank (1993) reported two night- montane forests in Puerto Rico closely correspond to jars in an eucalyptus plantation in the eastern bound- our species total. Miranda-Castro et al. (2000) re- ary of the Forest. Nightjar abundance tapered off ported 72 species in Los Tres Picachos State Forest, considerably to extremely low levels from a transi- and Wiley and Bauer (1985) reported 66 in Luquillo tional zone between dry coastal and dry limestone Forest. forest south of Susúa and Maricao State Forests (Vilella and Zwank 1993). The Puerto Rican Night- The largest taxonomic group in the Forest is repre- jar sighting we report is from a plantation near the sented by wood warblers (Parulidae), with 19 spe- northern boundary of the Forest. cies. Two species in this bird family are residents in Maricao State Forest, the Elfin Woods and Ade- The sightings of the naturalized exotic Indian laide’s warblers. Both are common in the Forest but Silverbill and Tricolored Munia in Maricao State segregate ecologically, selecting habitats that differ Forest seem to be the movement of flocks from low-

Page 52 El Pitirre 14(2) TOSSAS AND DELANNOY — BIRDS OF MARICAO STATE FOREST lands into the interior mountains during the non- ties. Caribb. J. Sci. 20:89–96. breeding period. Delannoy (pers. obs.) has seen DELANNOY, C. A. 1997. Status of the Broad-winged flocks of Tricolored Munias in farmlands close to the Hawk and Sharp-shinned Hawk in Puerto Rico. foothills of the Luquillo Forest at elevations of 700 Caribb. J. Sci. 33:21–33. m during the fall and winter months. Also, Wiley EWEL, J. J., AND J. L. WHITMORE. 1973. The eco- and Bauer (1985) reported the Bronze Mannikin logical life zones of Puerto Rico and the U. S. Vir- (Lonchura cucullata) from the Luquillo Forest as an gin Islands. US For. Serv. Res. Pap. ITF–18. uncommon resident in edges and interface of pas- tures, croplands, and fallow lands. Miranda-Castro et FIGUEROA COLÓN, J. C. 1992. La ecología de los al. (2000) reported naturalized exotic Orange- suelos ultramáficos. Acta Científica 6:49–58. cheeked Waxbill (Estrilda melpoda) and Bronze LEOPOLD, N. F. 1963. Checklist of birds of Puerto Mannikin to be common in Los Tres Picachos State Rico and the Virgin Islands. Univ. Puerto Rico Ag- Forest at elevations of 550 m. In addition, they re- ric. Exp. Sta. Bull. 168:1–119. ported the Pin-tailed Whydah (Vidua macroura) as LITTLE, E. L., JR., AND F. W. WADSWORTH. 1964. uncommon, but present, at an elevation of 690 m. Common trees of Puerto Rico and the Virgin Is- Maricao State Forest provides important habitat for lands. US For. Serv. Agric. Handbook No. 249, the endangered and endemic Puerto Rican Sharp- Washington, DC. shinned Hawk. This subspecies, whose population is MIRANDA-CASTRO, L., A. R. PUENTE-ROLÓN, AND estimated at 130 individuals, is one of the most en- S. VEGA-CASTILLO. 2000. First list of the verte- dangered on the island (Delannoy 1997). The Mari- brates of Los Tres Picachos State Forest, Puerto cao subpopulation, with 56 individuals, is the largest Rico, with data on relative abundance and altitud- in Puerto Rico. Other endemic birds common in the inal distribution. Caribb. J. Sci. 36:117–126. Forest, such as the Elfin Woods Warbler, Puerto Ri- can Tanager, and Puerto Rican Vireo, have disjunct RAFFAELE, H. 1975. Wildlife management in Mari- or relatively small populations elsewhere on the is- cao State Forest. Dept. Nat. Res., San Juan, Puerto land (Raffaele 1989). Therefore, the conservation of Rico. the habitat in Maricao State Forest, and the surround- RAFFAELE, H. A. 1989. A guide to the birds of ing buffer lands, will guarantee the subsistence of a Puerto Rico and the Virgin Islands. Princeton Univ. high species diversity, particularly of endangered, Press, Princeton, New Jersey. endemics, and rare birds. RAFFAELE, H., J. WILEY, O. GARRIDO, A. KEITH, AND J. RAFFAELE. 1998. A guide to the birds of the West Indies. Princeton Univ. Press, Princeton, New ACKNOWLEDGEMENTS Jersey. We are grateful for the comments and suggestions of R. A. Pérez-Rivera, J. A. Collazo, E. H. Williams, RIVERA-MILÁN, F. F. 1992. Distribution and relative Jr., J. Colón-López, J. M. Wunderle, Jr., and an abundance patterns of columbids in Puerto Rico. anonymous reviewer of an earlier version of this pa- Condor 94:224–238. per. SILANDER, S., H. GIL DE RUBIO, M. MIRANDA, AND M. VÁZQUEZ. 1986. Compendio enciclopédico de los recursos naturales de Puerto Rico. 10:210–236. LITERATURE CITED VILELLA, F. J., AND P. J. ZWANK. 1993. Geographic AMERICAN ORNITHOLOGISTS’ UNION. 1998. Check- th distribution and abundance of the Puerto Rican list of North American birds: 7 ed. American Orni- Nightjar. J. Field Ornithol. 64:223–238. thologists’ Union, Washington, DC. WILEY, J. W. 1979. The White-crowned Pigeon in BIAGGI, V. 1970. Las aves de Puerto Rico. Editorial Puerto Rico: Status, distribution, and movements. J. Univ. Puerto Rico, San Juan, Puerto Rico. Wildl. Manage. 43:402–413. COLÓN, E., D. J. POOL, F. RHOADES, AND R. DWOR- WILEY, J. W., AND G. P. BAUER. 1985. Caribbean SKY. 1976. The Master Plan for the Commonwealth National Forest, Puerto Rico. Am. Birds 39:12–18. forests of Puerto Rico. Dept. Nat. Res., San Juan, Puerto Rico. WUNDERLE, J. M., JR. 1995. Responses of bird popu- lations in a Puerto Rican forest to Hurricane Hugo: CRUZ, A., AND C. A. DELANNOY. 1984. Ecology of the first 18 months. Condor 97:879–896. the Elfin Woods Warbler (Dendroica angelae). I. Distribution, habitat usage, and population densi-

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