BIRDS OF MONTECRISTO NATIONAL PARK, EL · PDF fileBIRDS OF MONTECRISTO NATIONAL PARK, EL...

ORNITOLOGIA NEOTROPICAL 13: 167–193, 2002© The Neotropical Ornithological Society

BIRDS OF MONTECRISTO NATIONAL PARK, EL SALVADOR

Oliver Komar

University of Kansas, Department of Ecology and Evolutionary Biology, andNatural History Museum and Biodiversity Research Center, Lawrence, Kansas 66045 USA.

E-mail: [email protected]

Resumen. – Aves del Parque Nacional Montecristo, El Salvador. – A través del presente inventariode las aves de la estación reproductora en Parque Nacional Montecristo, se estimó las abundanciasrelativas de aves de bosque nebuloso, bosque pino-roble y plantaciones de ciprés (Cupressus lusitanica). Losmétodos utilizados incluyeron búsquedas intensivas de 4 horas diarias, conteos de punto con radio fijo, ycapturas con redes de neblina. Hay 233 especies de aves reportadas para el parque y la lista completapodría exceder las 275, dado muestreos incompletos y la distribución regional de la avifauna. El bosquepino-roble era el hábitat más diverso, aunque la densidad de aves era mayor en el bosque nebuloso. Lasplantaciones de ciprés eran intermedias en diversidad, pero forman el hábitat con las densidades menoresde aves y tienen menos especies residentes. Diez especies de aves en El Salvador están restringidas al Par-que Nacional Montecristo, y cinco de estas al bosque nebuloso. Los 52% de las aves del parque están ame-nazadas en El Salvador que incluye 22% que están en peligro al nivel nacional. De 18 aves restringidas alnorte de Centro-américa y reportadas en El Salvador, 15 habitan Montecristo, y tres no se encuentran enotro lugar del país. Comparado a un bosque nebuloso de mayor extensión en Chiapas, México (ElTriunfo), el bosque nebuloso de Montecristo tiene ~ 90% de las especies residentes, y tiene tres especiesno reportadas en El Triunfo. Se reportan cinco especies de aves nuevas para la lista nacional de las aves deEl Salvador.

Abstract. – An inventory of the breeding season birds of Montecristo National Park (El Salvador) esti-mated relative abundances of birds in cloud forest, pine-oak forest, and Mexican cypress (Cupressus lusita-nica) plantations. Methods included intensive searches (4 hours per day), fixed-radius point counts, andmist-netting. A list of 233 bird species was compiled for the park, and the actual species list may be higherthan 275, given incomplete sampling and regional bird distributions. Pine-oak forest was by far themost diverse habitat, although bird density was highest in cloud forest. Cypress plantations hadintermediate diversity, but lowest bird density, and fewer resident species than the other habitats.Many species at the park are habitat specialists restricted to cloud forest or pine-oak forest. Ten birdspecies in El Salvador are restricted to Montecristo National Park, and five of these to the cloud forest.Fifty-two percent (118 species) of the park's bird species are threatened in El Salvador, including 22% (53species) that are endangered at the national level. Of 18 regionally-endemic birds (restricted tonorthern Central America) reported from El Salvador, 15 are reported from Montecristo, and threeare not found anywhere else in El Salvador. Compared to the larger El Triunfo cloud forest in Chiapas,Mexico, Montecristo's cloud forest has about 90% as many resident species, and includes three speciesnot reported at El Triunfo. Five new species of birds are reported for the El Salvador national bird list.Accepted 7 November 2001.

Key words: Birds, communities, density, cloud forest, conservation, cypress plantation, inventory, pine-oak forest, El Salvador.

167

KOMAR

INTRODUCTION

Montecristo National Park protects the mostextensive cloud forest in El Salvador, as wellas montane pine-oak forest. The park is cur-rently the only Salvadoran protected area inthe ancient mountains of the Sierra Madre ofnorthern Central America. Montecristo andadjoining areas in El Salvador, Guatemala,and Honduras form the proposed La Frater-nidad Biosphere Reserve (Herrera et al. 1998).The park is home to numerous species offauna and flora not known to occur anywhereelse in El Salvador (Daugherty 1973, ReynaVásquez 1979, Thurber et al. 1987, Owen &Jones 1993, Schuster et al. 2000). A number ofspecies and subspecies were first described toscience from collections made at the park(Hincks 1953, Fryxell 1980, Hidalgo 1983, dela Maza E. & de la Maza E. 1984, Schuster1989, Zamudio 1997, Anderson & Ashe2000).

Cloud forest is a specialized habitat withmany endemic species – organisms uniquelyadapted to cloud forest that cannot live in anyother habitat. Many of the bird species withranges geographically restricted to northernCentral America are cloud forest endemics(Hernández-Baños et al. 1995, Long 1995).Detailed information on the distribution ofthese species is not available because siteinventories of birds are available for very fewNorthern Central American cloud forests.There are more than 50 separate cloud forestsin the biogeographic region of Northern Cen-tral America, which includes Chiapas (Mex-ico), Guatemala, Honduras, El Salvador, andNicaragua. Currently, only one site inventoryof cloud forest birds from this region iswidely available, for the El Triunfo cloud for-est in Chiapas (Parker et al. 1976, Gómez deSilva G. et al. 1999).

Several ornithological studies have beencarried out at Montecristo National Park,although none can be characterized as an

inventory, and none estimated relative abun-dances of different species. Steinbacher (1956,1958) collected 25 bird species at areas withinor adjacent to today's Montecristo park. PeterHamel (unpubl. report) observed 77 otherspecies in the park in 1974. Thurber (1978)documented 11 more species with photo-graphs. Hellebuyck (1983) collected two newspecies for El Salvador from the park. Pullen(1983) studied the Highland Guan (Penelopinanigra), but did not report additional species.Sixteen additional species were reported byThurber et al. (1987) for the park. In 1990,Néstor Herrera and Wilfredo Rodríguez(pers. com.) noted a single new species, bring-ing the bird list to 132. I visited Montecristobriefly in 1991, 1995, and 1996, recording 35species not previously reported from the parkand a few new records for El Salvador(Komar 2001). In 1996, Juan PabloDomínguez compiled an unpublished parkbird list, including 38 new species encoun-tered mostly in the tropical semi-deciduousforest zone (Domínguez, pers. com.). Visitorsto the park continued to record new speciesin unpublished field notes. In 1995 and 1997,Néstor Herrera found six; during 1997–1999Ricardo Ibarra recorded three; and in June1998, Wilfredo Rodríguez, Karla Pérez, Nés-tor Herrera, Roberto Rivera, and RicardoIbarra added four. In February–March 1999,Wilfredo Rodríguez, Ricardo Ibarra, and Rob-erto Rivera found nine more species for thepark's bird list.

Prior to the present study, 227 species ofbirds had been recorded in the park. Many ofthese species had not been reported from anyother site in El Salvador. The lack of a pub-lished list and relative abundance informationfor the park's birds, and a desire to under-stand the conservation importance of Monte-cristo National Park, motivated this iventory.As a result, the park bird list is now at 233species, and relative abundance estimates areavailable for birds in three major habitats

168

BIRDS OF MONTECRISTO NATIONAL PARK

within the park. This information permits ananalysis of the conservation importance ofthe park and its habitats.

STUDY AREA

Parque Nacional Montecristo is a 2000-haprotected area in the Department of SantaAna, northwest El Salvador (Fig. 1) that pro-tects an altitudinal transect from 800 m to2418 m above sea level. Cloud forest occu-pies all of the area between 2000 and 2350 m.Above 2350 m, the forest is replaced by ashrub (or elfin forest) community. Movingdownslope, the forest transitions to humidpine-oak forest in the zone between 1900 mand 1000 m. Below 1000 m, disturbed tropi-cal semi-deciduous forest and agriculturalplots dominate the landscape but these werenot surveyed. A 300-ha Mexican cypress

(Cupressus lusitanica) plantation occupies mostof the zone between 1750 and 2000 m. Sev-eral more plantations, mostly ocote pine(Pinus oocarpa) and Caribbean pine (Pinus ca-ribbea), occupy patches totalling less than 100ha. Some of these plantations were well-established by 1974, and others were estab-lished in the mid-1970s.

The cloud forest and pine-oak forests arecontiguous with similar forests outside parkboundaries. The size of the contiguous patchof cloud forest is roughly 2500 ha, of whichapprox. 825 ha is in El Salvador, with theremainder in adjoining Honduras and Guate-mala (Reyna Vásquez 1979). Only about 300ha of cloud forest are within park boundaries.I estimate a total patch size of 15,000 ha forpine-oak and pine forest, with about 1000 hawithin park boundaries. Two shaded coffeeplantations occupy 45 ha within the park,

FIG. 1. Montecristo National Park is located near the intersection of El Salvador, Honduras, andGuatemala.

169

KOMAR

between 900 and 1450 m, and a similar area isoccupied by other anthropogenic activities,such as agricultural plots and human habita-tions, mostly below 1200 m. The two villagesand scattered habitations within the park havea combined population of 704 (unpubl. data,Montecristo National Park, 1998). Tropicalsemi-deciduous forest occupies about 200 hain the lowest part of the park.

The cloud forest tree community includesat least 177 tree species and eight tree fernspecies (Reyna Vásquez 1979). Abundant can-opy trees (20–40 m tall) include Quercus aaata,Symplocos culminicola, Magnolia hondurensis, Cor-nus disciflora, and Brunellia mexicana (HeribertoLima, pers. com.). Abundant middle canopyspecies (5–20 m tall) include trees of the fam-ilies Araliaceae and Myrtaceae, as well as Hedy-osmum mexicanum and Saurauia sp., and the treefern Alsophila salvinii (Reyna Vásquez 1979).The shrub forest above 2350 m has a canopyof about 6 m, comprised of six species of thefamily Ericaceae (Reyna Vásquez 1979). Thedominant trees in the pine-oak forest arePinus oocarpa, Quercus eschineris, Quercus tristis,Diphysa robinioides, and Perymenium grande (Heri-berto Lima, pers. com.). There are patches ofmostly pine, and also mostly oak where bothof the Quercus spp. dominate. In the shrubbyunderbrush, a species of Agave with leaves upto 2 m long is common.

The base of operations for field work wasthe Cabaña Científica (1900 m, 14°23’N,89°22’W), a four-room log cabin at LosPlanes de Montecristo (“Los Planes”), a rec-reational area with picnic areas, campgrounds,and extensive flower gardens. Around LosPlanes, windbreaks and older plantations ofMexican cypress were probably planted wellbefore 1950.

METHODS

Relative abundances of birds were estimatedduring June and July 1999, using three sam-

pling methods: intensive searches with stan-dardized durations, fixed-radius point counts,and mist-netting. I also recorded birds casu-ally in the area of Los Planes de Montecristoon 15–17 June, 22–23 June, 27 June, 20–21July, and 27 July. I observed birds near thepark entrance, altitude 800 m, briefly whileentering or leaving the park. Documentaryphotographs are on-line at http://nhm.ku.edu/komar and have been deposited at theAcademy of Natural Sciences in Philadelphia,Pennsylvania (accession numbers v06/40/001–020).

Intensive searches. I conducted 4 h (06:00 to10:00 CST) surveys on seven days in cloudforest (during 16–26 June 1999), five days inhumid pine-oak forest (during 22–31 July1999), and four days in plantations ofMexican cypress (during 20–29 July 1999).Surveys consisted of walking slowlythrough the habitat, mostly along trails orroads but also off trails, and stopping fre-quently for periods of 10 min. Only in cloudforest did I visit some areas twice. All birdsheard and seen were noted, unless flying overand not evidently using the habitat. The dawnchorus began at 05:05; sunrise was at 05:30.Sampling began after the dawn chorus hadsubsided, to avoid bias in detection probabil-ity caused by increased singing during thechorus.

To avoid considering migrant or vagrantspecies as part of the community of breedingbirds, I defined members of the breedingcommunities as those that regularly breed orare presumed to breed in their respective hab-itats, and refer to such species as “core” spe-cies. Species detected casually and onlyoutside of intensive searches were conserva-tively considered non-core species, for com-parison purposes. All species detected by theintensive searches were assumed to be corespecies, since even species detected only oncewere also observed during casual observa-

170


tions or mist-netting, and did not appear tobe vagrants I conservatively considered addi-tional species reported by earlier studies(mostly opportunistic observations) to benon-core species, except when nesting wasreported.

Standardized point counts. I estimated densitiesof bird populations as an index of relativeabundance, using the fixed-radius point countmethod (Hutto et al. 1986). Points werelocated at 100 m intervals along transectschosen arbitrarily so as to be representativeof the major part of each habitat sampled,and visited during the intensive searchesdescribed above. Thus the data collected atpoints form part of the data for intensivesearches. I visited points once, and recordedall birds seen or heard within a 25 m radiusduring 10 minutes. The point count methodassumes that all birds detected were presentat a single moment in time, and thereforespecies recorded as a result of moving intothe count circle during the count inflatedensity estimates. This bias can be reducedby using shorter count durations, but lowdetectability in tropical forests due to infre-quent singing (author's unpubl. data) necessi-tate longer durations. Ten-minute durationshave been used widely in the tropics. Thesebiases cause density estimates (or other mea-sures of relative abundance) to be overesti-mated in some species and underestimated inothers.

For density estimates, I assumed that asingle individual recorded at a point repre-sented a breeding pair, and that additionalindividuals recorded were members of thesame pair or family unit, a conservativeassumption. Thus the units for density arepairs ha-1, and the estimated density of a givenspecies in a given habitat is the number ofcounts (points) where the species wasdetected in the given habitat, divided by thesum of the areas surveyed. The estimated

density for all birds in a given habitat is thesum of individual densities. The circle sur-veyed around each point (radius 25 m) mea-sures 0.196 ha. I visited 51 points in cloudforest (10.0 ha), 52 points in pine-oak forest(10.192 ha), and 39 points in cypress planta-tions (7.644 ha). The statistical significance ofdifferences in densities was calculated forspecies recorded at four or more points,using the presence-absence data for eachpoint, and 3 × 1 contingency tables(chi-square test). Apparent differences inestimated density should be interpretedwith caution, because they may reflect differ-ences in detectability and not density.Because different habitats were sampled atslightly different times of year, detectability ofbirds may have differed among habitats(Komar 1995).

Captures with mist nets. Field assistants and Icaptured under-story birds with mist nets incloud forest, pine-oak forest, and cypressplantation, using 9–12 nets per day, openedfor 6.5 h per day weather permitting, from05:30 until 12:00. The nets measured 12 × 2.5m, with 36 mm mesh. Nets were placed inlines along narrow roads and trails, withbreaks of 25 to 50 m between each pair ofnets. I combined mist-netting results fromFebruary and March, 1999 (W. Rodríguez,unpubl. data) with those from June and July1999, to present relative capture frequenciesfor all birds (very few migratory birds werecaptured during the earlier period whenNearctic migrants were present). In cloudforest, nets were operated for 716 net-hduring 18–19 February (100 net-h), 4 and 10–11 March (154 net-h), and 16–18 and 23–26June (462 net-h). In pine-oak forest, netswere operated for 597 net-h during 11–12February (82 net-h), 22 and 27 June (132 net-h), and 20, 22–23, and 27–30 July (383 net-h).Distribution of mist-netting effort in pine-oak forest was biased towards the upper ele-

171

KOMAR

TABLE 1. Mean number of birds detected per 4-h observation periods and captured per mist-nettinghour, by habitat.

Species Cloud forest Pine-oak forest Cypress plantations

Detected(n = 7)1

Captured(n = 716)2

Detected(n = 5)1

Captured(n = 597)2

Detected(n = 4)1

Captured(n = 216)2

Thicket Tinamou (Crypturellus cinnamo-meus)

Black Vulture (Coragyps atratus)Turkey Vulture (Cathartes aura)Zone-tailed Hawk (Buteo albonotatus)Red-tailed Hawk (Buteo jamaicensis)Barred Forest-Falcon (Micrastur ruficol-

lis)American Kestrel (Falco sparverius tropi-

calis)Highland Guan (Penelopina nigra)Singing Quail (Dactylortyx thoracicus)Red-billed Pigeon (Columba flavirostris)Band-tailed Pigeon (Columba fasciata)White-tipped Dove (Leptotila verreauxi)White-faced Quail-Dove (Geotrygon al-

bifacies)Squirrel Cuckoo (Piaya cayana)Fulvous Owl (Strix fulvescens)Chestnut-collared Swift (Streptoprocne

rutila)Lesser Swallow-tailed Swift (Panyptila

cayennensis)Violet Sabrewing (Campylopterus hemile-

ucurus)Green Violet-ear (Colibri thalassinus)White-eared Hummingbird (Hylocharis

leucotis)Azure-crowned Hummingbird (Ama-

zilia cyanocephala)Berylline Hummingbird (Amazilia be-

ryllina)Green-throated Mountain-gem (Lam-

pornis viridipallens)Amethyst-throated Hummingbird

(Lampornis amethystinus)Garnet-throated Hummingbird (Lam-

prolaima rhami)Magnificent Hummingbird (Eugenes ful-

gens)Sparkling-tailed Hummingbird (Tilma-

tura dupontii)

0.1

0.70.1

1.7

2.1

0.1

0.4

1.7

1.3

0.003

0.001

0.001

0.013

0.029

0.006

0.6

1.00.60.20.20.2

0.6

1.01.40.21.02.8

1.4

0.6

0.4

0.2

0.21.4

1.0

0.6

0.2

0.002

0.0050.012

0.018

0.002

0.007

0.010

0.30.8

1.0

0.8

0.3

0.3

0.3

0.0050.005

0.009

0.014

0.005

172


TABLE 1. Continued.


Detected(n = 7)1

Captured(n = 716)2

Detected(n = 5)1

Captured(n = 597)2

Detected(n = 4)1

Captured(n = 216)2

Violaceous Trogon (Trogon violaceus)Collared Trogon (Trogon collaris)Resplendent Quetzal (Pharomachrus mo-

cinno)Blue-throated Motmot (Aspatha gularis)Blue-crowned Motmot (Momotus momo-

ta)Emerald Toucanet (Aulacorhynchus

prasinus)Acorn Woodpecker (Melanerpes formici-

vorus)Golden-fronted Woodpecker (Melaner-

pes aurifrons)Golden-olive Woodpecker (Piculus ru-

biginosus)Northern Flicker (Colaptes auratus)Scaly-throated Foliage-gleaner (Anaba-

certhia variegaticeps)Ruddy Foliage-gleaner (Automolus rubi-

ginosus)Tawny-throated Leaftosser (Sclerurus

mexicanus)Ruddy Woodcreeper (Dendrocincla ho-

mochroa)Ivory-billed Woodcreeper (Xiphorhyn-

chus flavigaster)Spot-crowned Woodcreeper (Lepidoco-

laptes affinis)Barred Antshrike (Thamnophilus dolia-

tus)Mountain Elaenia (Elaenia frantzii)Yellow-olive Flycatcher (Tolmomyias sul-

phurescens)Greater Pewee (Contopus pertinax)Yellowish Flycatcher (Empidonax flaves-

cens)Dusky-capped Flycatcher (Myiarchus tu-

berculifer)Boat-billed Flycatcher (Megarynchus pi-

tangua)Brown-capped Vireo (Vireo leucophrys)Rufous-browed Peppershrike (Cycla-

rhis gujanensis)

1.11.4

2.4

2.1

0.3

0.4

0.9

0.7

2.7

5.6

9.6

2.1

0.001

0.004

0.007

0.42.4

1.02.2

1.8

2.0

0.4

3.6

2.4

3.8

2.0

0.2

0.40.4

3.22.2

3.4

0.4

0.42.4

0.002

0.002

0.012

0.008

1.8

1.0

1.5

0.5

0.5

2.5

9.5

1.0

0.005

0.005

173

KOMAR

TABLE 1. Continued.


Detected(n = 7)1

Captured(n = 716)2

Detected(n = 5)1

Captured(n = 597)2

Detected(n = 4)1

Captured(n = 216)2

Bushy-crested Jay (Cyanocorax melanocy-aneus)

Black-throated Jay (Cyanolyca pumilo)Unicolored Jay (Aphelocoma unicolor)Black-capped Swallow (Notiochelidon pi-

leata)Brown Creeper (Certhia americana)Band-backed Wren (Campylorhynchus

zonatus)Rufous-and-white Wren (Thryothorus

rufalbus)Plain Wren (Thryothorus modestus)House Wren (Troglodytes aedon)Rufous-browed Wren (Troglodytes rufoci-

liatus)Gray-breasted Wood-Wren (Henico-

rhina leucophrys)Eastern Bluebird (Sialia sialis)Brown-backed Solitaire (Myadestes occi-

dentalis)Slate-colored Solitaire (Myadestes unico-

lor)Orange-billed Nightingale-Thrush

(Catharus aurantiirostris)Ruddy-capped Nightingale-Thrush

(Catharus frantzii)Spotted Nightingale-Thrush (Catharus

dryas)Swainson's Thrush (Catharus ustulatus)*Black Robin (Turdus infuscatus)Mountain Robin (Turdus plebejus)Clay-colored Robin (Turdus grayi)Rufous-collared Robin (Turdus rufitor-

ques)Blue-and-white Mockingbird (Melanotis

hypoleucus)Olive Warbler (Peucedramus taeniatus)Crescent-chested Warbler (Parula super-

ciliosa)Black-throated Green Warbler (Dendro-

ica virens)*Grace's Warbler (Dendroica graciae)Ovenbird (Seiurus aurocapillus)*

0.40.30.1

7.7

10.3

17.9

9.7

7.1

8.30.7

0.7

0.003

0.013

0.015

0.053

0.025

0.003

11.8

6.4

0.61.0

1.8

6.21.8

1.010.0

3.0

0.4

0.2

1.4

0.4

1.43.2

3.6

0.002

0.003

0.02

0.008

0.007

0.002

0.0070.003

0.002

0.003

8.0

1.51.3

11.5

1.8

0.8

2.3

1.5

10.0

6.3

1.3

0.52.8

0.0050.009

0.005

0.005

0.046

0.0050.005

0.009

174


vations, where the forest begins a transitiontoward cloud forest. In cypress plantation,nets were operated for 216 net-h (4 February,50 net-h; 12 March, 40 net-h; 15 June, 60 net-h; and 21 July, 66 net-h). The selection of netsites in cypress plantation was biased toward

sites with extensive undergrowth, frequentlyat the edge of a plantation, or in narrow bor-ders between plantations. However, themajor part of these plantations had minimalundergrowth not conducive to mist-netting.Since the netting sites were not typical of the

TABLE 1. Continued.


Detected(n = 7)1

Captured(n = 716)2

Detected(n = 5)1

Captured(n = 597)2

Detected(n = 4)1

Captured(n = 216)2

MacGillivray's Warbler (Oporornis tol-miei)*

Wilson's Warbler (Wilsonia pusilla)*Painted Redstart (Myioborus pictus)Slate-throated Redstart (Myioborus mini-

atus)Rufous-capped Warbler (Basileuterus ru-

fifrons)Golden-browed Warbler (Basileuterus

belli)Common Bush-Tanager (Chlorospingus

ophthalmicus)Flame-colored Tanager (Piranga biden-

tata)Slaty Finch (Haplospiza rustica)Cinnamon-bellied Flower-piercer (Di-

glossa baritula)White-naped Brush-Finch (Atlapetes al-

binucha)Chestnut-capped Brush-Finch (Buarre-

mon brunneinucha)Rose-breasted Grosbeak (Pheucticus lu-

dovicianus)*Black-vented Oriole (Icterus wagleri)Yellow-backed Oriole (Icterus chrysater)Black-headed Siskin (Carduelis notata)Lesser Goldfinch (Carduelis psaltria)Hooded Grosbeak (Coccothraustes abeil-

lei)Subtotal identifiedUnidentifiedTotal

5.0

21.1

0.3

0.6

4.1

132.39.6

141.9

0.001

0.054

0.029

0.001

0.001

0.022

0.001

0.289

2.04.8

1.0

7.6

0.4

2.2

6.02.80.80.6

133.211.2144.4

0.003

0.008

0.002

0.028

0.007

0.018

0.0030.002

0.208

2.3

1.5

7.0

0.5

4.0

0.8

2.8

89.85.595.3

0.009

0.019

0.005

0.009

0.023

0.009

0.037

0.245

*Nearctic migrant, captured in February or March.1n = number of 4-h observation periods.2n = number of mist-netting hours.

175

KOMAR

overall habitat, results of mist-netting incypress plantations should not be consideredas .representative of the understory bird com-munity in that habitat.

RESULTS

The breeding season inventory recorded 115species of birds. Intensive searches providedrelative abundance information for more spe-cies (88) than other methods. Mist-nettingrevealed secretive species that may have beenmore common than observations indicatedbut captured only 38 species (Table 1), ofwhich one – Slaty Finch (Haplospiza rustica) –was never otherwise detected, and two others– Berylline Hummingbird (Amazilia beryllina)and Magnificent Hummingbird (Eugenes ful-gens) – were only detected casually. Twenty-four species were only detected during casualobservations. Six species were recorded forthe first time in the park, and the park birdlist, including species reported from a varietyof earlier sources (cited earlier), now reaches233 (Komar 2000).

Species diversity in three habitats, and completeness ofthe inventory. Pine-oak forest was the most spe-cies-rich habitat, followed by tropical semi-deciduous forest, then cloud forest, thencypress plantation (Table 2). Six of sevenpopular diversity indices (Magurran 1988)concurred that pine-oak forest was morediverse than cloud forest and cypress planta-tion. The tropical semi-deciduous forest, notincluded in this study, will not be treated fur-ther in this paper. Bird abundances for cloudforest, pine-oak forest, and cypress plantationare given in Table 1.

The species accumulation curves for theintensive search method and mist-net cap-tures nearly reached asymptotes for theinventory work in cloud forest, suggestingthat the inventory for that habitat is relativelycomplete (Fig. 2). The accumulation curvesfor pine-oak forest and cypress plantationwere still rising at he end of field work, how-ever, indicating that the inventories for thesehabitats are still far from complete (Fig. 2).

The fixed-radius point counts permittedestimates of density, as an index of relative

TABLE 2. Bird species diversity by habitat at Montecristo National Park.

Pine-oak forest Cloud forest Cypress plantation Tropical semi-deciduous forest

Total speciesSpecies richness with equal effort

(16 h observed)Core resident species

Abundant residents1 Common residents2 Uncommon residents3

Rare residents4 Regional endemic resident species

16664

7063529

N/A5

10

7334

4310111489

5735

3561712

N/A6

87N/A

N/AN/AN/AN/AN/A

2

1Abundant: average individuals detected per observation period (4 h) = ≥ 5.2Common: average ≥ 1 and < 5.3Uncommon: average ≥ 0.2 and < 1.4Rare: average < 0.2. 5No species in pine-oak forest or cypress plantation could be classified as rare because of fewer than sixsampling periods.

176


abundance, for species that were relativelycommon. I compared estimated densitiesamong habitatst for 26 species that weredetected within the fixed radius at four ormore points (Table 3). Seventeen species

showed statistically significant differences.Eleven species were more abundant in cloudforest; five species were more abundant inpine-oak forest. One species, Black Robin(Turdus infuscatus), was abundant in cloudforest and cypress plantations (no significantdifference between those two habitats), whilebeing virtually absent in pine-oak forest. Thedensity estimates may be biased by differen-ces in song behavior and detectability relatedto time of year (Komar 1995) and to habitat.Sampling of cloud forest was completed inJune, while sampling of the other two habi-tats was completed one month later. Junesampling would be expected to increase theoverall bird density estimates because ofmore frequent song activity.

Intensive searching was statistically lesspowerful than fixed-radius point counts,given smaller sample sizes, but detected farmore birds and more species. Some species,probably among the rarest encountered, wereonly observed outside of standardized sam-pling. I summarize such observations inTable 4.

The most frequently detected species(recorded more than five times per 4 hobservation) in each major habitat's birdcommunity (Table 1) are presented herein order of frequency of detection. Forcloud forest they were: Common Bush-Tana-ger (Chlorospingus ophthalmicus), Slate-coloredSolitaire (Myadestes unicolor), Gray-breastedWood-Wren (Henicorhina leucophrys), Ruddy-capped Nightingale-Thrush (Catharus frantzii),Yellowish Flycatcher (Empidonax flavescens),Black Robin, Rufous-browed Wren (Troglo-dytes rufociliatus), Spotted Nightingale-Thrush(Catharus dryas), Mountain Elaenia (Elaeniafrantzii), and Golden-browed Warbler(Basileuterus belli). For pine-oak forest theywere: Bushy-crested Jay (Cyanocorax melanocya-neus), Brown-backed Solitaire (Myadestes occi-dentalis), Flame-colored Tanager (Pirangabidentata), Black-capped Swallow (Notiochelidon

FIG. 2. Accumulation curves for the bird inven-tory at Montecristo National Park, El Salvador, in1999 show that (A) the intensive search methoddetected most species in cloud forest, but sam-pling was incomplete in cypress plantations and inpine-oak forest; (B) mist-net sampling of birds wasrelatively complete in cloud forest, and incompletein pine-oak forest.

177

KOMAR

pileata), Plain Wren (Thryothorus modestus), andYellow-backed Oriole (Icterus chrysater). Forcypress plantations, they were: Brown-backedSolitaire, Black Robin, Mountain Elaenia,Bushy-crested Jay, Flame-colored Tanager,and Rufous-collared Robin (Turdus rufitorques).

Species accounts. The following section includesnatural history observations, movementsderived from netting recaptures, and new spe-

cies reported for El Salvador. Abundanceinformation contained in tables and appendi-ces is not repeated here. A complete list of allspecies recorded at Montecristo NationalPark is given elsewhere (Komar 2000). Ireport movements (or lack thereof) forWhite-naped Brush-Finch (Atlapetes albi-nucha), Golden-browed Warbler, Ruddy-capped Nightingale-Thrush, Spotted Nightin-gale-Thrush, Plain Wren, and Brown-backed

TABLE 3. Estimates of densities (pairs ha-1) of birds at Montecristo National Park, El Salvador, as anindex of relative abundance. The number of points (PTS) with detections is assumed to be equivalent tothe number of pairs per total area sampled. Species detected at < 4 points are not presented. Significantlydifferent densities are indicated by *. N represents the number of points used to estimate densities in eachhabitat. English names are given in Table 1.

SpeciesCloud forest

(n = 51)Pine-oak forest

(n = 52)Cypress plantation

(n = 39)χ2 P

PTS Density ± SD PTS Density ± SD PTS Density ± SDHylocharis leucotis Lampornis viridipallens Lampornis amethystinusLamprolaima rhami Colaptes auratus Lepidocolaptes affinisElaenia frantziiEmpidonax flavescens Vireo leucophrys Cyanocorax melanocyaneus Notiochelidon pileataTroglodytes rufociliatus Henicorhina leucophrys Myadestes occidentalis Myadestes unicolor Catharus frantzii Catharus dryas Turdus infuscatus Peucedramus taeniatusParula superciliosa Myioborus miniatus Basileuterus belliChlorospingus ophthalmicusPiranga bidentata Buarremon brunneinuchaIcterus chrysater

164

414163

119

241498

3

837

4

0.10 ± 0.010.60 ± 0.030.40 ± 0.03

0.40 ± 0.031.40 ± 0.051.60 ± 0.050.30 ± 0.02

1.10 ± 0.040.90 ± 0.04

2.40 ± 0.051.40 ± 0.050.90 ± 0.040.80 ± 0.04

0.30 ± 0.02

0.80 ± 0.043.70 ± 0.05

0.40 ± 0.03

43

33

4287

5

438

235

0.39 ± 0.030.29 ± 0.02

0.29 ± 0.020.29 ± 0.02

0.39 ± 0.030.20 ± 0.020.78 ± 0.040.69 ± 0.03

0.49 ± 0.03

0.39 ± 0.030.29 ± 0.020.78 ± 0.04

0.20 ± 0.020.29 ± 0.020.49 ± 0.03

1

128

2

3

118122

134

0.13 ± 0.02

0.13 ± 0.020.26 ± 0.031.05 ± 0.05

0.26 ± 0.03

0.39 ± 0.04

0.13 ± 0.020.13 ± 0.021.05 ± 0.050.13 ± 0.020.26 ± 0.030.26 ± 0.03

0.13 ± 0.020.39 ± 0.040.52 ± 0.04

7.1242.9238.2467.3443.1430.31915.8520.752.2779.6112.7421.2817.154.90651.5324.1813.99.814.630.0269.6115.1385.212.3490.3198.97

0.028*0.2320.016*0.025*0.2080.8530.001*0.001*0.3200.008*0.002*0.001*0.001*0.0860.001*0.001*0.001*0.007*0.0990.9870.008*0.001*0.001*0.3090.8530.011*

178

179

BIRDS O

F MO

NTE

CRISTO N

ATION

AL PA

RKTABLE 4. Species observed during casual observations in 1999, but not during standardized sampling.

Hab NotesClouPine

Cyp

Trop

Non

ing overult flying over, 1500 m served by R. Ibarra Planes Planesserved by R. Ibarra

Planescks of 10 and 20 birds Planes Planes

coffee plantation (R. Ibarra)served by R. Ibarra and W. Rodríguezund 1450 m served by R. Ibarracoffee plantation (R. Ibarra)

ing overing overing overaging in cypress tree, 1900 mck of 7 at Los Planesk entrancek entrancek entrancek entrancek entrancek entrancek entrance Planes Planes

itats Species Dates d forest-oak forest

ress plantation

ical semi-deciduous forest

-forested areas

Black Vulture (Coragyps atratus)King Vulture (Sarcoramphus papa)Solitary Eagle (Harpyhaliaetus solitarius)Short-tailed Hawk (Buteo brachyurus)Collared Forest-Falcon (Micrastur semitorquatus)Buffy-crowned Wood-Partridge (Dendrortyx leucophrys)White-faced Quail-Dove (Geotrygon albifacies)Mottled Owl (Ciccaba virgata)Chestnut-collared Swift (Streptoprocne rutila)Vaux's Swift (Chaetura vauxi)Amethyst-throated Hummingbird (Lampornis amethystinus)Turquoise-browed Motmot (Eumomota superciliosa)Tufted Flycatcher (Mitrephanes phaeocercus)Social Flycatcher (Myiozetetes similis)Black-and-white Warbler (Mniotilta varia)Bar-winged Oriole (Icterus maculialatus)Yellow-billed Cacique (Amblycercus holosericeus)Zone-tailed Hawk (Buteo albonotatus)Chestnut-collared Swift (Streptoprocne rutila)Vaux's Swift (Chaetura vauxi)Azure-crowned Hummingbird (Amazilia cyanocephala)Blue-crowned Chlorophonia (Chlorophonia occipitalis)Great Kiskadee (Pitangus sulphuratus)White-throated Magpie-Jay (Calocitta formosa)Rufous-naped Wren (Campylorhynchus rufinucha)Scrub Euphonia (Euphonia affinis)Grayish Saltator (Saltator coerulescens)Black-headed Saltator (Saltator atriceps)Melodious Blackbird (Dives dives)White-winged Dove (Zenaida asiatica)Yellow-faced Grassquit (Tiaris olivacea)

26 June 23 July 29 July 24 June 24 June 29 July

23 June; 21, 23 July 15-16, 23-24 June

23 and 27 July 16 June 21 July 30 July 22 July 30 July 29 July 30 July 16 June 30 July 30 July 30 July

28-29 July 22 June 26 July 26 July 26 July 26 July 26 July 26 July 26 July

15-16, 27 June 15, 22, 27 June

FlyAdObLosLosOb

LosFloLosLosIn ObAroObIn

FlyFlyFlyForFloParParParParParParParLosLos

KOMAR

Solitaire. I present breeding information for26 species. Breeding was reported elsewherefor Collared Trogon (Trogon collaris) andResplendent Quetzal (Pharomachrus mocinno,Thurber et al. 1987), Slate-throated Redstart(Myioborus miniatus, Thurber 1978), and Black-vented Oriole (Icterus wagleri, Komar et al.2000). “Reproductive condition” refers todeveloped brood patches and/or swollen clo-acal protuberances. Taxonomy follows theAmerican Ornithologists' Union (1998). Newspecies reported for El Salvador, with respectto the list published by Komar (1998), areindicated with an asterisk (*).

King Vulture (Sarcoramphus papa). An adultflew across the park on 23 July, in the pine-oak zone between 1450–1550 m. Park guardsoccasionally see the “Rey Zope” but say itdoes not nest in the park.

Solitary Eagle (Harpyhaliaetus solitarius). W.Rodríguez and R. Ibarra (unpubl. data)observed a pair calling and taking turns land-ing on 12 February 1999 and again on 19March in pine-oak forest between 1200–1300m. They thought the birds may have beenbreeding, but could not confirm a nest. R.Ibarra observed one nearby on 29 July, flyinglow over pine-oak forest at 1490 m.

Highland Guan (Penelopina nigra). Park guardsregularly encounter this species from about1200 m (near the village of Las Majaditas)upwards into the cloud forest. The Penelopinavocalized frequently throughout June andJuly. A nest was found in March 1999 incloud forest, in a tree known locally asChemís (Juan José Aldana, pers. com.). On 17June a family group included the adults andone young male that appeared full-grown butwas brown with some black feathers. On 21July, I found a female sitting on a nest atabout 1910 m, in a patch of very old cypresstrees that were natural (E. Ramos, pers.

com.). The nest tree was 20 m from a small(25 m across) marshy pond. A brook that fedthe pond ran almost directly below the nest.The female did not leave the nest during ourvisit. The nest was in a cypress tree, on a hor-izontal branch, 3 m from its terminus and 4.5m from the trunk, 10 m above the ground.The tree itself was only about 15 m tall, but itsbase was right next to a much larger cypresstree about 35 m tall. The understory of theforest was dense with shrubbery less than 2 mhigh, and a middle stratus of trees 8–12 mtall. The nest was ~60 cm in diameter and~40 cm deep, made of fine cypress branches,pine needles, and a large green moss. E.Ramos had encountered three nests of thisguan since 1974, twice in forks of branches oflarge oaks in cloud forest, and once in a treefern in cloud forest. On 30 July, I saw afemale with a ¾ grown juvenile that hadblackish wings and a darker tail (male?), in anatural cypress grove at about 1800 m, east ofLos Planes.

Band-tailed Pigeon (Columba fasciata). A nestwas being built in a tall cypress tree adjacentto the Cabaña Científica (1900 m) on 21–22June. On 25 June, one or more pigeons weremaking long circular flights, presumably aspart of courtship, above the cloud forest(2300 m). I found broken egg shells, con-firmed of this species by E. Ramos, in cypressplantation on 21 July at 1880 m. A pigeon wassinging from the tree above.

Berylline Hummingbird (Amazilia beryllina). N.Herrera (pers. com.) found a nest of this spe-cies at Los Planes in April 1997.

Green-throated Mountain-gem (Lampornis viridi-pallens). The lowest encounter of this specieswas at 1450 m, on 30 July in open pine forest.

Amethyst-throated Hummingbird (Lampornisamethystinus). W. Rodríguez observed a male

180


carrying nest material in its beak on 16 June,in cloud forest.

Garnet-throated Hummingbird (Lamprolaimarhami). A female carried material to a nest incloud forest, and tightly guarded the sitealong the edge of the road, harassing me as Iwalked by, on 17 June. One week later (24June), the nest was a deep cup of mosses andfine fibers, measuring 9 cm from top to bot-tom. It hung from roots under an overhang-ing bank in the road cut.

Magnificent Hummingbird (Eugenes fulgens). Amale was seen at Santos Martínez's gardenthroughout June and July, possibly the sameindividual that occasionally flew through theflower gardens below the Cabaña Científica.Despite frequent sightings, the species maybe rare (captured just once, not recorded dur-ing censuses).

*Broad-tailed Hummingbird (Selasphorus platy-cercus). Although not recorded during thebreeding season inventory, two females werereported by R. Ibarra (unpubl. data) on 19March 1999, at a small shade coffee planta-tion near 1300 m. The species was also foundon 4 June 1998, by Ibarra and N. Herrera(unpubl. data), at Hotel La Palma, La Palma,Chalatenango.

Blue-throated Motmot (Aspatha gularis). Birdswith brood patches were captured in cypressplantation on 15 June, and in pine-oak foreston 22 July. The typical call of this species,heard frequently in those habitats and incloud forest, is a long series of ringing hoots.On 17 June, one constantly repeated a squir-rel-like scolding call. On 28 July, two birdswere duetting in pine-oak forest, at about1650 m.

Acorn Woodpecker (Melanerpes formicivorus).Mist-nets captured a juvenile on 30 July near

the fire tower at 1400 m. In the same area on31 July, R. Ibarra observed a nestling peekingout of a hole 10 m high in a pine tree.

Northern Flicker (Colaptes auratus). Hamel(unpubl. report) observed flickers nesting in awindbreak of mature cypress trees at LosPlanes de Montecristo during the summer of1974. Birds were present in 1999 but nestingwas not detected.

*Tawny-throated Leaftosser (Sclerurus mexica-nus). I detected this bird in June, by sight andvocalizations, six times in cloud forest, 2050–2350 m. W. Rodríguez also saw one at theedge between cypress plantation and brushysecondary scrub, about 100 m from cloudforest. The species was first detected at Mon-tecristo (and in El Salvador) on 11 June 1998,when one flew into a mist net in the cloudforest (Herrera et al. 1998), and R. Rivera(pers. com.) photographed one at night roost-ing on the road bank.

Spot-crowned Woodcreeper (Lepidocolaptes affi-nis). On 23 June, an adult with recedingbrood patch and a juvenile with a not-yetfully-grown bill (presumably a mother withher young) were captured in the same net, incloud forest (photo v06/40/004).

Mountain Elaenia (Elaenia frantzii). Femaleswith brood patches were caught in the upperpine-oak forest (1930 m) on 20 July.

Yellowish Flycatcher (Empidonax flavescens).An adult with brood patch was captured inpine-oak forest (1900 m) on 22 June.

Bushy-crested Jay (Cyanocorax melanocyane-us). At Los Planes, a family group had fiverecently-fledged juveniles on 15 June. Oneadult had three white head feathers. The larg-est group observed was 22 birds in cypresstrees below the Cabaña Científica on 29 July.

181

KOMAR

Black-capped Swallow (Notiochelidon pileata).Nest holes were present in the road cutthroughout the upper parts of the pine-oakzone of the park. Swallows flew into nestholes on 18 June at 1750 m, and on 28 July at1825 m. A pair was nesting in the road cut incloud forest, around 2250 m. We captured thepair on 16 June; one had a brood patch.Thurber et al. (1987) reported nesting in thepark in May and June, 1973. Swallows werecommon around 1500 m during July. On 22July, an aggregation of 20 perched in an oldpine tree near km 16. During the non-breed-ing season, swallows probably wander tolower elevations. I noted six below 900 m on20 February 1991, inside the park.

Band-backed Wren (Campylorhynchus zonatus).On 15 June, a family group at Los Planesincluded four birds in adult plumage and onefledged juvenile. On 20 July, a group in thesame place included four juveniles. W.Rodríguez and R. Ibarra saw an adult carrymosses to a nest on 22 July, at the edge of theorchid garden parking lot. On 30 July, I notedthe domed nest was built over a fork on thelowest branch of a Mexican cypress, 4.5 mabove the ground, 1.5 m in from the tip of thehorizontal branch, and 6 m out from thetrunk. The nest shape was somewhat flat-tened and oval (longer than wide), with theentrance in the middle of the narrow end,measuring approx. 20 cm × 15 cm × 10 cm(l × w × h).

Plain Wren (Thryothorus modestus). Onebanded in 1996 by J. P. Domínguez (yellow legband number 282) was recaptured (near theCabaña Científica) on 20 July 1999.

Rufous-browed Wren (Troglodytes rufociliatus).This species was much less frequentlydetected at Montecristo than at the Santa AnaVolcano cloud forest (unpubl. data). In Mon-tecristo, the species lives among epiphytes (I

observed one singing from 18 m aboveground). The forest floor niche, where thespecies was abundant on the Santa Ana vol-cano, appeared to be occupied exclusively atMontecristo by the Gray-breasted Wood-Wren, a species absent from the volcano.

Gray-breasted Wood-Wren (Henicorhina leu-cophrys). On 17 June, in cloud forest, Iobserved a nest with young, in tangledbranches 4 m above the road, but just 1.5 mfrom the hillside. The same day, an adult withbrood patch was captured. Thurber et al.(1987) reported finding nests in the cloud for-est during May and June, 1974 and 1975, anddetermined that one pair's territory measuredapproximately 32 × 20 m, and was boundedby three other territories. If that territory sizewas typical, it suggests a density of 15.6 pairsha–1, much higher than the estimate of 0.9 ±0.04 pairs ha–1 generated by point-count data(Table 3). If the wrens vocalized infrequentlyduring my survey, then detection probabilitywould have been low, and the resulting den-sity estimate could be far below the true den-sity.

*Brown Creeper (Certhia americana). One wassinging on 30 July in open pine woods at 1450m on the top of a ridge about 300 m north ofthe fire tower. I photographed one (Photov06/40/007) and heard two at the same placeon 31 July. Three park guards with 25 years'experience in the park were not previouslyfamiliar with the species. The habitat was anopen grove of pine trees about 25 m tall and> 40 years old. Although reported for El Sal-vador by Thurber et al. (1987), the location ofthat 1973 record now pertains to Honduras.

Eastern Bluebird (Sialia sialis). In 1974, P.Hamel (unpubl. report) observed adults withfledged young near 1400 m, in open pinewoods. W. Rodríguez (unpubl. data) found apair nesting in almost the same place in 1998.

182


Birds were present in 1999, although nestingwas not detected.

Brown-backed Solitaire (Myadestes occidentalis).This species was an abundant nester in theearthen road cut throughout the pine-oak andcypress forest zone. The lower distributionlimit, according to park guards, is around 900m. On 23 June, three nests had three eggsand another had two nestlings. On 21 July, anest constructed of only dry pine needlescontained one egg. Another that day,constructed of mosses and dried earth, con-tained two nestlings. On 29 July, a nest 1.3 mabove the road in the bank had three eggs,and another nest with two eggs was found inthe wall of a house, about 2 m above theground. Both nests contained mostly pineneedles. In pine-oak forest, we captured afemale with brood patch on 22 July and alocal juvenile on 23 July 1999. Two capturedat Los Planes de Montecristo on 3 March1999 were recaptured there in June 1999. Iobserved skylarking on one occasion (23 June1999).

Slate-colored Solitaire (Myadestes unicolor).Adults in reproductive condition were cap-tured on 23–25 June in cloud forest and on20 and 27 July in the uppermost pine-oak for-est (1930 m), in the zone of transition tocloud forest. Thurber (1978) reported a nestat Hacienda Montecristo on 4 May 1973. Alocal juvenile was captured on 25 June incloud forest. On one occasion I observed anadult skylarking to about 8 m above the can-opy (16 June). The bird flew straight up anddown, singing on the way up.

Ruddy-capped Nightingale-Thrush (Catharusfrantzii). Adults in reproductive conditionwere captured on 15–25 June (cypress planta-tion and cloud forest) and 21 July (cypressplantation). A recently-fledged juvenile wascaptured in cloud forest on 18 June. A pre-

sumed floater captured near El Trifinio on 16June was captured about 1000 m away on 18June. Another banded in 1996 (yellow plasticleg band number 130) by J. P. Domínguez wasrecaptured in nearly the same place on 23June 1999.

Spotted Nightingale-Thrush (Catharus dryas).This species was nearly absent above 2100 m,abundant in the lower cloud forest (below2100 m), and uncommon in upper pine-oakforest down to 1750 m. One captured on 11March was recaptured on 24 June, havingmoved a few hundred meters. A presumedfloater captured on 17 June was recapturedon 23 June more than 500 m away. Adults inreproductive condition were captured incloud forest on 16–26 June. A local juvenilewas captured in pine-oak forest (1930 m) on27 July. Thurber et al. (1987) reported an adultwith brood patch collected 15 August1975. In El Salvador, this species has onlybeen documented from Montecristo, but R.Ibarra (pers. com.) saw one – possibly avagrant – at Volcán de San Vicente on 18April 1998.

Black Robin (Turdus infuscatus). On 17 June, afemale in the cloud forest was carryingfood. A female with brood patch was cap-tured on 27 July in pine-oak forest. The low-est observation of this species was onesinging from a cypress plantation at about1660 m on 23 July.

Clay-colored Robin (Turdus grayi). A nest wasfound at Los Planes, at 1900 m on 18 June, ina windbreak of mature cypress. A fledgedjuvenile was observed on 22 July. Hamel(unpubl. report) reported a nest in the samelocation in 1974.

Rufous-collared Robin (Turdus rufitorques).Juveniles with spotted plumage wereobserved near Los Planes on 15 and 22 June,

183

KOMAR

and on 20 July. On 27 June, a female visited acypress near the Cabaña Científica with foodseveral times, indicating a nest. Hamel(unpubl. report) found a nest in the sameplace in 1974.

Blue-and-white Mockingbird (Melanotis hypoleu-cus). On 15 June, a family group at Los Planeshad two juveniles, identified by dark grayplumage with dull whitish streaks on the darkunderparts and a white malar streak. Ialso noted breeding at Los Planes on 12August 1995, when I found a fledgling aboutone day out of the nest.

Crescent-chested Warbler (Parula superciliosa).On 23 July, a flock in undergrowth of pine-oak woodland (1750 m) included at least 15 ofthese warblers, with many immatures beggingfood from females, and small numbers ofSpot-crowned Woodcreeper, Mountain Elae-nia, Greater Pewee (Contopus pertinax), Dusky-capped Flycatcher (Myiarchus tuberculifer),Brown-capped Vireo (Vireo leucophrys),Grace's Warbler (Dendroica graciae), and Slate-throated Redstart. The lowest Crescent-chested Warbler recorded was observed by R.Ibarra at an oak grove around 1550 m on 28July.

*Tropical Parula (Parula pitiayumi). Althoughnot observed during the inventory, W.Rodríguez (pers. com.) observed three inMontecristo's cloud forest on 11 June 1998. Itwas first reported for El Salvador on 5 June1998, when N. Herrera (unpubl. data)observed two at 2045 m in thickets along theroad to Cerro El Pital, department of Chala-tenango. R. Ibarra (unpubl. data) foundanother at La Montañona, Chalatenango, 11November 1999.

Golden-browed Warbler (Basileuterus belli).Adults in reproductive condition and localjuveniles were captured in cloud forest on 16–

24 June. Two adults banded in March 1999were recaptured in June 1999. One was inalmost the same place, the other had movedseveral hundred meters.

Common Bush-Tanager (Chlorospingus oph-thalmicus). In cloud forest on 16 June, anadult was feeding a fledged juvenile andanother juvenile was captured in mist nets.Adults in reproductive condition werecaptured in cloud forest on 16–26 June,in upper pine-oak forest on 22 June, 20and 27 July, and in cypress plantation on 15June and 21 July. Local juveniles were cap-tured on 22 June and 27 July in upper pine-oak forest.

*Scarlet Tanager (Piranga olivacea). This Nearc-tic migrant was observed twice by R. Rivera(W. Rodríguez, pers. com.) between 12 and 15April 1999, one near the park headquartersand the other near the village of Las Majadi-tas. It has not been reported previously in ElSalvador.

White-winged Tanager (Piranga leucoptera).Although not recorded during this study, N.Herrera (pers. com.) recorded a nest at LosPlanes in April 1997.

Blue-crowned Chlorophonia (Chlorophonia occi-pitalis). Usually considered a cloud forest spe-cies, the chlorophonia may be more commonin the upper pine-oak forest around LosPlanes than in the cloud forest. I did not findit in cloud forest, but Thurber et al. (1987)stated that it was “rarely seen in the cloud for-est at Montecristo.”

Slaty Finch (Haplospiza rustica). This specieswas detected just once, a male not in breedingcondition, captured in a mist net (photo v06/40/005) at about 2120 m in cloud forest on23 June. Although two were collected atMontecristo in 1975 (in July and November),

184


neither was reported in breeding condition(Thurber et al. 1987).

White-naped Brush-Finch (Atlapetes albinucha).Adults in reproductive condition and a juve-nile were captured in pine-oak forest on 20–22 July. A bird marked with a colored legband on 4 February at Los Planes wasobserved there again on 15 June, although itwas not recaptured.

Chestnut-capped Brush-Finch (Buarremon brun-neinucha). Adults in reproductive conditionwere captured on 15–25 June (cloud forest,pine-oak forest, and cypress plantation).Local juveniles were captured in pine-oak for-est on 27 July. Thurber (1978) reported a nestat Hacienda Montecristo on 3 May 1973,containing a day-old chick.

Bar-winged Oriole (Icterus maculialatus). R.Ibarra observed an adult male on 30 July, atthe edge of pine-oak forest and shaded coffeeplantation at 1450 m. Two were observed on10 June 1998 at the same locality (W.Rodríguez, unpubl. data), and N. Herreraobserved one on 26 April 1995 in the park(Sector La Joya; pers. com.). These are thefirst records of this regional endemic speciesin El Salvador's northern cordillera. The spe-cies is not known from Honduras.

Black-headed Siskin (Carduelis notata). On 22June, at Los Planes, a female was feeding arecently-fledged juvenile.

Hooded Grosbeak (Coccothraustes abeillei).This species was observed mostly in cypressplantations around Los Planes de Monte-cristo. It was also found lower down theslope, including at a cypress plantationaround 1700 m (22 July) and in pine forestnear the fire tower (1400 m, 31 July). Breed-ing behavior was first noted on 23 June, whena male and female were observed together at

the edge of a cypress plantation at 2000 m.The female was collecting small twigs on adirt road that separated the cypress treesfrom the lower edge of cloud forest. Themale sat and watched from a nearby bush.On 24 June, near the Cabaña Científica, a pairwas accompanied by a begging juvenile,which was fed by the male. On 27 July, nestbuilding was observed at the Cabaña Cientí-fica, along the edge of an old (>50 years)cypress plantation. I suspect the pair was pre-paring to renest. On 29 July, the female was inthe final stages of building the nest in a 35 mtall cypress. The nest was a cup of unknownconstitution, about the size of a Clay-coloredRobin nest [outside dimensions for that spe-cies' nest are given as 15.2 × 20.3 cm diame-ter and 10.2 cm height by Skutch (1960)]. Itwas nestled on top of the dense and fineouter branches of the lowest major branch ofthe cypress, 7 m above the ground. The densefoliage around the nest made it impossible todescribe the nest more clearly. The nest wasabout 5 m out from the trunk and 1 m fromthe outermost limit of foliage. When thefemale visited the nest, the male perchedabout 5 m away. The male accompanied thefemale constantly, but did not directly partici-pate in nest building. The female took the ini-tiative in all 12 flights that I observed. Thiswas true for four visits to the nest (the femalewould arrive first, followed closely by themale), for the five times I observed the pairleave the nest, and for three times I observedthe pair fly among trees while away from thenest. Most grosbeaks detected during surveyswere vocalizing; however, males and femalescollecting nesting material and nest-buildingwere silent.

DISCUSSION

Due to lack of published site lists of birds inEl Salvador and Honduras, a recent analysisof montane bird distribution in Mexico and

185

KOMAR

Central America (Hernández-Baños et al.1995) used regional treatments (Monroe1968, Thurber et al. 1987) to assemble a list of173 montane forest species resident in inter-ior Honduras and El Salvador, and predictedthe presence of 10 additional bird species ongeographic grounds. The present study con-firms the presence of three species predictedby Hernández-Baños et al. (1995) to occur inthe region: Blue-and-white Mockingbird (Me-lanotis hypoleucus), Rose-throated Becard(Pachyramphus aglaiae), and Rufous-and-whiteWren (Thryothorus rufalbus). The becard, how-ever, was not found above 1000 m and maynot be a “montane” species in this region.

Almost half (46%) of the 233 bird speciesrecorded for Montecristo National Park aremontane forest resident species. Also presentare species of open habitats and lowlandareas, and 49 (21%) migratory visitors thatbreed in the Nearctic region. Numerous addi-tional open area species, lowland species, andmigratory species may occur. Future fieldwork may determine that the overall speciesrichness for the park is perhaps 275–290 spe-cies, given incomplete sampling in the parkand known regional bird distributions. Themost diverse forest area now recognized in ElSalvador, El Imposible National Park, is morethan twice as large as Montecristo, containsdifferent habitats, and has approx. 277 speciesof birds (Ramírez Sosa & Komar 1996; W.Rodríguez, unpubl. data).

Threatened and endemic species. Komar (1998)classified 508 Salvadoran bird species for con-servation status, finding that 254 species werethreatened at the national level, including 117in danger of extirpation in the short tomedium term (due to very reduced numbersor geographic restriction to one or two smallor medium-sized populations). Of thosethreatened species, 119 occur in MontecristoNational Park, including 50 endangered spe-cies. Two new species discovered recently at

the park and not included in the earlier analy-sis, Brown Creeper and Tawny-throated Leaf-tosser, meet the criteria for nationallyendangered. The Hairy Woodpecker (Picoidesvillosus) and Band-backed Wren, classified asthreatened, proved to be rare and probablyare endangered. The woodpecker was notrecorded, and perhaps is no longer present inthe park. Thus, 121 (52%) of the park's birdsare threatened and 54 (23%) are endangered.

The two forest corvids, Black-throated Jay(Cyanolyca pumilo) and Unicolored Jay (Aphelo-coma unicolor), are probably among the rarestand most threatened birds in El Salvador,being restricted to cloud forest (in El Salva-dor) and seen infrequently. Another endan-gered corvid is the Common Raven (Corvuscorax), which has not been recorded in thepark since 1995 and may now be extirpated inEl Salvador. Some of the other “endangeredspecies,” such as Amethyst-throated Hum-mingbird, Gray-breasted Wood-Wren, Slate-colored Solitaire, Spotted Nightingale-Thrush, and Golden-browed Warbler, areabundant in the cloud forest. These specieswere classified “endangered” because theirhabitat is endangered in El Salvador.Although these species have just one or tworelatively small populations in El Salvador,they may not be threatened on a regionalscale.

Ten species of birds in El Salvador aremostly restricted to Montecristo NationalPark, and are threatened due to reduced habi-tat (Table 5). Four of these species were oncefound in and near the cloud forests of CerroEl Pital (“Los Esesmiles” of Dickey & vanRossem 1938) in El Salvador's Chalatenangodepartment, but were not detected there dur-ing two weeks of field work in 2001 (author'sunpubl. data). Sixteen other species areknown in El Salvador only from highland for-ests at Montecristo and Cerro El Pital. Eigh-teen species of birds that are endemic to thebiogeographic regions of Northern Central

186


America occur in El Salvador (Komar 1998).All but three are found in MontecristoNational Park, and three are apparentlyrestricted in El Salvador to the park (Table 6).

Three habitats. Bird communities of the threeforest habitats surveyed differed profoundly.The present study confirms the conservationimportance of the Montecristo cloud forest(Daugherty 1973). Nine regionally-endemicbird species are found there, and five birdspecies in El Salvador are restricted to it.Eight other bird species restricted to cloudforest are also found at Cerro El Pital, 24 kmto the east, but lack adequate protectionthere. At least 11 bird species had signifi-cantly higher densities in the cloud forestthan in any other habitat in the park. Theimportance of the cloud forest is apparent,despite relatively low species richness.

The study has also demonstrated the con-servation importance of the park's pine-oakforest; its avifauna is largely different fromthat of the cloud forest. Species richness inpine-oak forest was higher than the other

habitats surveyed, and the species accumula-tion curve was still far from reaching itsasymptote (many more species can beexpected). Ten regionally-endemic bird spe-cies occupy this habitat, and two bird speciesare known in El Salvador only from this hab-itat in the park (Table 5). Moreover, threeother species restricted (in El Salvador) toMontecristo are residents of pine-oak forestsin addition to cloud forest. At least nine birdspecies were significantly more abundant inpine-oak forest than in cloud forest orcypress plantation.

The artificial tree plantations within thepark, such as the extensive Mexican cypressplantations, probably have little conservationvalue. A number of cloud forest and pine-oakforest bird species, including regionally-endemic and threatened bird species, werefound in the cypress plantations, althoughbreeding was noted for very few. A few spe-cies were detected more frequently in theplantations, e.g., Mountain Elaenia, BlackRobin, Rufous-collared Robin, and HoodedGrosbeak. Nonetheless, higher densities are

TABLE 5. Ten species of birds restricted in El Salvador to Montecristo National Park.

Species Habitats (forest types)Fulvous Owl (Strix fulvescens)1

Blue-throated Motmot (Aspatha gularis)1

Scaly-throated Foliage-gleaner (Anabacerthia variegaticeps)Ruddy Foliage-gleaner (Automolus rubiginosus)1

Tawny-throated Leaftosser (Sclerurus mexicanus)Belted Flycatcher (Xenotriccus callizonus)2

Unicolored Jay (Aphelocoma unicolor)1

Brown Creeper (Certhia americana)Spotted Nightingale-Thrush (Catharus dryas)3

Hooded Grosbeak (Coccothraustes abeillei)

Cloud Cloud, pine-oak

Cloud Cloud Cloud

Pine-oak Cloud

Pine-oak Cloud, pine-oak Cloud, pine-oak

1Reported from cloud forest on Cerro El Pital, department of Chalatenango, prior to 1943 (Dickey & vanRossem 1938, Marshall 1943) but not recorded there during 2001 (author's unpubl. data).

2Found in 1998 in the department of Chalatenango (R. Ibarra, unpubl. data) but status or future there isnot certain.

3A single Spotted Nightingale-Thrush was reported from the San Vicente Volcano in 1998 (R. Ibarra,unpubl. data); its status there is unknown.

187

188

KO

MA

RTABLE 6. Fifteen of El Salvador's 18 regionally-endemic bird species live in Montecristo National Park. These species are endemic to the Northern CentralAmerica biogeographic region (Nicaraguan Depression to the Isthmus of Tehuantepec, Mexico).

Spec MontecristoWhi

HighOce

PaciFulvRufoGre

Slen

Win

BlueBeltBush

BlacBlacRufo

RufoBlue

Bar-

d from lower elevations in the park (J.ínguez, pers. com.)n above 1200 morded, but likely to occur in mid-ele-onesordedmon in cloud forestordedn above 1450 m

ot recorded this study)

ot recorded this study)

n above 1600 mot recorded this study)n in pine-oak forest; absent or rare inrest

mon or rare in cloud forestn above 1400 m

mon, only in cloud forest

n in open areasn in young second growth and dis-reas

mon or rare in coffee plantation at

ies Distribution in El Salvador Status inte-bellied Chachalaca (Ortalis leucogastra)

land Guan (Penelopina nigra)llated Quail (Cyrtonyx ocellatus)

fic Parakeet (Aratinga strenua)ous Owl (Strix fulvescens)us Sabrewing (Campylopterus rufus)

en-throated Mountain-gem (Lampornis viridipallens)

der Sheartail (Doricha enicura)

e-throated Hummingbird (Atthis ellioti)

-throated Motmot (Aspatha gularis)ed Flycatcher (Xenotriccus callizonus)y-crested Jay (Cyanocorax melanocyaneus)

k-throated Jay (Cyanolyca pumilo)k-capped Swallow (Notiochelidon pileata)us-browed Wren (Troglodytes rufociliatus)

us-collared Robin (Turdus rufitorques)-and-white Mockingbird (Melanotis hypoleucus)

winged Oriole (Icterus maculialatus)

Lowlands

Montecristo and Cerro El PitalNorthern cordillera

Volcanic chain and coastMontecristo only (formerly Cerro El Pital)Volcanic chain Santa Ana Volcano, Montecristo, and Cerro ElPitalSanta Ana Volcano, Montecristo, and (formerly?)Cerro El PitalSanta Ana Volcano, Montecristo, and Cerro ElPitalMontecristo only (formerly Cerro El Pital)Montecristo onlyHighlands above 900 m (also reported at sea levelin Parque Nacional Deininger)Montecristo and Cerro El PitalMontecristo and Cerro El Pital Santa Ana Volcano, Montecristo, and Cerro ElPitalHighlands above 1700 m Highlands above 1600 m, occasional down to1200 m Middle elevations, mostly on volcanic chain

ReporteP. DomCommoNot recvation zNot recUncomNot recCommo

Rare? (n

Rare? (n

CommoRare? (nCommocloud foUncomCommoUncom

CommoCommoturbed aUncom1400 m


not necessarily evidence of habitat quality(van Horne 1983). The cypress plantationsseemed to be a transition zone between cloudforest and pine-oak forest, with bird speciesrepresentative of both the natural forest habi-tats.

Species not observed and possibly locally extirpated.Only 115 bird species were detected duringthe study, out of 233 species reported in thepark. Fifty-six species on the park's list areNearctic migrants or vagrants, thus I detected65% of 177 resident or potentially residentspecies. Some of the species not detected mayhave disappeared from the park in recentyears. Of concern are Hairy Woodpecker,Lineated Woodpecker (Dryocopus lineatus), andCommon Raven. Park guards recall seeing thelatter two species regularly up until 1995 andnot afterward (Silvestre Carranza, pers. com.).Two species reported breeding in the cloudforest by Thurber et al. (1987), Scaled Ant-pitta (Grallaria guatimalensis) and Eye-ringedFlatbill (Rhynchocyclus brevirostris), were notdetected, although both species maintainpopulations elsewhere in El Salvador (pers.observ.). Thurber et al. (1987) observed“White-breasted” Hawk (Accipiter striatus chio-nogaster), the distinct resident race that isendemic to northern Central America, fre-quently during 1974 at Los Planes, and Iobserved one on 13 August 1995; none wasrecorded in 1999.

Three species previously fairly commonaround Los Planes but apparently absent dur-ing the present study were White-throatedFlycatcher (Empidonax albigularis) and Buff-breasted Flycatcher (Empidonax fulvifrons)(Thurber et al. 1987), and Rufous-collaredSparrow (Zonotrichia capensis) (pers. observ.).The latter species was present in 1999 atHacienda Montecristo, just outside the park(W. Rodríguez, pers. com.). All three speciesprefer open habitat, and could eventuallyrecolonize clearings at Los Planes from habi-

tat outside the park.Belted Flycatcher (Xenotriccus callizonus) is

a regional endemic species reported from justa few sites in Chiapas (Mexico), Guatemala,and from Montecristo (Hellebuyck 1983). InJune 1998, R. Ibarra (unpubl. data) observedone about 30 km east of the park, near CerroEl Pital, department of Chalatenango. It isperhaps one of the rarest birds in El Salvador.My inability to find this species may havebeen due to lack of sufficient time spent in itsoak-forest habitat, or perhaps I visited whenbirds were not vocalizing and were most dif-ficult to detect. I was unable to discover if theMontecristo population persists.

I hoped to find the El Salvador Crested(Steller's) Jay (Cyanocitta stelleri lazula), a racerestricted to highlands of northwestern ElSalvador, described from the pine-oak andcloud forests of Cerro El Pital in Chala-tenango by van Rossem (Dickey & van Ros-sem 1938). The persistence of the species inEl Salvador is in question.

Comparison with El Triunfo cloud forest. Becausecloud forests tend to be geographically iso-lated, they are home to many endemic andthreatened species of birds (Long 1995). It isinteresting to consider how widely distributedare bird species ecologically restricted tocloud forests. Unfortunately, bird lists forcloud forests are difficult to find in the litera-ture. Birds of the El Triunfo cloud forest,located about 360 km to the west-northwestin Chiapas, Mexico, in a similar altitudinalrange as the cloud forest at Montecristo, wereinventoried by Parker et al. (1976) andGómez de Silva et al. (1999). I have found noother published cloud forest bird inventoriesfor the biogeographic region of northernCentral America (between the Isthmus ofTehuantepec and the Nicaraguan Depres-sion).

The El Triunfo cloud forest studied hasan overall extension that is more than twice

189

KOMAR

that of the patch where Montecristo islocated. More area was sampled at El Triunfo(about 1200 ha vs. about 300 ha), samplingwas carried out at all times of year, and totalobservational sampling effort at El Triunfowas greater (302 h vs 28 h). The Montecristoinventory included 716 net-hours of mist-net-ting understory birds; no mist-netting wasreported for El Triunfo. Unequal samplingefforts make comparisons difficult, butbecause sampling was so extensive at El Tri-unfo, I assume that it was virtually complete.Given that the Montecristo species accumula-tion curve had leveled off (for cloud forestonly), and that a few rare breeding species hadbeen previously reported, I also assume thatsampling at Montecristo was virtually com-plete. Complete inventories are comparable,even with unequal sampling.

Gómez de Silva et al. (1999) reported 80core species for El Triunfo, including migra-tory visitors (non-breeders). Twelve non-breeding visitors and two species recordedonce or twice should be removed to make thelist comparable to the 43 core speciesreported for Montecristo. The 66 residentspecies reported for El Triunfo include 12species present in the pine-oak forests ofMontecristo but not recorded in its cloudforest or too rare to be considered corespecies. These are Black Vulture (Coragypsatratus), Red-tailed Hawk (Buteo jamaicensis),Squirrel Cuckoo (Piaya cayana), Chestnut-collared Swift (Streptoprocne rutila), GreenViolet-ear (Colibri thalassinus), White-earedHummingbird (Hylocharis leucotis), MagnificentHummingbird, Tufted Flycatcher (Mitre-phanes phaeocercus), Rose-throated Becard,Brown-backed Solitaire, Slate-throated Red-start, and White-naped Brush-Finch. Thesespecies are not cloud forest specialists, andperhaps some should not be considered corespecies.

The remaining list of 54 core species at ElTriunfo includes four species (Wine-throated

Hummingbird Atthis ellioti, Hairy Wood-pecker, Elegant Euphonia Euphonia elegantis-sima, and Blue-crowned Chlorophonia) thathave been reported at Montecristo, but werenot recorded in the present cloud forestinventory; they may be rare residents at Mon-tecristo, or recently extirpated species. Thereare 10 species at El Triunfo that are evidentlyabsent at Montecristo (Table 7). The Stub-tailed Spadebill (Platyrinchus cancrominus) andWhite-throated Robin (Turdus assimilis) maynot actually be core species as they were notrecorded during the breeding season (Parkeret al. 1976, Gómez de Silva et al. 1999). Threespecies at Montecristo were absent at ElTriunfo (Table 7). One of these, Brown-capped Vireo, probably occurs at El Triunfo(Gómez de Silva et al. 1999). The other two,Mountain Elaenia and Slate-colored Solitaire,are among the most abundant species atMontecristo. The greater diversity (speciesrichness) at El Triunfo may be due to thelarger area of cloud forest available, or moreextensive adjacent forest.

Conclusions. Although the inventory of thebirds of Montecristo National Park is still notcomplete, I have provided the first relativeabundance and reproductive status informa-tion for birds in the park and several of itsprincipal habitats. This information permitsan interpretation of the importance of thepark for the conservation of El Salvador'sbiodiversity. The present contributionincreased the inventory of the park's birds to233 species, and includes five new species forEl Salvador's national bird list. More fieldwork at Montecristo National Park is neededto determine the conservation value of thepark's habitats, and the conservation needs ofits species. At present, very little is knownabout movements of threatened bird popula-tions in and around the park. A number ofthe cloud forest species, such as the Resplen-dent Quetzal but also many others, may move

190


out of the cloud forest to other habitats in oroutside the park, as part of annual migrations.For example, at some cloud forests in Mex-ico, 25% of the species withdraw to lower ele-vations during the coldest months (Binford1989). Year-round survey work may berequired to adequately describe these move-ments. Furthermore, at present we have noinformation on population trends in the park.A long-term monitoring program is neededto determine which species may be declining.The present study may serve as a baseline forsuch monitoring.

ACKNOWLEDGMENTS

I am most grateful to the technicians whoassisted with mist-netting, WilfredoRodríguez, Ricardo Ibarra, Roberto Rivera,and Ricardo Pérez. I thank the El SalvadorServicio de Parques Nacionales y Vida Silves-tre for permission to conduct this study inMontecristo Park, and for the collaborationof park personnel, especially Jaime Latín,park director, and park guards AlejandroAguilar, Lazaro Aguilar, Juan José Aldana, Sil-vestre Carranza, José Luís Ladino, Heri-bertoLima, Gabino Posada Molina, Eduardo Orel-lana, and Eugenio Ramos. At Los Planes deMontecristo, Santos and Olivia Martínez

assisted with hospitality and equipment stor-age. I thank Juan Pablo Domínguez, NéstorHerrera, Ricardo Ibarra, and WilfredoRodríguez for sharing their unpublished data.Funding for the study was provided by agrant from the National Fish and WildlifeFoundation, a National Science FoundationGraduate Research Fellowship, and a Madi-son A. and Lila Self Graduate Research Fel-lowship from the University of Kansas. Ithank SalvaNATURA and the Ministerio deMedio Ambiente y Recursos Naturales forassistance with transportation. Miguel OrtegaHuerta assisted with preparation of the map.Finally, I am grateful to A. Townsend Peter-son and two anonymous reviewers for valu-able comments on the manuscript.

REFERENCES

Anderson, R. S., & J. S. Ashe. 2000. Leaf litterinhabiting beetles as surrogates for establishingpriorities for conservation of selected tropicalmontane cloud forests in Honduras, CentralAmerica (Coleoptera; Staphylinidae, Curcu-lionidae). Biodivers. Conserv. 9: 617–653.

American Ornithologists' Union. 1998. Check-listof North American birds. 7th ed. AmericanOrnithologists' Union, Washington, D.C.

Binford, L. C. 1989. A distributional survey of thebirds of the Mexican state of Oaxaca. Orni-

TABLE 7. Species found at El Triunfo cloud forest, in Chiapas, Mexico, not recorded for Montecristo, ElSalvador, and vice versa.

Species only at Montecristo Species only at El TriunfoMountain Elaenia (Elaenia frantzii)Brown-capped Vireo (Vireo leucophrys)Slate-colored Solitaire (Myadestes unicolor)

Horned Guan (Oreophasis derbianus)Barred Parakeet (Bolborhynchus lineola) Emerald-chinned Hummingbird (Abeillia abeillei)Mountain Trogon (Trogon mexicanus)Spotted Woodcreeper (Xiphorhynchus erythropygius)Paltry Tyrannulet (Zimmerius vilissimus)Stub-tailed Spadebill (Platyrinchus cancrominus)White-throated Robin (Turdus assimilis)Gray Silky-flycatcher (Ptilogonys cinereus)Chestnut-sided Shrike-Vireo (Vireolanius melitophrys)

191

KOMAR

thol. Monogr. 43: 1–405.Daugherty, H. E. 1973. The Montecristo cloud for-

est of El Salvador – a chance for protection.Biol. Conserv. 5: 227–230.

de la Maza E., J., & R. de la Maza E. 1984. NuevosDismorphiinae de México y El Salvador(Pieridae). Rev. Soc. Mex. Lepidopterol. A. C. 9:3–12.

Dickey, D. R., & A. J. van Rossem. 1938. The birdsof El Salvador. Field Mus. Nat. Hist. Zool. Ser.23: 1–609.

Fryxell, P. A. 1980. A new species of Hampea (Mal-vaceae) from El Salvador. System. Bot. 5: 442–444.

Gómez de Silva G., H., F. González-García, & M.P. Casillas-Trejo. 1999. Birds of the upper cloudforest of El Triunfo, Chiapas, Mexico. Ornitol.Neotrop. 10: 1–26.

Hellebuyck, V. 1983. Three new specimen recordsof birds for El Salvador. Wilson Bull. 95: 662–664.

Hernández-Baños, B. E., A. T. Peterson, A. G.Navarro-Sigüenza, & B. P. Escalante-Pliego.1995. Bird faunas of the humid montane for-ests of Mesoamerica: biogeographic patternsand priorities for conservation. Bird Conserv.Int. 5: 251–277.

Herrera, N., R. Rivera, & R. Ibarra. 1998. Estudiode la fauna vertebrada en la reserva de la bios-fera La Fraternidad (El Salvador, Guatemala yHonduras). Informe de Consultoría. Fun-dación para la Conservación de los Ecosiste-mas Mayas, Ciudad de Guatemala.

Hidalgo, H. 1983. Two new species of Abronia(Sauria: Anguidae) from the cloud forests of ElSalvador. Occ. Pap. Mus. Nat. Hist. Univ. Kans.105: 1–11.

Hincks, W. D. 1953. The Passalidae (Ins. Col.) of ElSalvador. Senckenb. 34: 29–35.

Howell, S. N. G., & S. Webb. 1995. A guide to thebirds of Mexico and northern Central America.Oxford Univ. Press, New York.

Hutto, R. L., S. M. Pletschet, & P. Hendricks. 1986.A fixed-radius point count method for non-breeding and breeding season use. Auk 103:593–602.

Komar, O. 1995. Variation in bird song detectionwithin the rainy season in El Salvador. WorkingPaper No. 4: 63–68, Wildlife Conservation

Society, Bronx, New York.Komar, O. 1998. Avian diversity in El Salvador.

Wilson Bull. 110: 511–533.Komar, O. 2000. Lista de las aves del Parque

Nacional Montecristo, El Salvador. Sal-vaNATURA, San Salvador.

Komar, O. 2001. Contribuciones a la avifauna deEl Salvador. Cotinga 16: 40–45.

Komar, O., W. Rodríguez, & R. Ibarra. 2000. Black-vented Oriole nests inside a cabin in El Salva-dor. Wilson Bull. 112: 551–553.

Long, A. J. 1995. The importance of tropical mon-tane cloud forests for endemic and threatenedbirds. Pp. 79–105 in Juvik, J. O., & F. N. Scatena(eds.). Tropical montane cloud forests.Springer-Verlag, New York.

Magurran, A. E. 1988. Ecological diversity and itsmeasurement. Princeton Univ. Press, Prince-ton, New Jersey.

Marshall, J. T., Jr. 1943. Additional informationconcerning the birds of El Salvador. Condor45: 21–33.

Monroe, B. L., Jr. 1968. A distributional survey ofthe birds of Honduras. Ornithol. Monogr. 7: 1–458.

Owen, J. G., & J. K. Jones. 1993. The red brocket,Mazama americana (Artiodactyla: Cervidae), inEl Salvador. Texas J. Sci. 45: 106.

Parker, T. A., III, S. Hilty, & M. Robbins. 1976.Birds of El Triunfo cloud forest, Mexico, withnotes on the Horned Guan and other species.Am. Birds 30: 779–782.

Pullen, T., Jr. 1983. Sound production and repro-ductive biology of the Highland Guan in ElSalvador's Montecristo cloud forest. Am. Birds37: 948–950.

Ramírez Sosa, C. R., & O. Komar. 1996. Plan parala conservación de la biodiversidad del ParqueNacional El Imposible. USAID Proyecto Pro-tección del Medio Ambiente, Colección Con-sultoría. San Salvador, El Salvador.

Reyna Vásquez, M. L. 1979. Vegetación arbórea delbosque nebuloso de Montecristo. Tesis delicenciatura. Univ. El Salvador, San Salvador.

Schuster, J. C. 1989. Petrejoides salvadorae sp. nov.(Coleoptera: Passalidae) from El Salvador. Fla.Ent. 72: 693–696.

Schuster, J. C., E. B. Cano, & C. Cardona. 2000. Unmétodo sencillo para priorizar la conservación

192


de los bosques nubosos de Guatemala, usandoPassalidae (Coleoptera) como organismosindicadores. Acta Zool. Mex. (n. s.) 80: 197–209.

Skutch, A. F. 1960. Life histories of Central Amer-ican birds II. Pacific Coast Avifauna 34: 1–593.

Steinbacher, J. 1956. Uber eine kleine Vogel sam-mlung aus El Salvador. Senckenb. Biol. 37:371–375.

Steinbacher, J. 1958. Weitere Beitrage zur Vogel-fauna von El Salvador. Senckenb. Biol. 39: 11–40.

Thurber, W. A. 1978. Cien aves de El Salvador.Ministerio de Educación, San Salvador.

Thurber, W. A., J. F. Serrano, A. Sermeño, & M.Benítez. 1987. Status of uncommon and previ-ously unreported birds of El Salvador. Proc.West. Found. Vertebr. Zool. 3: 109–293.

van Horne, B. 1983. Density as a misleading indi-cator of habitat quality. J. Wildl. Manage. 47:893–901.

Zamudio, S. 1997. Una especie nueva de Pinguicula(Lentibulariaceae) de Centroamérica. Acta Bot.Mex. 40: 65–69.

193

BIRDS OF MONTECRISTO NATIONAL PARK, EL · PDF fileBIRDS OF MONTECRISTO NATIONAL PARK, EL...

Documents

Transcript of BIRDS OF MONTECRISTO NATIONAL PARK, EL · PDF fileBIRDS OF MONTECRISTO NATIONAL PARK, EL...